chapter 5dilapidated bridge locations. and suddenly, i knew the answer. bridges and roads are built...

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Chapter 5

Mussel Beds

“Connecting dots is central to answering even the simple questions. I was once asked, ‘Why do you collect mussels at bridges where roads intersect with streams?’ The answer was apparent to me, ‘Easy access.’ The follow-up question related to biodiversity, ‘How can you be confident that you have a good sample of the stream’s mussels and mites?’ This question required a bit of thought and research. I certainly did not want to have a huge hole in the logic of my plan for sampling. With a bit of serendipity, I connected two seemingly unrelated pieces of information. First, I made several forays into streams searching for mussels and mussel beds more than a mile upstream and likewise downstream. Occasionally I found small beds of mussels, but usually not. Secondly, several stories that I had heard mentioned early settlers looking for mussel beds because these made good crossings for wagons. I also recalled that when asking people for help to find mussels, they often took me to washed-out and/or dilapidated bridge locations. And suddenly, I knew the answer. Bridges and roads are built on top of pre-existing roads and trails. Early trails led to river crossings, where large mussel beds had once been and in many cases still persist, at least at the time I sampled. Thus by sampling at bridges, I was sampling the largest beds, aka the best mussel locations, in the streams as found by the early settlers. I know of no Cajun Prairie stories of mussel beds and bridges, but finding mussel beds in the Cajun Prairie was no different than in other areas of the country. I thus assume the same logic was applied when early settlers were looking for safe crossing areas in streams.” (from Vidrine 2010—The Cajun Prairie: A Natural History)

“Freshwater mussels (Bivalvia: Unionoida: Margaritiferidae and Unionidae) make up a predominant benthic community in most waterways in North America, with 50 genera and more than 300 species (Roe and Hoeh 2003, Roe and Hartfield 2005, Graf 2002, and Graf and Cummings 2005).

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Freshwater mussels are among the most endangered animals in freshwaters (Bogan 1993 and Strayer et al. 2004). Freshwater mussels once cobbled the stream bottoms in Louisiana (Vidrine 1993). The development of canal systems and lakes during the last century provided additional habitat—often likewise cobbled with mussels that formed diverse communities (Vidrine and Vidrine 1987, Vidrine and Cordes 1994, Vidrine and Quillman-Vidrine 1994, and Vidrine and Borsari 1994).” (from Vidrine 2008—Red River paper) “The future of freshwater mussels has become entirely dependent upon the further actions of man. The best possible scenario for the future of mussels depends upon the development of an informed public and a supported scientific effort. Although Louisiana contains a very diverse faunal assemblage, literally no funds are available for the study of mussels. In a recent effort to develop a controlled mussel harvesting industry in the state, the over-riding concern was the tremendous lack of information on the locations of this state's communities and populations, and, further, the near total absence of biological information on the life histories and requirements for the survival of the mussels. This book begins the education process and provides general information in regards to the first deficiency, the lack of mussel distribution records. However, the "historical" in the title means just that in the largest sense of the word. Most of the records provided are at least ten years old. During the last couple of years, I have revisited some of the stations that contained the most diverse mussel assemblages. The results were devastating, as in many of these stations, few mussels remained. For example a striking memory involves the return to the Louisiana Irrigation Canal system in southwest Louisiana, where my son, Macky, and I had sampled during the early 1980's. This irrigation canal system was abandoned by local rice farmers, and the canal bed was dry and densely overgrown with willow seedlings taller than I am. Unfortunately, most bayous and rivers are used as sewers and agricultural runoff ditches, and my return to many of these now channelized canals deepened my concern.” (from Vidrine 1993—The historical distributions of freshwater mussels in Louisiana)

General Introduction

Mussels once cobbled the streams of the eastern half of the United States, in many locations forming beds with thousands if not millions of individuals. Streams, even those of southern Louisiana called bayous and known for

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their muddy waters, were clear and for the most part gravel or sand-bottomed and served as the source of drinking water for animals and man. These clear waters were the product of these mussel beds as each mussel filtered gallons of water each day—what an ecological service! The beds latter would provide a passageway for wagons, seeds for pearls, food for animals as once they had been food for Native Americans as evidenced in the shell middens, shells and natural pearls for jewelry and money as once they were a form of exchange and numerous other ecological services. These mussel beds are among the least remembered of all the treasures of the North American wilderness, and although mussels and mussel beds were as abundant as the passenger pigeons, they are essentially gone. The largest mussel bed ever found was Muscle Shoals, Alabama (Haag 2012—pages 319-322). These mussel shoals extended more than 50 miles and were a complex of rocky ledges and shallow, gravelly shoals in the Tennessee River. Estimates of width of the river before damming was 2-3 miles with many small islands surrounded by shallow shoals with beds of mussels—altogether 70 species. First humans constructed a canal through it (early 1900s) as it was too shallow to navigate, then we constructed a dam (Wilson Dam in 1924) on top of it, thus placing the beds under a hundred feet of water and eventually nearly 20 feet of silt. While the first dam destroyed only a small half of the beds, 2 additional dams (Wheeler and Pickwick in the 1930s) did in most of the rest of the beds. Countless other mussel beds have succumbed to similar treatment or served as crossings in small creeks and rivers as they provided a solid foundation for wagons. Sedimentation from urbanization and agricultural activities smothered many more beds. Overharvesting by pearl fishermen and modern day shell fisherman for seed pearls do/did in many beds. Chemical pollution from a variety of sources continues to eliminate mussel populations. Aside from this endless stream of tragedies, we want to focus on the idea that mussel beds are the evolutionary units, when thinking about mussels and mites, and as such these are primary topics in a book on mussels and mites. Communities evolve and as such their constituent organisms serve as the intimate units in this evolution, much as the nucleic acid bases in evolving DNA/RNA sequences or amino acids in protein sequences—the differences are mainly in terminology, but the idea of community evolution is foreign and unfortunately readily discounted by organismal biologists. Unlike organisms, communities are at present just too complex to

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genuinely ponder from an evolutionary perspective, but that will not prevent some commentary. Evolution within the context of a community simply implies that mussels and mites are coevolving within a community consisting of many species of mussels and mites. The evolution of mussels and mites is far more complex in that the mussels are parasites of fish as young, while the mites are parasites of chironomids as young. Thus, we have at minimum 4 very disparate groups of organisms intricately interwoven—fish, mussels, mites and chironomids—and this is still a fairly linear outline of this community contained within a mussel bed. The fish are famous for gathering for mating in shoals, and mussels have evolved mechanisms to literally ensnare the fish into hosting their young—the most extravagant of these mechanisms involves modification of the mussel mantle flaps into fish, worm and insect mimics to aggressively attract fish interested in mates and/or food. Much is now known about these mussel adaptations (Haag 2012 and Zanatta and Murphy 2006). Unfortunately, much less is known about the mite and chironomid associations, but mussel and mite associations are our focus. While there are many known associations between mussels and mites, in many ways the search has just begun. Andrew C. Miller and Barry S. Payne have described numerous mussel beds using a variety of statistical methods of sampling. Check out some of their references. Haag (2012) explores both mussel assemblages and mussel bed creation as a function of macrohabitat and microhabitat levels using abiotic factors including stream size, upland vs lowland, substrate type, lenthic vs lotic, geographical history, stability of the bottom and impacts of extremes such as frequency and extent of flooding and drying down. He also explores numerous biotic factors, including fish hosts and coevolution of mussel-fish communities. And finally, he discusses man-made impacts, such as sedimentation, exotic invasives, agriculatural and industrial and urban pollution, channelization, and much more.

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Studying Mussel Beds

Possibly the best way to introduce mussel beds is to tell stories about them. Since Vidrine has sampled a fair number of them, examples of beds Vidrine has searched are here described from his perspective. The actual data from these beds are published in a number of papers and books. Several stations are included from which mite data from Canada and Mexico can be added to our discussion. For comparison, we will initially follow the Appendix 5--Descriptions of mite-mollusk associations in sampled stations in Edwards and Vidrine 2013c.

These stations were sampled usually several times. The data is here

collated into single tables. The goal is to demonstrate the complex

communities that occurred in streams and still occurs in several streams.

We could also show a series of companion locations that have been greatly

altered, and the communities are significantly altered, e. g., the mussels

are greatly reduced in variety and/or numbers; the mussel community

remains but the mites are significantly decreased or entirely extirpated; or

the stream has been entirely changed by engineering (damming,

channeling, diversion, sedimentation, pollution, etc.).

