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Page last updated
12 October 2017

Mussel of the Month

The 2017 Mussels of the Month, so far...

September 2017

LanceolariaLanceolaria gladiola (Unionidae, Indotropica)

The September 2017 Mussel of the Month is Lanceolaria gladiola. Lanceolaria is a genus of 10 species found in eastern Asia, from Vietnam north to Japan.

Back in November 2012, Lanceolaria grayana was the Mussel of the Month, and we used it as an opportunity to discuss the subfamily classification of the freshwater mussel family Unionidae (at least as we understood it then). Among those subfamilies is the Unioninae, which includes mussels like Unio, Lanceolaria, and Anodonta. As we pointed out back then, the subfamily is distinctive because it is easily recognized by morphological characteristics. The combination of hooked-type glochidia (parasitic larvae) and ectobranchy (brooding in the outer demibranchs) is diagnostic of the Unioninae. The other subfamilies haven't really beeen characterized except as clades in molecular analyses.

Back in 2006, we were unable to classify most of the Old World freshwater mussel genera to the rank of subfamily because we didn’t have molecular data for them (Graf & Cummings, 2006). Bouchet & Rocroi (2010) proposed a novel subfamily classification based on input from the community but only scant additional phylogenetic data. For the most part, it turns out their classification has been largely accurate in the sense that it has been reflected in recent phylogenetic studies — with a few modifications here and there (Whelan et al., 2011; Pfeiffer & Graf, 2015; Lopes-Lima et al., 2017). However, it is still the case, based on the recent genus-rich sampling of Lopes-Lima et al. (2017), that if we don’t have the molecules, the mussels can’t be classified. Any genus not included in their analysis was regarded as incertae sedis — even the North American Pegias.*

Interestingly, Lopes-Lima et al. (2017) proposed a revision to the classification of the Unioninae. Since Y2K, the subfamily has been divided among two tribes, Unionini and Anodontini. The anodontines** are easily recognized by the modifications of their marsupium to accommodate lateral expansion (i.e., gravid gills look like inflated mattresses) and the presence of tripartite water tubes. The unionine genera lack these characteristics.

Lopes-Lima et al. (2017) discovered that “unionines” like Lanceolaria and Arconaia share a more recent common ancestor with Anodonta than Unio. That is, the Unionini as we have traditionally viewed it, is paraphyletic. These authors proposed splitting the Unioninae into two subfamilies: Unioninae and Anodontinae (including the new tribe, Lanceolariini). That is a reasonable proposal that names every clade that needs a name. However, the problem with this scheme is that it separates the named taxa from their non-molecular synapomorphies (i.e., diagnostic derived characters). The unionid clade diagnosed by hooked-type glochidia would no longer have a name — mussels with that character could belong to either the Unioninae or the Anodontinae. Even the distinctive characteristics of the anodontines no longer refer to a named taxon. The subfamily Anodontinae includes genera like Lanceolaria that lack tripartite water tubes and marsupia capable of expansion, and the mussels that do have those characters are split between two tribes, Anodontini and Cristariini.

DLG re-analyzed the Lopes-Lima et al. (2017) data set (with additional genes) as a test case for his updated code for Conflict Clade Analysis. He presented that work, "Conflict Clade Analysis (CCA) of the freshwater mussel subfamily Unioninae (Bivalvia: Unionoida)," last summer at the American Malacological Society meeting in Delaware last July. In his talk (the source of these slides), he argued that a more conservative solution would be to simply add the tribe Lanceolariini to the traditional Unioninae. Such a modification accommodates these new results without loosening the useful morphological characters from the taxonomy. That is the classification we will follow on this web site.

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* Pegias is unquestionably a member of the tribe Anodontini. Even more specifically, it belongs to the eastern Nearctic clade we like to call subtribe Alasmidontina. At least, based on its anatomy and geography (Ortmann, 1914; Williams et al., 2008).

** Like Pegias.*

August 2017

StrophitusStrophitus undulatus (Unionidae, Nearctic)

The August 2017 Mussel of the Month is Strophitus undulatus. Strophitus is a genus of three species, widespread in eastern North America.

There are three species currently classified in the genus Strophitus. S. undulatus, our Mussel of the Month, is widespread in the Interior Basin, Hudson Bay, Great Lakes, and the Northern Atlantic Slope drainages in eastern North America. Its genus-mates, S. connasaugaensis and S. subvexus, are both endemic to waters of the Gulf Coastal Plain. There is some interesting biogeography to discuss there, but we are going to focus on systematics and classification this month. Strophitus is emblematic of the remaining work yet to do on American mussels.