Understanding these mites involves some exposure to them in the wild, i.e.,

in nature. It is exciting to visit a stream or lake and spend an afternoon

raking the sediment with your hands. The first excitement is finding the

mussel. The further excitement comes upon the discovery of these

remarkable mites in the mussel. And the coup de gras is the discovery of

possibly 3 or more species in the mussel, where each species is doing

something entirely different in the mussel—resource partitioning. And the

ultimate experience is to discover that one of the mites is a new record for

the host species or for the location, but the über-ultimate experience is to

find a species new to science. But just the visit to the stream or lake is a

great place to begin. And now that we can expect to find ‘cryptic species’

not only among the mussels but also among the mites that we collect, the

impacts of our efforts take on exponentially greater importance and

potential usefulness in understanding the nature of mussel beds.

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Map xxx. Stations (25) examined in this appendix.

Stations

Twenty-five (25) stations across North America are presented. Many of

them are ‘natural’ or modified as reported and present community

structures that we suspect would occur. Several stations are obviously

impacted either by agriculture or by coal mining. In some cases, the mussel

populations of rarer species are thought to be so diminished as to

negatively affect parasite mite community diversity. In other cases, the

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mussel community appears to be healthy, but the mite community is greatly

impacted if not completely extirpated.

1. Louisiana. Allen Parish. Calcasieu River at Indian Village boat landing.

2. Louisiana. Rapides Parish. Calcasieu River along LA Highway 112 between LA Highways 121 and 113 (ca. 10 miles of river).

3. Louisiana. Jefferson Davis Parish. Louisiana Irrigation Canal along U. S. 90 and above I-10.

4. Texas. Hardin and Jasper Counties. Neches River at U. S. 96. 5. Texas. Hardin County. Village Creek at U. S. 96. 6. Louisiana. St. Landry Parish. Bayou Courtableu (West Atchafalaya Basin levee

burrow canal) ca. 0.5 miles north of U. S. 190. 7. Louisiana. Rapides Parish. Brown and Loving Creeks. 8. Louisiana. Natchitoches, Red River and DeSoto Parishes. Bayou Pierre. 9. Louisiana. Morehouse Parish. Bayou Bartholomew at U. S. 425. 10. Arkansas. Montgomery and Polk Counties. Ouachita River at U. S. 270 and up-

stream to McQuire Public Landing. 11. Arkansas. Sharp County. Strawberry River at U. S. 167, ca. 2 miles north of Evening

Shade. 12. Tennessee. Bedford, Coffee, Marshall, Maury and Hickman Counties. Duck River at

several stations. 13. Tennessee. Davidson and Rutherford Counties. Stones River at several stations. 14. Tennessee. Hancock County. Clinch River at Kyles Ford. 15. Tennessee. Claiborne County. Powell River near Hoop, ca. 10 miles northeast of Tazewell. 16. Mississippi. Amite County. East and West Fork Amite River. 17. Louisiana. St. Helena Parish. Twelve Mile Creek (Tangipahoa River drainage) at LA 1045. 18. Florida. Holmes and Washington Counties. Holmes Creek (Choctawhatchee River

drainage) at U. S. 90. 19. Florida. Gadsden and Leon Counties. Ochlockonee River at FL 263, U. S. 27 and U. S. 90. 20. Florida. Sumpter and Hernando Counties. Little Withlacooche River at U. S. 301. 21. Florida. Okeechobee and Highlands Counties. Kissimmee River at U. S. 98. 22. Canada. Ontario. Dufferin County. Amaranth Twp. Willow Brook 2.2 miles east-northeast of

Grand Valley. 23. Canada. British Columbia. Vereux Lake 8.5 miles north of Oliver. 24. Texas. Brazos, Grimes, Madison, Robertson, Leon and Limestone Counties. Navasota

River at 26 stations. 25. Mexico. Tamaulipas. Small river (locally called Arroyo los Gatos and presumably a

headwater of the Casas Viajas River system) at MX 80 in Nuevo Morelos.

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Descriptions of sites and communities:

1. Louisiana. Allen Parish. Calcasieu River at Indian Village boat landing.

September 3, 1983. Gail Quillman, Macky Vidrine and M. F. Vidrine.

October 20, 1991. M. F. Vidrine.

July 15, 2011. John Hamlin and M. F. Vidrine.

October 25, 2012. M. F. Vidrine, Dale Edwards and Blaine Vidrine.

Host # searched Mite # infected

Plectomerus dombeyanus 54 Unionicola megachela 20

U. tupara 10

Leptodea fragilis 3 U. hoesei 2

U. austalindistincta 2

U. abnormipes 1

Lampsilis hydiana 26 U. hoesei* 26

U. abnormipes 26

Lampsilis teres 30 U. hoesei* 30

U. abnormipes 30

Lampsilis satura 1 U. hoesei* 1

U. abnormipes 1

U. serrata 1

Potamilus purpuratus 10 U. hoesei* 10

U. austalindistincta 9

U. fulleri 9

Glebula rotundata 30 U. hoesei 28

Amphinaias mortoni 50 U. vikitra 31

U. vikitrella 19

Amblema plicata 52 U. amandita 26

U. tupara 6

Villosa lienosa 10 U. gailae 6

U. serrata 2

Obliquaria reflexa 2 U. vikitra 2

Pyganodon grandis 9 U. formosa 2

U. mitchelli 3

U. megachela 1

Toxolasma texasiensis 6 U. abnomipes 1

Quadrula apiculata 1 U. vikitra 1

Quadrula verrucosa 7 U. vamana 7

U. hoesei 2

U. abnormipes 1

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Utterbackia imbecillis 3 U. foili 2

Campeloma decisum 15 U. campelomaicola 1

Obovaria jacksoniana 28 none

Pleurobema riddelli 5 none

Toxolasma parvum 10 none

Ligumia subrostrata 5 none

Fusconaia askewi 3 none

*Four cryptic species of Unionicola hoesei were recognized by Ernsting et al. (2014) from the collection in 2012 from this station.

This free-flowing stream is one of the most natural remaining in Louisiana. This fairly large sandy-bottomed stream with muddy backwaters hosts a diverse assemblage of mussels and mites. The river and its watershed are among Vidrine’s favorite stations to visit and sample. Every visit has brought to light a new record for a mussel or a mite. Beds of mussels usually 3-5 feet wide line the banks usually on one side or the other, while shifting sands known locally as sand bars moves in the center of the stream and host few mussels. The station is illustrated in a photo taken by D. D. Edwards on the back cover of the 2013 Mites of Freshwater Mollusks book. Not only are the mussels parasitized by a diverse fauna, but also a single snail, Campeloma decisum, was found in 2012 with a single Unionicola campelomaicola (Wei Li, personal communication 2013)—these associations not only extend throughout this river drainage but apparently extend to the east and west in other drainages. The Calcasieu River drainage is a complex network of both small and large creeks and man-made canals that flood rice fields, including Louisiana Irrigation Canal (below). This sandy system contains numerous small creeks with headwaters in the Kisatchie National Forest and in the Fort Polk Military Reserve Training Posts. A number of these creeks had small beds of mussels. The river has shifting sand bars, but linear mussel beds form on one or the other bank and around the bases of Bald Cypress trees in the river or on its banks. The series of beds at the Indian Village landing has 21 species of mussels in linear narrow beds (ca. 5 feet wide) that we sampled for at least 200 yards. The center of the stream had shifting sand bars and few mussels. The mussels are buried deeply in the sand, with Plectomerus dombeyanus being most numerous, and Amblema plicata and

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Amphinaias mortoni forming close seconds. Many of the mussels are infected with mites. This station in one very specific way is the impetus for this book. Four cryptic species of Unionicola hoesei were recognized by Ernsting et al. (2014) from the collection in 2012. Each cryptic species was associated with a single host species at this location. Glebula rotundata and Leptodea fragilis also had populations of U. hoesei in this location but these were not evaluated in that study. The implications of this finding involve re-evaluation of the numerous reports of U. hoesei from Canada to southern Mexico in more than 15 recognized genera of mussels. Could literally dozens of cryptic species be involved? However, the discovery of these cryptic species indicates that morphologically similar populations currently placed in a single species could not only represent many species but also provide a mechanism for evaluating the host groups. Host groups with similar mites may be hypothesized to comprise a single evolutionary radiation; thus we can approach mussel systematics by in part delineating mite species cryptic among currently recognized morphological species. 2. Louisiana. Rapides Parish. Calcasieu River along LA Highway 112 between LA Highways 121 and 113 (ca. 10 miles of river). July 20, 1974. Mark S. DeRouen, Blake Vidrine and M. Vidrine. October 5, 1974. Blaine Vidrine and M. Vidrine. August 25, 1978. Daniel J. Bereza, Selwyn Roback, and M. Vidrine. October 4, 1986. Macky Vidrine and M. Vidrine. October 11, 1986. M. Vidrine. August 3, 1991. Daniel Vidrine and M. Vidrine. September 21, 1991. Bruno Borsari and M. Vidrine. May 27, 2005. M. Vidrine.

Host # searched mite # infected

Strophitus undulatus 43 Unionicola dimocki 39

U. abnormipes 1

Campeloma decisum 171 U. campelomaicola 14

Pleurobema riddelli 37 U. gowani 16

N. ingens 3

Lampsilis teres 19 U. hoesei 19

U. abnormipes 11

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Uniomerus declivus 35 U. stricta 6

N. ingens 9

Lampsilis satura 1 U. hoesei 1

U. abnormipes 1

Leptodea fragilis 1 U. serrata 1


N. ingens 3

Ligumia subrostrata 2 U. serrata 1

Fusconaia askewi 486 U. parkeri 154

U. serrata 119

N. ingens 7


U. serrata 8

U. abnormipes 1


U. serrata 2

U. abnormipes 1

N. ingens 3

Lampsilis hydiana 229 U. hoesei 220

U. abnormipes 79

U. serrata 3

N. ingens 1


U. serrata 86

U. abnomipes 1

U. laurentiana* 33

Toxolasma parvum 66 U. kavanaghi 43

U. serrata 2

N. ingens 9

U. laurentiana* 1

Obovaria jacksoniana 100 U. gailae 1

U. abnormipes 1

Pyganodon grandis 3 none

*Unionicola (Unionicola) laurentiana is typically a sponge mite—its occurrence in these mussels, where they were numerous chrysalids and newly hatching deutonymphs and adults in the suprabranchial chambers.

These 3 stations are combined and represent another free-flowing and near natural section of the stream in its upper to middle part of the watershed. The sand and mud bottom apparently shifts, while the mussels lodge themselves in beds along the banks and around fallen timber or

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cypress stumps—the places one usually finds schools of fish. Both mussel and mite diversities are noteworthy in this classic bayou-like stream. Vidrine spent his early years studying this and nearby streams (altogether more than 100 stations) in order to unravel the links between mussel and mite diversity. Along this 10 mile stretch of serpentine river, many areas have Bald Cypress trees in the stream or along the banks. This section of river is about 30 miles north of Indian Village—much of the intervening 30 miles has few mussels as it was obviously polluted by the lumber/paper mill at Elizabeth, Louisiana. Mussels form very small beds along the banks and around the trees. The large bed forming mussels like Plectomerus dombeyanus and Amblema plicata were absent or uncommon, but many Lampsilis hydiana, Villosa lienosa, Fusconaia askewi, Obovaria jacksoniana, Toxolasmus parvum, Amphinaias mortoni, Strophitus subvexus and Uniomerus declivus populate the sandy bottom. This cool, clear stream is readily canoed, but we sampled at the intersection of the river with bridges. Again many mussels were parasitized by mites. Another station (Calcasieu River at U. S. 190 ca. 8 miles north of Indian Village and west of Kinder) was actually measured and reported in Vidrine and Clark (1981), in which they characterized the freshwater mussel bed as follows: “The Calcasieu River is moderately large and is characterized by a shifting sand bottom. Mussel beds (aggregations) occur along the banks of the river; one such bed was analyzed in order to characterize the nature of the diversity (species composition) and relative abundance of the mollusks present. A 28 m2 area was sampled, and the introduced Asian bivalve, Corbicula fluminea (Müller) (Corbiculidae), was the most common mollusk. Twenty-one endemic freshwater mussel species occur in this bed.” The species in the list include those listed in stations 1 and 2 above. 3. Louisiana. Jefferson Davis Parish. Louisiana Irrigation Canal along U. S. 90 and above I-10. May 2, 1981. Macky Vidrine and M. Vidrine. February 1, 1984. M. Vidrine. February 11, 1984. Gail Vidrine, Macky Vidrine and M. Vidrine. May 1, 1984. M. Vidrine.


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Utterbackia imbecillis 2 Unionicola foili 2

U. megachela 1

Toxolasma texasiensis 24 U. megachela 3

U. causeyae 3

U. aculeata 2

Arcidens confragosus 4 U. megachela 1


U. abnormipes 12

U. aculeata 3

Plectomerus dombeyanus 37 U. tupara 8

U. megachela 31

Quadrula apiculata 36 U. megachela 22

U. vikitra 29

U. aculeata 2

Amphinaias mortoni 12 U. megachela 1

U. vikitra 9

U. aculeata 3


U. megachela 1




U. fulleri 12

U. megachela 1

Potamilus purpuratus 17 U. hoesei 3

U. australindistincta 13

U. fulleri 17

Viviparus subpurpureus 43 U. viviparaicola 7

Truncilla donaciformis 3 none

I drove by this canal several times a week while working for the Jefferson Davis Parish Mosquito Abatement District (1980-1984) as I searched the parish for mosquito populations and monitored district activities attempting to moderate their populations. I spent the winter months thinking that there must be a noteworthy mussel community in this canal, and Macky and I decided to find out. This man-made canal is one of nearly a hundred named canals built during the late 1890s and the early 1900s as rice was developed as a primary crop in the Cajun Prairie region of southwestern Louisiana. The need for water required a network of canals, which are/were owned and cared for by local corporations or individuals even to this day, to carry water from local rivers and now pumps to individual rice fields. These

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canals host dynamic mussel communities; however, they are very varied from only a few species to highly diverse as in the Louisiana Irrigation Canal. The bottom was a silty-mud with literally thousands of mussels scattered throughout the slow flowing, uniformly bottomed canal. The mussel and mite assemblage of this canal was very similar to local bayous, but its source was the Calcasieu River itself. This canal is no longer managed—underground irrigation has replaced it in the region—the mussels and mites are gone as most of the land beneath the canal has been reclaimed for agriculture. My son, Macky, and I spent 2 days sampling this canal at several highway intersections and other locations (Vidrine and Vidrine 1987). We could easily have filled many bags with mussels and snails. Large populations of Rangia cuneata, a estuarine clam that can survive in freshwater if their larvae are transported into these waters, were found at several locations. Since the water was pumped out of the lower Calcasieu River, we assumed that the larvae were pumped into the canal. These clams had no mites. We were amazed by the miles of canal with mussels and overwhelmed with the idea of searching the entire canal in order to find all of the species that may have been living there. Each time we were ready to give up and head home, we would find a population of another species.

4. Texas. Hardin and Jasper Counties. Neches River at U. S. 96. January 5, 1978. Daniel J. Bereza, Darryl Clark, Mary Curry, Jim Leemann, Betty Everitt and M. Vidrine. June 2, 1978. Bill Bell, Don Gowan, Darryl Clark and M. Vidrine. August 22, 1978. Daniel J. Bereza, Selwyn Roback and M. Vidrine. August 15, 1980. Darryl Clark, Mark LaSalle, Macky Vidrine and M. Vidrine.





U. fulleri 5

U. abnormipes 1


U. abnormipes 33

U. aculeata 5


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U. abnormipes 37


U. abnormipes 3


Uniomerus declivus 5 U. stricta 1

U. abnormipes 1

N. ingens 2


Megalonaias nervosa 13 U. tupara 8


Amblema plicata 109 U. tupara 81



U. abnormipes 3



Truncilla donaciformis 15 U. hoesei 1

U. amandita 1

Arcidens confragosus 6 U. belli 2



U. vikitrella 26

Quadrula quadrula 12 U. vikitra 8

U. vikitrella 10


U. vikitrella 35

Potamilus amphichaenus 9 U. hoesei 2


Viviparus subpurpureus 50 U. viviparaicola 25

Ligumia subrostrata 1 none


Fusconaia askewi 7 none

Amphinaias nodulata 4 none


Utterbackia imbecillis 1 none

Although impounded upstream, the river is free-flowing in this section with a sandy bottom. The river commonly overflows, and several canal-like, backwater streams are present with mussels and mites. Both the river and these backwaters were sampled. Similar to the Calcasieu and Sabine

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River to the east and the San Jacinto River to the west, this stream and its tributaries, including Village Creek, indicate the diversity of mussels and mites in the western Gulf drainages. This is a big river with mussels embedded in the banks and backwaters. Twenty-five species of mussels were found. The snail Viviparus subpurpureus was also present and parasitized. 5. Texas. Hardin County. Village Creek at U. S. 96. August 12, 1977. Macky and M. Vidrine. December 28, 1977. Daniel J. Bereza and M. Vidrine. January 5, 1978. Daniel J. Bereza, Darryl Clark, Mary Curry, Betty Everitt, Jim Leeman and M. Vidrine. June 2, 1978. Bill Bell, Don Gowan, Darryl Clark and M. Vidrine. August 21, 1978. Daniel J. Bereza, Selwyn Roback and M. Vidrine. August 15, 1980. Darryl Clark, Mark LaSalle, Macky and M. Vidrine.