North American freshwater mussels are among the best studied in the world, and Strophitus is well known owing to its broad range. We would have expected that its classification was planted on as firm of a foundation as any other genus. It turns out that is true, but if you read the Mussel of the Month frequently, you know that many genera are pretty rickety. Strophitus is one of those.

Strophitus is a Rafinesque (1820) genus created solely for S. undulatus. The genus was mentioned throughout the 19th century in various lists, as often as not synomyzing it with the then-current concept of Anodonta. Conrad (1853) used Strophitus for a hodge-podge of species now classified in Anodontoides, Alasmidonta, Simpsonaias, Lasmigona, Pegias, as well as S. undulatus. Our current concept of Strophitus dates from Simpson (1900, 1914). He included the three species listed above, plus Anodontoides radiatus and Alasmidonta wrightiana.

Simpson (1900, 1914) recognized Strophitus as distinct based on the arrangement of the glochidia in the marsupium. Lefevre & Curtis (1912:122) described it this way:

“[Strophitus] is unique among the Unionidae in that the embryos and glochidia are embedded in gelatinous cords (called ‘placentae’ by Sterki, ‘placentulae’ by Ortmann), which lie transversely in the gills, whereas in all other cases the egg masses are placed vertically, each one occupying an entire water tube. In Strophitus, on the other hand, the cords are packed closely together, like chalk crayons in a box, a variable number being contained in a single water tube, while the blunt ends of the cords are distinctly seen through the transparent external lamella of the outer gill.”

Both Simpson (1900) and Ortmann (1912) provided descriptions of these structures, but neither were as colorful as the this! See Watters (2002) for a detailed description of those “cords” of glochidia.

These descriptions of such striking reproductive characters were based on S. undulatus, and Simpson (and everyone else) attributed these diagnostic traits to the genus. However, the other species of Strophitus were classified as such based on similarities of the hinge. A new grouping of species was established because they shared such a strikingly unique arrangement of larvae in the marsupium that Simpson coined a new word (“Diagenae”) to describe it, but it was only observed in one species.

That alone does not mean that Strophitus is not a good genus — that it is not monophyletic. After all, having a curved hinge and weak hinge teeth may in fact be shared derived homologies (i.e., synapomorphies) among S. undulatus, S. connasaugaensis, and S. subvexus. However, it turns out that they are not. At least two phylogenetic analysis to-date have included more than one species of Strophitus (Chong et al., 2008; Inoue et al., 2014), and each species appears to share a more recent common ancestor with members of other anodontine genera than with each other. However, for neither of these studies was the phylogeny of Strophitus the objective, and the sleeping dog was left to lie.

So, add Strophitus to the list of genera that would benefit from more attention by taxonomists.

July 2017

VirgusVirgus beccarianus (Hyriidae, Australasian)

The June 2017 Mussel of the Month is Virgus beccarianus. Virgus is a monotypic genus endemic to New Guinea.

Once again, our Mussel of the Month is classified in a monotypic genus. This has been a pattern this year (e.g., Plectomerus, Pseudodontopsis, Protunio, Elliptoideus) as we work our way through the genera we have not yet covered. The classification of Virgus has actually been pretty straight-forward. Simpson (1900) introduced the genus with Unio beccarianus as the type species, and everyone since then has agreed (e.g., McMichael, 1956; McMichael & Hiscock, 1958; Haas, 1969; Graf & Cummings, 2007; Walker et al., 2014). But, that is about the extent of our knowledge: there is a mussel from New Guinea that goes by that name. We know of fewer than 20 museum lots (including the types in Italy, which we haven’t handled), and there is no IUCN Red List assessment for this species.

V. beccarianus is one of eleven species (in seven genera) of freshwater mussels found on New Guinea and nearby islands. Two of the species, in the genus Haasodonta, are currently regarded as unionids, but the rest are classified in the family Hyriidae. Haasodonta, Microdontia, and Virgus are endemic to New Guinea (and adjacent islands), but the genera Alathyria, Hyridella, Velesunio, and Westralunio are also represented on Australia. In fact, two species (see table below) are shared between New Guinea and Australia.

June 2017

DisconaiasDisconaias disca (Unionidae, Nearctic & Neotropical)

The June 2017 Mussel of the Month is Disconaias disca. Disconaias is a largely Mesoamerican genus of six species, found from the Rio Grande of Texas south into Mexico.