U. megachela 14


Fusconaia askewi 82 U. parkeri 48

U. serrata 39

N. ingens 2

Pleurobema riddellii 165 U. gowani 90

U. aculeata 2

N. ingens 1


U. abnormipes 9


U. abnormipes 7

N. ingens 1


U. abnormipes 22

U. serrata 23

Potamilus purpuratus 2 U. serrata 1

Ligumia subrostrata 1 U. hoesei 1



U. megachela 3

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U. hoesei 1

Megalonaias nervosa 1 none




Quadrula quadrula 1 none

Quadrula apiculata 2 none

Quadrula verrucosa 9 none

Uniomerus declivus 1 none

This sand and occasionally gravel-bottomed stream was shallow and free-flowing. The stream appeared as many do in southeastern Texas and western Louisiana, and it had a similar assemblage of mites and mussels. Mussels were numerous and diverse. However, the mite community was obviously impacted by the presence of larval Ablabesmyia janta, a predatory chironomid that routinely occurs within the mantle cavity (usually between the gills) of mussels in this locality. As a result, many mussels that were inhabited by the chironomid had no mites in contrast to those of the same species with mites and without the chironomid (usually one per mussel). The station was the central focus of 3 papers (Roback 1982; Roback et al. 1980; Vidrine 1990a).

6. Louisiana. St. Landry Parish. Bayou Courtableu (West Atchafalaya Basin levee burrow canal) ca. 0.5 miles north of U. S. 190. August 18, 1978. Robert S. Parker, Beverly Williams and M. Vidrine. August 24, 1979. Daniel J. Bereza, Darryl Clark and M. Vidrine.


Amphinaias nodulata 4 U. vikitra 3

U. megachela 3

Utterbackia suborbiculata 5 U. foili 2

U. aculeata 2

U. megachela 2

U. viviparaicola 1

Potamilus purpuratus 12 U. aculeata 11

U. hoesei 11


U. megachela 11

U. fulleri 12

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U. abnormipes 15

U. megachela 9

Toxolasma texasiensis 8 U. causeyae 4



U. megachela


U. megachela 8


U. abnormipes 8

U. megachela 4



U. abnormipes 7

U. aculeata 7

U. fulleri 3



U. aculeata 1

U. megachela 1

U. abnormipes 1


Amphinaias pustulosa 8 U. vikitra 8

U. megachela 3


U. megachela 2


U. megachela 3


U. megachela 8

Arcidens confragosus 17 none

This man-made branch of the bayou was dug to build the western levee of the basin—it was shallow and literally filled with mussels prior to harvesting in the mid and late 1980s and 1990s. Our sampling occurred before the second harvesting. Many of the mussels were very large in size and commonly had more than one species of mites. The stream bottom was a silty mud, and the ‘canal’ was only a few feet deep in late summer.

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We musseled/pollywogged the lower end just north of Rte. U. S. 190 and west of the Atchafalaya levee. We had knowledge and pictures of some mussel harvesting during the mid-1970s in the mid-reaches of this canal (George et al. 1993--Louisiana Conservationist—I was working for GSRI on leprosy in armadillos and did not sample until I attended ULL then USL), so we sampled the lower end. This lower end apparently was not harvested in that foray of harvesting, and the mussels were abundant in a muddy bottom. Whereas our sampling along the lower reaches of this canal south of Rte. U. S. 190 and all the way down to Interstate 10 and south, most of the beds were small and dominated by Glebula rotundata. Diving showed that mussels were common at depths beyond our reach, but a large amount of boat traffic caused a lot of wash on banks and shallow shorelines. These inimical conditions were absent at this station, and mussels were commonly implanted in the mud side by side. Although no measurements were made, at least 50 mussels per square meter were present throughout the relatively flat bottom of the canal, which was less than 3 feet deep in this end of the canal and extremely difficult to move in with a large sack of mussels. We would sit on the bottom or drag along with our hands identifying the mussels by their size and ridges, pimples, roundedness, pointedness or sharp edges and only removing those individuals that we wanted to collect or individuals that our fingers could not identify. While smaller mussels could easily be removed from the substrate, larger mussels required both hands and a good yank. This most remarkable bed was a 5 mile stretch along the western levee borrow canal (an arm of Bayou Courtableu). Mussels cobbled the bottom; unfortunately, shell fishermen arrived in the mid-1970s (photo of mussel shells in burlap sacks after boiling and drying in piles in George et al. 1993 in the Louisiana Conservationist magazine) and removed approximately a hundred tons of shells for the cultured pearl industry. I recall piles of shells nearly 20 feet high and the horrible smell of the boiled, rotting flesh piled along the banks. The stench and the loss of life overwhelmed me. But my goal is to describe the bed before it was harvested. Megalonaias, the size of my open hand, were common, but there were some that were almost impossible to remove from the soft mud surface as they were dug into the clay bottom and were the size of both of my hands put together. Very large Plectomerus and Potamilus were initially ignored by the shell fishermen as these shells have purple nacre, which greatly reduces the value of the shells—pearly white nacre is prized. Pyganodon, Lampsilis, Utterbackia, Toxolasma and other smaller shells were ignored at first by the shell

20

fishermen, as the shells were not suitable for pearl seeds nor within the legal sizes. However, they wised up at some point and took all the mussels. When I returned to the site, the piles of shells were gone, but there now was several hundred 55 gallon drums neatly arranged along the road between the canal and the Atchafalaya levee road. Naturally, I asked, ‘what was in the barrels?” The drums/barrels were filled with the non-commercial mussels pickling in formalin for use by high school students across the country. Recall high school and college students were required to take a course in biology, and most schools could afford a barrel of mussels such that each student could cut open a pegged mussel and look at the strange parts inside. I am sure many still marvel at the experience and tell stories of the event to their grandchildren. Thus the mussels not harvested for pearl seeds went into the barrels—damn near every one of them. My only solace from these days was a trip in the midst of all this harvesting where the place was nearly vacant. Again I asked. Many of the divers were hospitalized as the canal apparently hosted a fairly nasty worm locally called ‘nutria itch,’ which burrows into the skin commonly around the waist and moves around looking for a good location, meanwhile causing a terrible itching and obvious linear lesions under the skin. In a weird kind of way, this may have been the only defense that the mussel bed had to resist the invasion of the shell fishermen. Again in the 1980s and 1990s, the shell fishermen returned to remove any new mussels that survived or colonized the area. All in all, the bed literally evaporated and got shipped either to Asia as seed pearls or to high schools for informal dissection. I have never returned for fear that the bed would not have returned—maybe I can add a return visit to my bucket list. Of the 18 species of mussels commonly found there, mites were common in all species except Arcidens confragosus.

Exposed posterior surfaces of the shells of mussels are commonly covered with a lush growth of organisms. This condition was extremely common here and among small beds in the eastern borrow canal near Ramah, Louisiana and extending from U. S. 190 to below I-10 in my searches. The canal is just outside the Atchafalaya Basin levee and served as a source of soil for levee construction. I recall a colony of Pectinatella magnifica nearly 2 inches in diameter on a shell of Pyganodon grandis, but noted larger colonies, some 8-10 inches along branches submerged in the canal—not to mention lots of sponges, entoprocts, ectoprocts, hydroids and algae. These slow moving or nearly stationary waterways with less boat traffic developed extraordinary mussel beds with luxuriant and diverse growths on exposed shell surfaces (Vidrine 2008).

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7. Louisiana. Rapides Parish. Brown and Loving Creeks. October 1, 1973. Samuel L. H. Fuller and M. F. Vidrine. August 17, 1977. James Jackson, J. Alan Pounds and M. F. Vidrine. Host # searched mite # infected

Margaritifera hembeli 31 none

Fusconaia flava 12 none

Villosa lienosa 2 none

Rapides Parish District Kisatchie Forest creeks had Margaritifera hembeli, Villosa lienosa and Fusconaia flava beds in central Louisiana. The beds were Margaritifera beds for the most part and contained hundreds to thousands. The mussels were found in the center of the sandy creeks, with hundreds of posterior ends sometimes obvious in the shallow water. Other clusters of mussels were in deeper holes into banks, such that you could reach up and under banks and feel 4-5 mussels with a single hand. My first visit was with Sam Fuller in 1973 was seminal in shaping my interest in mussels and their mites—he spoke at length about how important such research would be, and he went out of his way to cite my first paper in his book (Hart and Fuller 1974—Pollution Ecology of Freshwater Invertebrates). Both the Villosa and Fusconaia occasionally had mites in these and other creeks in the Kisatchie Forests, but none were found in Margaritifera. In the end, hundreds of margaritiferid mussels were studied by Doug Smith, and he reported that he had never seen mites in North America—also note that margaritiferid mussels from Eurasia and Africa also have had no reports of Unionicola infesting them. Many creeks were searched in the Vernon District of the Kisatchie Forest, where many species of mussels occur with impressive mite communities—however, the mussel beds were small and usually covered a couple of square meters at best. 8. Louisiana. Natchitoches, Red River and DeSoto Parishes. Bayou Pierre at Hwys. 485, 509, U. S. 84, 177 and 174—a 40 mile stretch between Shreveport and Natchitoches. August 22, 1974. M. F. and Blake Vidrine. September 1, 1974. M. F. and Blake Vidrine. April 25, 1976. Samuel L. H. Fuller, Daniel J. Bereza and M. F. Vidrine.

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Amphinaias nodulata 4 U. vikitra 2




Plectomerus dombeyanus 17 none




U. abnormipes 12

Potamilus purpuratus 71 U. abnormipes 2

U. hoesei 39


U. fulleri 12


U. abnormipes 2







Arcidens confragosus 25 U. tupara 1

Toxolasma texasiensis 2 none



Truncilla truncata 5 none

Truncilla donaciformis 5 U. vamana 1

Bayou Pierre, an arm of the Red River, is a small river that was nearly dry in 1974, when my brother, Blake, and I decided to sample it. Several significant bridges crossed the bayou and provided easy access. Although very muddy, the stream was shallow and fordable with some effort in the deep mud sediments. The river had been dredged in the 1950s and resembled a canal in its appearance.

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Mussels were cobbled in so thick that it was difficult to pull one from the substrate—you had to yank 3 out at a time. The mussels were mostly large and overgrown with luxuriant growths on any exposed surface. Lorraine Screven Frierson (see my popular article) had written a number of articles on his life on the bayou, where he owned and ran a cotton gin and plantation. Megalonaias, Plectomerus, Amblema, Lampsilis, Potamilus, Amphinaias, Glebula and Quadrula were large and abundant. In a few minutes, you could have filled a sack that you could not carry out in the deep mud. So we moved about while digging our fingers into the beds in search of mussels different from those above—the big bed forming species. Corbicula, the Asian weedy clams, were thick in and among the native mussels. A good number of dead shells on the muddy banks signaled the activity of local mammals feeding on mussels or at least dislodging them. I was reminded of Frierson’s story of Uniomerus declivus drying on the banks and embedded in the banks and surviving there during dry seasons for months. However, I could not find a single Uniomerus among the tens of thousands of specimens that I handled or at least touched. I did bring home several dozen mussels from another canal (Quadrula apiculata and Glebula rotundata) on a bet with Thomas Dietz, who was interested in dessication physiology. I laid the mussels out on my desk and weighed them each day for a month—they lost about a gram a day. After that month, I dropped them in Tom’s aquarium, and within several hours they gapped and sprung to life—and I won the bet. African mussels are famous for living an entire year buried in dry soil that once was a stream only to return to life as if by magic upon the return of the wet season and the stream—some of our mussels would give them a run for the money. By contrast, Villosa lienosa cannot survive but for an hour out of running water—there is always an exception to every rule. So Bayou Pierre, Frierson’s arm of the Red River, runs more than 60 miles and had a remarkable mussel community. I have not returned to the river except for a short foray with Dan Bereza and Sam Fuller in search of material for molecular assay back in 1976. They too marveled at the dense beds of mussels along the 20 foot wide main stream bottom. The mussels were of no interest to shell fishermen as the shells were usually heavily eroded and possessed dark lines in their nacre—a no-go for pearl seed production. Apparently the drying down and extreme conditions during the summer has a lasting impact on the periostracum (often colored protein covering of the nacre of the shells)—these mussels for the most part were jet-black in color and the typical yellow to straw coloring common in many

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mussels was stained dark with tannins and other chemicals in the mud and water. In this cotton country, it was obvious that many mussels lacked mites. I would not understand this circumstance until years later when I studied the Bayou Bartholomew and Tensas River in detail—these lie at the heart of modern cotton production and simply lack mite populations in their mussels. We were lucky in 1974 in that we did find some mussels with mites in several of the stations, although the overall infection rates were exceptionally low. One location had no mites at all, and two others were generally with low infection rates. Two stations, Rte. 174 and U. S. 84, had relatively high infection rates (ca. 50%). These long beds, with lots of diversity, were drying down but with some flow. This river with incredible mussel history has mussels with exposed posterior surfaces of the shells of the mussels commonly covered with a lush growth of organisms. Vidrine commented on this in his Red River paper: “Freshwater Mussels and Community Structure. --The freshwater mussels in these benthic communities represent the large reef-building (megafauna) of the benthos—many other kinds of organisms live in or on these mussels (Curry et al. 1981, Fuller 1974, Hendrix et al. 1985, Roback et al. 1980, and Vidrine 1996b, c). The communities are complex. The diverse variety of haptobenthic organisms and parasitic/mutualistic organisms includes algae, sponges, hydroids, entoprocts, ectoprocts, leeches, trematodes, chironomids, and mites.” (from Vidrine 2008—Red River). 9. Louisiana. Morehouse Parish. Bayou Bartholomew at U. S. 425 and upstream. This location has a mite community that is apparently extirpated. Like so many streams draining agricultural land farmed in cotton, the mites literally disappear and leave a community of hosts that are not infested with mites. September 26, 1992. Steven George, Charles Allen and his ecology class and M. Vidrine. Host # searched mite # infected

Amblema plicata 8 none


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Arcidens confragosus 7 none

Elliptio dilatata 28 none

Ellipsaria lineolata 1 none

Reginaia ebena 20 none


Lampsilis abrupta 2 none

Lampsilis cardium 2 none

Lampsilis satura 4 none

Lampsilis siliquoidea 1 none

Lampsilis teres 8 none

Leptodea fragilis 4 none

Megalonaias nervosa 8 none

Obliquaria reflexa 3 none

Obovaria olivaria 2 none

Plectomerus dombeyanus 7 none

Pleurobema rubrum 7 none

Potamilus purpuratus 5 none

Ptychobranchus occidentalis 5 none

Theliderma cylindrica 3 none

Theliderma metanevra 33 none

Amphinaias pustulosa 19 none

Quadrula quadrula 10 none

Toxolasmus parvus 3 none

Quadrula verrucosa 9 none

Villosa lienosa 2 none

Campeloma decisum 10 none

Viviparus subpurpureus 10 none

Pleurocera canaliculata 10 none

This sand and gravel bottom river contains the most diverse mussel assemblage remaining in Louisiana. This location had a mite community that was apparently extirpated. Like so many streams draining agricultural land farmed in cotton in north Louisiana and Mississippi, the mites literally disappear and leave a community of hosts that are not infested with mites. The station is provided as an example of the impact of human activity on a freshwater community. The station was originally a ‘fantastic’ fish diversity station as described by Steven George, when he first encountered its rich mussel diversity.

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Vidrine (1995) wrote a report on Bayou Bartholomew, which described the beds found in the Louisiana reaches of the river. A recent study on the Arkansas reaches of this river by Peacock et al. (2013) focused on the Recent and prehistoric mussel fauna. Vidrine (1996f) also wrote a report on Tensas River, which described the mussel beds of this river. The Tensas River stations had been recently deforested. Deforestation along banks left only a thin row of trees, which are commonly blown over thus breaking the bank and leading to massive sedimentation, which literally smothers the mussel beds. Abundant shells of dead mussels and numerous stranded dead and dying mussels along a great length of the river during 1990s study. Few mites were found in that survey—massive cotton farming enterprise caused the deforestation noted above. We do have data from one station in 1970s when there were some mites. The Tensas River has much in common with Bayou Bartholomew and Bayou Pierre, in that they collectively show the impact of agricultural and dredging and deforestation on the mussel and mite communities in rivers in northern Louisiana. All of these rivers historically had huge mussel beds, which appear to have contained diverse mite communities. 10. Arkansas. Montgomery and Polk Counties. Ouachita River at U. S. 270 and upstream to McQuire Public Landing.