Mesoamerican freshwater mussels like Disconaias disca are poorly understood (a frequent complaint on this web site, e.g., Disconaias). Our ignorance is largely the result of a lack of modern attention. The last time the freshwater mussels of Mexico and Central America were “revised” was by Frierson in 1927. Subsequent lists of those species basically just followed him, with minor updates in genus-level classification (Haas, 1969; Graf & Cummings, 2007).

However, over the past semester, University of Wisconsin-Stevens Point undergraduate Elena Hausmann has been reinvigorating the MUSSEL Project effort to update the state of our knowledge of Mesoamerican mussels. For us, this project got underway with some seriousness in 2003 when we started collecting specimen records for the MUSSELp database. For more than a decade, we have been gathering museum records of tropical freshwater mussels, including those of Neotropical Central America and Mexico. In 2014, this long-term project saw a jolt of activity when we helped to curate a large collection of Mexican specimens in the Bereza collection at the Smithsonian. Caitlin Luebke and Rachel Sommer (former UWSP undergraduates) presented the results of that work at the Mollusca 2014 meeting in Mexico City.

Elena’s effort since the start of 2017 has been to work through the MUSSELpdb records from Mexico and assignment to the freshwater ecoregions of the area. The findings were quite interesting. Based on 908 specimen records (as well as the literature), there are 75 specimens in Mexico and adjacent Guatemala. However, these records are not uniformly distributed. In fact, of the 26 relevant freshwater ecoregions, only four account for 84% of the specimen records. These four ecoregions are also the ones with the highest species richness (20-29 species in each).

It was also found that the majority of freshwater mussel species in the Mexico (39 of 75) are endemic to a single ecoregion, and 20 more are limited to two adjacent ecoregions. These results demonstrate just how clumped the diversity of freshwater mussel diversity is in this part of the world.

Elena presented the results of her research to-date at the 2017 UWSP College of Letters and Sciences Undergraduate Research Symposium with a poster, “Mesoamerican Freshwater Mussels (Bivalvia: Unionidae & Mycetopodidae). Phase I: Mexico & Adjacent USA and Guatemala.” We will continue to provide updates as this project progresses. Watch this space!

May 2017

ElliptoideusElliptoideus sloatianus (Unionidae, Nearctic)

The May 2017 Mussel of the Month is Elliptoideus sloatianus. Elliptoideus is a monotypic genus endemic to the southeastern United States.

Elliptoideus is another one of those monotypic genera that may require a little scrutiny. As we have discussed previously, monotypic genera have their time and place. But, on the other hand, there are some such taxa that are just hold-overs from the pre-cladistic era.

For much of the 19th century, there were basically only two genera of mussels that concerned most people: Unio (for species with hinge teeth) and Anodonta (for those without). Those genera are still with us, though over the last century, their constituent species have been whittled down by the removal species to their own genera (as well as tribes, subfamilies, and families).

The earliest placement of Unio sloatianus Lea, 1840 into a different genus was accomplished by Conrad (1853): Plectomerus sloatianus. The genus Plectomerus is still in use, but only for P. dombeyanus, rather than the whole slate of original species that have since been removed to Amblema, Megalonaias, and Elliptoideus. The important point is that Conrad’s system of genera was largely ignored, and U. sloatianus remained in Unio – even after it was classified in the "section" Elliptio Rafinesque, 1819 by Simpson (1900).*

Ortmann (1912) raised Elliptio to a genus, and from that Frierson (1927) split out E. sloatianus into its own subgenus, Elliptoideus, because unlike other species of Elliptio, our Mussel of the Month broods in all four demibranchs. Elliptoideus was raised to a full-on genus by Modell (1942), then he sank it back into Plectomerus (Modell, 1964). Haas (1969a) returned Elliptoideus to a subgenus of Elliptio, though Haas (1969b) classified it as a subgenus of Nephronaias. Heart & Guckert (1970) reinstated Elliptoideus has a genus, and that rank as generally stuck ever since.** Given all the taxonomic flux described in this paragraph, it should be emphasized that these changes were not based so much on new data (that largely ended with Ortmann). Rather, these taxonomists were using classification to reflect their own views of how the data could/should be organized.