August 12, 1978. Bill Bell, Darryl Clark and M. Vidrine. August 15, 1985. Gail, Macky and M. Vidrine.



Alasmidonta marginata 12 U. bishopi 9

U. arcuata 6

Potamilus purpuratus 1 U. australindistincta 1


Pyganodon grandis 4 U. serrata 1

U. wolcotti 1

Lasmigona costata 42 U. dimocki 25

U. clarki 20

U. smithae 20

Villosa iris 7 U. hoesei 7

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U. dimocki 1

Actinonaias ligamentina 112 U. bakeri 15

U. serrata 5

U. clarki 1


Fusconaia flava 46 U. parkeri 27

Strophitus undulatus 8 U. dimocki 2

U. bishopi 2

Lampsilis cardium 9 U. hoesei 9

U. abnormipes 8

U. serrata 2

Obovaria jacksoniana

(formerly Villosa arkansasensis) 18 U. serrata 1

Ptychobranchus occidentalis 50 U. hoesei 50

U. abnormipes 25

U. serrata 25

Cyprogenia aberti 3 U. serrata 1

Theliderma metanevra 2 U. sakantaka 1

U. abnormipes 1

Elliptio dilatata 4 U. parkeri 3

Lampsilis siliquoidea 1 U. fossulata 1

Pleurobema rubrum 2 none


The Ouachita River drains the western portion of the Ouachita Mountains and shares the mussel fauna of these mountains with streams draining the eastern portions, as is evidence in the next station, the Strawberry River. This free-flowing portion of the river above the impoundment had a sand and gravel bottom with areas with large numbers of mussels. The area was routinely canoed by tourists. The stations had very diverse assemblages of mussels and mites, and as such, they represented the best locations in western Arkansas to study mussels and mites. We had the opportunity to canoe this part of the river. Stopping occasionally to search for mussels was so rewarding that the trip made for a high rating as one of the most enjoyable mussel adventures. The river was fairly slow flowing and clear and shallow during our trip, all which made it easy to search for mussels in scattered beds.

11. Arkansas. Sharp County. Strawberry River at U. S. 167, ca. 2 miles north of Evening Shade.

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August 14, 1979. Daniel J. Bereza, C. Vidrine and M. Vidrine. August 13, 1985. Gail, Macky and M. Vidrine. October 14, 1992, June 28, 1993, July 1, 1993 and July 11, 1993. Veryl Board and his students.

Host # searched mite

#

infected

Ptychobranchus occidentalis 14 U. hoesei 4

U. abnormipes 1

Lampsilis streckeri 16 U. hoesei 16

U. abnormipes 16

U. serrata 15

Lampsilis cardium 10 U. hoesei 8

U. abnormipes 5

N. ingens 2


U. belli 1

Lasmigona complanata 1 U. dimocki 1

Theliderma metanevra 3 U. sakantaka 2

U. vikitra 1



U. serrata 3

Lampsilis siliquoidea 3 U. fossulata 2

U. abnormipes 2

Elliptio dilatata 6 U. parkeri 2

U. serrata 2

Potamilus purpuratus 3 U. australindistincta* 3

U. hoesei 1

U. serrata 1


U. laurentiana

deutonymphs 1

Villosa iris 3 U. hoesei 1

Cyprogenia aberti 4 Unionicola larvae 1


U. hoesei 1

U. abnormipes 1

U. australindistincta* 1

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Lampsilis reeviana brevicula 2 U. hoesei 1




Cyclonaias tuberculata 1 U. sakantaka 1

Epioblasma triquetra 2 none

*Unionicola australindistincta in Potamilus purpuratus in this stream differs from the type material in that the genital fields have 7-9 pairs of acetabula similar to U. indistincta; however, the males have a pair of large spines on their tibias of their fourth pair of walking legs—U. indistincta males have only a single large spine.

The Strawberry River drains into the upper White River, a major eastern tributary of the Red River of Arkansas and obviously draining the eastern Ouachita Mountains. The free-flowing sand and gravel-bottomed stream in northeastern Arkansas provided a good example of the diversity of mussels and mites in the eastern part (Ouachita Mountains area) of the Red River in Arkansas. This near natural stream had a highly diverse assemblage. Veryl Board and his students also conducted a survey of this stream, and he shared his mites. Board’s records greatly extended our knowledge of the stream’s fauna, although the numbers in our chart do not represent the mussels examined by Board and his students. This was another remarkable stream with a complex community with diverse interactions between the mites and mussels.

12. Tennessee. Bedford, Coffee, Marshall, Maury and Hickman Counties. Duck River at several stations. July and August 1962. James Lester Wilson.



U. smithae 7


U. tupara 1



U. vikitra 1

U. vamana 1


U. sakantaka 1

Theliderma cylindrica 2 U. sakantaka 2

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U. vamana 5


Lampsilis fasciola 6 U. hoesei 3

Villosa fabalis 16 U. hoesei 13

Actinonaias pectorosa 8 U. hoesei 5


Potamilus alatus 10 U. hoesei 3

U. fulleri 3


U. serrata 1


Alasmidonta marginata 2 none

Lasmigona complanata 5 none

Strophitus undulatus 3 none

Elliptio crassidens 3 none

Lampsilis ovata 1 none

Lampsilis teres 2 none

Before his death, Jim Wilson gave his mussel and mite collection to Vidrine for study. The collection contained large lots from namely the Duck and Stones Rivers before their impoundment and represent very important insight into the nature of streams in central and eastern Tennessee during the mid-20th century. The collections represent highly diverse mussel and mite assemblages. Wilson has sent some of the mites to Rodger Mitchell—these turned out to be new species (Mitchell and Wilson 1965). He had hundreds of vials with mites from hundreds of mussels—a collection never to be repeated. He called one day in 1988 and simply invited my family up to Tennessee for a visit and a gift of a lifetime. We arrived late in the afternoon and enjoyed several beers and a number of tales. I accepted his gift of boxes of labeled vials of mites and promised to figure out what he had discovered. Unfortunately, the effort took nearly 3 years, and Jim passed from lung cancer during the interim—so these mites are his legacy. Sadly, I called him the night that the paper came out—only to find out that he was gone. Jim took great pleasure in describing the beautiful streams and their remarkable mussel beds. His tales of the streams before they were impounded made me want to weep; large dams now obstruct the flow and have long since buried the mussel fauna that was the source of these

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remarkable mites. My experiences in the Clinch and Powell Rivers clearly had prepared me for appreciation of Jim’s tales. I only wish I had had the opportunity to sample with him in the early 1960s—what a pleasure this must have been. 13. Tennessee. Davidson and Rutherford Counties. Stones River at several stations. August 1962. James Lester Wilson.


Pyganodon grandis 1 U. smithae 1


U. smithae 8


U. tupara 33

U. serrata 11



Elliptio dilatata 35 U. serrata 1

U. tupara 2


U. abnormipes 2

U. causeyae 6

U. serrata 2

U. vamana 1

Amphinaias pustulosa 22 U. causeyae 1



Actinonaias ligamentina 16 U. hoesei 12

U. abnormipes 5

U. serrata 3


U. abnormipes 4

U. serrata 8

Lampsilis abrupta/orbiculata 4 U. hoesei 1

/higginsi

U. abnormipes 2

Lampsilis ovata 26 U. hoesei 22

U. abnormipes 6

U. serrata 3


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U. fulleri 4


U. serrata 1

Ligumia recta 6 U. serrata 1

U. abnormipes 1

Villosa taeniata 12 U. hoesei 2

Medionidus conradicus 3 U. hendrixi 3


Ptychobranchus fasciolaris 33 U. hoesei 8

U. causeyae 1

Lampsilis fasciola 4 none

Obovaria subrotunda 1 none



Epioblasma lenior 1 none

Epioblasma interrupta 4 none

Potamilus purpuratus 1 none

Obliquaria reflexa 1 none

Elliptio crassidens 2 none

These stations are part of the same donation by Jim Wilson, an acarologist who was better known for his studies of members of the genus Arrenurus, a wildly colorful and diverse group of free-swimming mites. Vidrine and Wilson (1991) summarized the mite assemblages of Wilson’s collections in the Duck and Stones Rivers. I cannot understate the importance of these collections and Jim’s contribution. There are thousands of mussels in liquid collections from streams around the world—these mussels contain mites preserved for study. The mites are probably not useful for DNA analyses, but they are extremely useful for morphological analyses even though many are preserved in formalin and suffer from breakage during handling and from difficulty in clearing for examination. The fount of information and evidence of associations in these collections could go a long way to answering lots of questions about the evolution of these mites and mussels. Art Bogan has sent me mites from mussels collected and preserved in the 1920s from southern Africa that revealed remarkable new species to science. 14. Tennessee. Hancock County. Clinch River at Kyles Ford. July 8, 1971. Daniel J. Bereza and J. Homziak.