In the cladistic era, molecular phylogenetic analyses have recovered E. sloatianus in various positions (e.g., Serb et al., 2003; Campbell et al., 2005), and thus the sister lineage remains unclear. Elliptoideus is apparently a good use of a monotypic genus – at least until we get this stuff sorted out.

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* Despite the obvious phylogenetic distance between true Unio and Elliptio even on anatomic grounds, the legitimate placement of Unio wasn’t correctly sorted until Graf (2002).

** Oesch (1984) inexplicably synonymized the species E. sloatianus with Plectomerus dombeyanus —the species, not just the genera — and this was followed by Howells et al. (1996), and then we were done with that.

April 2017

ProtunioProtunio messageri (Unionidae, Indotropical)

The April 2017 Mussel of the Month is Protunio messageri. Protunio is a monotypic genus from northern Vietnam.

We chose Protunio messageri as Mussel of the Month because we are busy: something easy for which there isn’t much to say. It seems like a perfectly valid species, and we have 23 lots in our database of specimens, all from northern Vietnam.

Ever since Haas (1912) described the genus (the original description was really just applying the name in a plate caption), the species, Protunio messageri has turned up on all the lists (e.g., Thiele, 1934; Haas, 1969; Dang et al., 1980; Graf & Cummings, 2007). Although we are fuzzy with how to classify it in the Unionidae, this is definitely a thing.

But, P. messageri also has an IUCN Red List entry with this interesting taxonomic note:

“The type specimens of Unio messageri do not match the species recorded from Viet Nam, although they come from the same locality. These are possibly two species under the same heading. This assessment refers to the more recently described of the two species.”

So, P. messageri is apparently endangered but it is not the species that has its type pictured here (which matches the other specimen records in our database)? Well, the specimen figured here is Protunio messageri, so the Red List might be talking about something else. Perhaps the species figured by Dang et al. (1980)?

March 2017

DelphinulusDelphinonaias delphinulus (Unionidae, Neotropical)

The March 2017 Mussel of the Month is Delphinonaias delphinulus. Delphinonaias is a genus of four species found in Central America.

Delphinonaias delphinulus is another one of those Central American mussels about which we know very little. That is about all we can say when we honor mussels from that part of world in our monthly showcase (e.g., Psorula, Sphenonaias).

Delphinonaias caught our eye because it has been so unambiguously classified as a lampsiline since Simpson (1900). Even Martens (1900) — who was apparently unaware that Simpson was simultaneously moving mussel generic classification into the modern era — placed those mussels in Unio (Metaptera). Metaptera is a Rafinesque name for the genus Potamilus (also a Rafinesque genus). Presumably, these early classifications with Lampsilis and Potamilus were based on the alate* shape of Delphinonaias, but we wondered if anyone has actually looked at the anatomy. Does Delphinonaias exhibit the characteristic traits of the Lampsilini, like the marsupium limited to a portion of each outer demibranch and capable of expansion when gravid and long-term brooding?

There really aren’t that many references that mention the classification of Delphinonaias, and our quick scan of those did not produce an explicit description of the anatomy of D. delphinulus or any congener. However, Goodrich & van der Schalie (1937) came close. They lumped D. delphinulus under Leptodea paludosa and stated,

“On the whole, this species has all the characters of a Leptodea, both in soft parts and in shell characters.” (p. 47)

Leptodea (also alate*) is a lampsiline. Frierson (1927) and Haas (1969) even arranged Delphinonaias as a subgenus of Lampsilis. We recognized Delphinonaias as a genus (Graf & Cummings, 2007), and we will continue to do that until we have some data to the contrary.

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* Alate refers to the presence of hinge wings.

February 2017

PseudodontopsisPseudodontopsis euphraticus (Unionidae, Palearctic)

The February 2017 Mussel of the Month is Pseudodontopsis euphraticus. Pseudodontopsis is a monotypic genus from the Tigris-Euphrates Basin in the Middle East.

P. euphraticus has a big, good-looking shell, but we don’t know very much about the species. It’s distribution in the Fertile Crescent means shells turn up occasionally in the archaeological literature (e.g., Ridout-Sharpe, 2015). But, the biological classification of the genus Pseudodontopsis is a house of cards.

The species was originally described in the 19th century in the genus Unio. Because of its reduced hinge teeth, P. euphraticus was classified for a while as Pseudodon, Leguminaia, etc., until it got its own genus Pseudodontopsis. Kobelt (1913) gave a long bit of hand-waving to make the case that P. euphraticus (and the various synonyms that he described) was distinct from the other genera in the region, and the name Pseudodontopsis has stuck since then.

However, as we explained last month, monotypic genera often seem like a cop-out — a place-holder for species of unknown affinities. Pseudodontopsis is one of those genera.

Today, we classify Pseudodontopsis in the subfamily Gonideinae based on argumentation rather than data. Both Modell (1964) and Starobogatov (1970) classified Pseudodontopsis in a subfamily called Pseudodontinae,* along with other genera like Pseudodon, Leguminaia, and Microcondylaea. Lopes-Lima et al. (2017) recovered those three genera in a clade with Gonidea, and ipso facto we conclude the Pseudodontopsis also belongs in a clade with the Gonideinae. Unfortunately, the transitive property is not a good basis for phylogeny. What we know about the glochidia of P. euphraticus is consistent with this classification (Al-Mahdawi & Al-Dulaimi, 2009).**

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* We’ll ignore for this post that Modell and Starobogatov classified the Pseudodontinae in the family Margaritiferidae!

** There was a brief, pre-cladistic period in the late 20th century when some Palearctic mussels (including Pseudodontopsis) in the Nearctic genus Pleurobema, but we digress.

January 2017

PlectomerusPlectomerus dombeyanus (Unionidae, Nearctic)

The January 2017 Mussel of the Month is Plectomerus dombeyanus. Plectomerus is a monotypic genus from the western Gulf Coastal Plain of North America, including the lower Mississippi Basin.

Plectomerus is a monotypic genus. A single species in its own genus separate from all other genera,* and that classification has been widely accepted since Ortmann & Walker (1922). That work was among the earliest formal attempts to reconcile the current nomenclature of freshwater mussels with the International Code of Zoological Nomenclature.

We are generally skeptical of monotypic genera. All species are distinct from other species, but genera should reflect information about relationships — groups of species. Sometimes monotypic genera are helpful, but sometimes they are not (e.g., Arkansia). What makes P. dombeyanus so distinctive that two classical anatomists like Ortmann and Walker thought it should be set apart from genera like Amblema and Megalonaias?

Ortmann (1912) had regarded the species now classified in Amblema and Megalonaias as well as P. dombeyanus to belong in the genus Crenodonta based on their shared soft-anatomical characters and similar shell morphology. At the time, though, Ortmann had noted that Frierson (a source of Ortmann’s specimens) had reported that some females of P. dombeyanus brooded their larvae in only the outer gills. The other species of Crenodonta brooded in all four gills. In technical terms, some P. dombeyanus are apparently ectobranchus while others are tetragenous, like the other species of Crenodonta.

Frierson (1914) — an early apologist of crazy Frenchmen — argued that Amblema Rafinesque, 1820 had priority over Crenodonta of Schlüter, 1838. And then Utterback (1915) sequestered the species with zig-zag umbo sculpture into a new genus, Megalonaias. This splitting of mussels with plicate (i.e., broadly wrinkled) shells into new genera forced Ortmann & Walker (1922) to choose a genus for P. dombeyanus. And, they decided — given Frierson’s claim of possibly different brooding anatomy — that it would be better to resurrect Plectomerus Conrad, 1853** than to make the classify it one of the other genera.

Fortunately, Ortmann & Walker (1922) actually fell backwards into the correct answer — or at least the “correctest” answer to-date. P. dombeyanus is not closely related to Megalonaias (Tribe Quadrulini). In fact, in some analyses, P. dombeyanus shares a more recent common ancestor with Lampsilis (Tribe Lampsilini) than Amblema (Tribe Amblemini) (Chapman et al., 2008). Plectomerus dombeyanus is a species with unclear phylogenetic affinities, and to park it in another genus would be more confusing.

Plectomerus is monotypic, and that is the way we like it.

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* Frierson (1927) and others regarded Plectomerus as a subgenus of Amblema, but the sentiment is still the same.

** Conrad (1853) clearly intended Plectomerus as a genus equal in scope to the old Crenodonta, and it included the species now classified as Plectomerus, Amblema, and Megalonaias. However, Conrad neglected to assign a type species, and Ortmann & Walker (1922) applied their ICZN-given right to just pick one. The alternative would have been to pull from obscurity another questionably recognizable Rafinesque† genus, Bariosta Rafinesque, 1820.

† “Rafinesque” is used here as an adjective to describe poorly-described, should-have-been-suppressed taxa. We hope the practice catches on.

 
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