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September 3, 1972. Robie Hensley and Daniel J. Bereza. September 30, 1974. Samuel L. H. Fuller and Daniel J. Bereza. September 21, 1975. Steven Hensley, Daniel J. Bereza and M. Vidrine.


Fusconaia cuneolus 31 U. bogani 10



U. indistincta 12


U. aculeata 1






Leptodea leptodon 1 U. hoesei 1

Margaritifera monodonta 22 none

Epioblasma capsaeformis 69 none

Ptychobranchus subtentum 32 none


Villosa iris 12 none

Alasmidonta marginata 10 none

Lampsilis fasciola 2 none

Ptychobranchus fasciolaris 2 none

Hemistena lata 5 none

Medionidus conradicus 21 none

Ligumia recta 1 none

Epioblasma brevidens 1 none

Amblema plicata 5 none

Dromus dromas 2 none

Plethobasus cyphyus 2 none

Lemiox rimosus 4 none

Amphinaias pustulosa 7 none


Cyprogenia stegaria 6 none

Io fluviatilis 2 none

Campeloma sp. 2 none

At the time of sampling the river was already showing damage as a result of coal mining upstream; however, this region of the river has long been

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heralded as possessing a rich fauna. The stream is fast flowing with many rare species—this rarity may account for the fairly large number of mussels lacking mites.

This reach of Clinch River is very famous for its large variety of mussels. Dan Bereza, Steven Hensley and Robie Hensley sampled the mussel bed several times prior to my visit. The mussels were imbedded in the gravel bottom among boulders and very difficult to dig out on occasion. The stream was very cold and fast when I visited the bed, so collecting was difficult. The water was only 1-2 feet deep and the mussels were fairly common. Many species now rare or endangered were present in these swift waters. Mites were not commonly found, but they were found in the more common mussels, but not in the Margaritifera (Cumberlandia), Epioblasma, Ptychobranchus, Medionidus, Ellliptio, Villosa and Alasmidonta. Really uncommon mussels also did not have mites. All in all, the mussel bed appeared natural to me and very different from those in Louisiana, Mississippi, Texas and Arkansas. Building a campfire on a gravel bar to warm up between intervals in the cold water—now there is an adventure. I was very fortunate to have experience hands working with me. The diversity of this bed revealed itself grudgingly as the mussels were entrenched and scattered into small clusters throughout the river. We spent hours that September day—much too cool for my experienced Louisiana life—working and warming up. New species kept appearing as the work continued, and I would have guessed that if we returned for another day, we would have found another dozen species. But mites were not so common, but I was surprised to see as many as we did in this cold, fast flowing water with mussels buried deep into the rocky substrate—another eye-opening experience. There seemed to be no end to the ability of these mites, and my interest had been tweaked to the maximum. Years later, my dear friend from graduate school, Don Gowan, left Lafayette and ended up working with mussels on the Clinch River. After Don spent several days on excursions to collect mussels with Darryl Clark and me, the surprise that he ended up working on mussel conservation gave me great pride. More than a dozen fellow students from ULL spent several days on excursions with me and made significant contributions to the effort.

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15. Tennessee. Claiborne County. Powell River near Hoop, ca. 10 miles northeast of Tazewell. July 3, 1971. Daniel J. Bereza and J. Homziak. August 16, 1971. Daniel J. Bereza and J. Homziak. September 22, 1975. R. T. Hensley, Steve Hensley, Daniel J. Bereza and M. Vidrine.








Fusconaia cuneolus 27 U. bogani 7

Lasmigona costata 6 none


Plethobasus cyphyus 1 none

Dromas dromus 5 none

Theliderma intermedia 5 none

Ligumia recta 1 none

Lemiox rimosus 1 none

Ptychobranchus fasciolaris 6 none

Ptychobranchus subtentum 4 none

Medionidus conradicus 3 none

Epioblasma triquetra 2 none

Although similar to Clinch River, this river was obviously more heavily impacted by coal mining (with numerous small chunks of coal in the river bottom along with gravel). The mussel community appeared to be ancient, and there was little evidence of recruitment. While a community of mites remained, it was obvious that the mites were far less common, especially in mussels that were rare. Many of these mussels are now considered rare and/or endangered. Taken together, the Clinch and Powell Rivers and the stations of Wilson on the Duck and Stones Rivers provide an excellent window into the historical mussel and mite assemblages of the western Appalachian drainages of Tennessee, Virginia and Kentucky.

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Every station has a story. My visit to this station was on September 22, 1975 with Daniel Bereza and the Hensleys. Dan retold stories of seeing mussels in this stream as a boy when he spent summer months with the Hensleys in their Tennessee home. This is one of the locations that initiated and tweaked his passion for freshwater mussels, and this passion was on fire and freely shared throughout the drive to and from the river. Cutback after cutback was cross in order to get over the mountain to the Powell River. A shallow gravel bar with fast flowing water (fast for me after my experiences in the bayous of Louisiana) with occasional boulders was my first impression. Mussels were common and deeply buried for the most part in gravel making the task very unlike my experiences in the coastal streams. Rake the gravel with your fingers and as the water wash the rakings, mussels appear as if by magic. On occasion a mussel’s siphons or posterior end could be seen as your eye discriminated among the other rocklike entities—many of the mussels were as colorful as the rocks and similarly shaped. But the real story here is one of recruitment. In late September, the cool north breeze shocked me as did the colors breaking out all around—fall in Tennessee felt and looked more like winter in Louisiana. The chill in the water accentuated my sense of season, as it was damn cold. Dan described his youth in the mountains, where he visited the Hensleys for an opportunity to get out of Philadelphia and to get into nature. The Hensleys were not only the most friendly out of state people I had ever met, since they fed me and housed me for a week even roasting chestnuts, albeit Chinese chestnuts, for me in their fireplace. Then they drove us around and jumped into the cold water with us and picked mussels with skills that they appeared to developed over a lifetime of outdoor activity. Dan’s central story revolved around his first trip to Powell River some 15 years earlier as a youth. He had lifted mussels out of the water, and they had as many as 10 juvenile mussels hanging by threads on them. Haag (2012, p. 33) said “The byssus is present in most Lampsilini but is absent in most Margaritiferidae, Amblemini, Anodontini, Pleurobemini, and Quadrulini (Isely 1911; Coker et al. 1921; Smith 2000). The byssus appears within the first 38 days of life and disappears by the end of the first or, rarely, the second year (Coker et al. 1921; Howard 1922). The byssus is retained in the adult stage only in a few small species, including

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Medionidus spp., the fawnsfoot, Truncilla donaciformis, and the rayed bean, Villosa fabalis (Howard 1922; Smith 2000; D. Woolnough, personal communication), for which it evidently remains an effective anchor.” So let’s quickly refresh. Mussels produce larvae called glochidia and release these to infect fish. A myriad of techniques are used to attract fish and infect them. After a brief parasitic life on the gills and fins of fish, the mussels drop off and relocate as juveniles onto the surface of older mussels, sticks and even rocks. Nowhere is this more evident than in a mussel bed, and for Dan’s experience in the Powell River, this was an eye-opener. The juvenile mussel secrete a byssal thread, which adheres to just about any substrate—the thread can be 8-10 inches long—and the mussel is tethered to the line. The mussel juvenile quickly grows a fleshy, muscular foot to hang on to substrate, but if it is dislodged, the tether keeps it from being washed downstream. The juvenile grows quickly and maneuvers itself into a crevice in the substrate where it grows for much of the rest of its life. If disturbed or removed, the mussel can re-enter the substrate and regain a foothold or as occurs on occasion remain loose on the surface. In muddy or sandy substrates, mussels move about to some extent often leaving a visible trail; in rocky substrates, dislodging is life threatening and often fatal. After Dan’s story, I was fully expecting to see juvenile mussels all over the bed, but juveniles were uncommon. Most rocks and mussels had no juveniles. They were almost completely absent. I did pick up one shell with 3 juveniles—picture a 4 inch mussel with 3 small mussels (roughly a centimeter to a half inch each) dangling on strings. These juveniles are similar to their parents obvious even at a remarkably small size, but they are brilliantly colored red, yellow, blue, green, sometimes with ridges, pustules, and other striking surface features. I don’t think anyone could have seen this without holding their breath for a moment—I was not prepared for the impact that it had on my life. Recruitment of juvenile mussels at Powell River was on a severe decline. The immediate suspect was the intense coal mining upstream, which was fouling the water and causing chunks of coal to move downstream and mix with the gravel. We had no way of knowing what kinds of chemicals were being leached out of the numerous pieces of coal evident in the river bed. I remember seeing juveniles in mussel beds in Louisiana, but they were never common.

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Dan lamented at the loss of recruitment since his earlier visits, but he did not seem surprised only disappointed. The impact of this event for me is larger than I imagined. From that day forward, I marveled at each juvenile that I saw dangling from a mussel or stick. Every young mussel of any kind was a celebration of recruitment for me. But they were uncommon, and I never saw even 3 at one time again—I know this because I’ve always counted them and seldom saw more than one at a time. Looking for small mussels to indicate recruitment became a passion, and my fingers grew sensitive as I constantly searched for the smallest mussels, and I routinely found them. But an ever-growing concern developed and disappointment followed, year after year of sampling produced fewer and fewer juvenile mussels, even in streams with well-developed beds. I also looked for mussels that were less than 5 years of age, age is sometimes easy to discern in young mussels as they have obvious growth lines on the shell surface. In many stations, these 5 year old mussels were becoming rare even though they could move about and avoid some fairly extensive sedimentation events. The trend that disappointed Dan at Powell River soon developed into grave concern on my part as the trend became obvious all around. Mussel beds were dying—the young could not survive to adulthood for any one or more of a dozen human generated progressive venues from mining to agriculture to grazing to urbanization not to mention outright point source pollution. I have not returned to Tennessee to see mussels, and I often wonder about the current status of these mussel beds. What about the recruitment of mites? Without being remarkable, it is obvious that the loss of the mussel beds heralds the loss of the mite fauna. Other impacts likewise may readily wipe out the mites. The impacts on fish are for another story.

16. Mississippi. Amite County. East and West Fork Amite River. November 30, 1974. Ed Stern. December 21, 1977. Daniel J. Bereza, Darryl Clark and M. Vidrine. July 25, 1977. Daniel J. Bereza and M. Vidrine. July 2, 1978. Beth and Darryl Clark, Bill Bell and M. Vidrine. November 8, 1981. Darryl Clark and M.Vidrine.


Pleurobema beadleanum 18 U. gowani 1

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U. serrata 6

Elliptio crassidens 88 U. alleni 84

Fusconaia cerina 135 U. parkeri 66

U. serrata 22

N. ingens 1

Amphinaias refulgens 59 U. vikitra 43

U. serrata 8

Lampsilis ornata 4 U. hoesei 4

U. serrata 2

Obovaria unicolor 11 U. guilloryi 3

U. serrata 2

Lampsilis straminea 36 U. hoesei 32

U. abnormipes 23

U. serrata 15

N. ingens 4


U. serrata 15

U. abnormipes 4

Villosa vibex 2 U. hoesei 2

U. abnormipes 2


Amite River drains southwestern Mississippi and contains a very diverse assemblage of mussels and mites. The headwaters are free-flowing, small, sand and gravel bottom streams that contained beds of mussels both in slower flowing areas and in riffle areas. The stations that we examined were in both of the major headwaters called forks. In 1981, Darryl Clark and I studied a bed with lots of Elliptio crassidens in the west fork. Vidrine and Clark (1983) characterized the bed as follows: “West Amite River is a medium-sized stream with a gravel and sand bottom. We sampled a 105 m2 area in a small rivulet connecting two branches in the winding river approximately 1.5 miles north of the Louisiana border. The rivulet varied between three and five meters wide and had a mixed mud, sand and detritus substrate. The mussel bed was not well formed, since mussels were spread about the area collected. We analyzed the area in order to characterize the species composition and relative abundance of the mussels present. Of the 610 mussels encountered, the species and their percentage of the total were: Elliptio crassidens

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(Lamarck 1819) 56.6%, Villosa lienosa (Conrad 1834) 16.6%, Lampsilis claibornensis (Lea 1838) 12.0%, Fusconaia flava group (Rafinesque 1820) 7.9%, Quadrula pustulosa group (Lea 1831) 1.5%, Elliptio beadliana (Lea 1861) 1.5%, Tritogonia verrucosa (Raf. 1820) 1.2%, Obovaria jacksoniana Frierson 1812 1.0%, Obovaria unicolor (Lea 1845) 1.0%, Villosa vibex (Conrad 1834) 0.5%, and Lampsilis excavata Lea 1857 0.2%.” Many names changed for the mussels; thus, a list is provided here:

Name name changes percent Elliptio crassidens Elliptio crassidens 56.6% Villosa lienosa Villosa lienosa 16.6% Lampsilis claibornensis Lampsilis straminea 12.0% Fusconaia flava Fusconaia cerina 7.9% Quadrula pustulosa Amphinaias refulgens 1.5% Elliptio beadliana Pleurobema beadleanum 1.5% Tritogonia verrucosa Quadrula verrucosa 1.2% Obovaria jacksoniana Obovaria jacksoniana 1.0% Obovaria unicolor Obovaria unicolor 1.0% Villosa vibex Villosa vibex 0.5% Lampsilis excavata Lampsilis ornata 0.2% We also studied a bed in east fork in 3 inches of water in a gravel bar with Obovaria unicolor. It was amazing to see mussels thick as thieves in clear fast moving water just 3 inches deep.

17. Louisiana. St. Helena Parish. Twelve Mile Creek (Tangipahoa River drainage) at LA 1045. December 19, 1976. Daniel J. Bereza and M. Vidrine. December 21, 1977. Daniel J. Bereza and M. Vidrine. April 12, 2006. M.Vidrine. May 23, 2006. M. Vidrine. June 7, 2007. John Hamlin, Lindsey Thomas, Dustin Joubert, Benjamin Dubansky and M. Vidrine.



U. abnormipes 19

U. serrata 1


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U. abnormipes 8

U. serrata 7


Pleurobema beadleanum 59 U. gowani 36

U. aculeata 1

N. ingens 5

Fusconaia cerina 3 U. parkeri 3

Anodontoides radiatus 13 U. arcuata 10

U. dimocki 1

U. ernstingi 8

Strophitus subvexus 7 U. arcuata 6

U. dimocki 2

U. tumida 6

This small, sandy creek contained a diverse and apparently unique assemblage. It provided us with a clue as to the hidden potential in the creeks of the northern Gulf drainages. This is a creek that provided mites for analyses in several of the papers on DNA analyses by Ernsting and Edwards including the 2014 paper on cryptic species as well as the source of U. ernstingi. The diversity of mussels and mites and their associations clearly demonstrates how communities evolve in each stream. Although this stream is only a few feet deep and slow flowing and seemingly deep in the woods, it is a window into the potential diversity in our streams. This location was one of the first creeks that Dan Bereza collected with me in Louisiana. He was amazed by the mussels and the ease of collecting in the dead of winter in 1976. I think he was as amazed as I was in September in Tennessee in the Clinch River for just the opposite reasons. 18. Florida. Holmes and Washington Counties. Holmes Creek (Choctawhatchee River drainage) at U. S. 90.

September 1965. Robert Dobson. June 13, 1975. M. Vidrine.

July 7, 1977. M. Vidrine.


Pyganodon gibbosa 9 U. formosa 1

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U. wolcotti 4

U. serrata 3


U. serrata 25


U. serrata 8

U. sakantaka 1

Elliptio pullata/arctata 22 U. alleni 10

U. causeyae 2

Fusconaia burkei 21 U. gowani 14

Utterbackia peggyae 17 U. foili 16

U. wolcotti 3

U. serrata 3

Toxolasma paulum 22 U. latipalpa 21

U. vamana 1

U. sakantaka 1

U. formosa 1

Campeloma geniculum 40 U. dobsoni 25

Pomacea paludosa 5 none

This small, sandy creek with a variety of snails and mussels was free-flowing and apparently near natural in character. The station had been studied by Robert Dobson in the 1960s. It was a good source of information on the mites and mussels in creeks in the eastern Florida panhandle. Here is another creek with huge diversity, including large snails, Pomacea paludosa, with amazing pink egg sacks posted on the pilings of the bridge. This was my first encounter with U. (Berezatax) latipalpa, a fascinatingly different mite from any that I had ever seen. Robert Dobson had found this mite and used the name ‘latipalpa’ as an informal, manuscript name that marvelously described the mites’ pedipalps as ‘dorsoventrally flattened’—a characteristic typical of the South American mites belonging to the then genus Atacella. The same mite was later found in Texas in another species of Toxolasma.

19. Florida. Gadsden and Leon Counties. Ochlockonee River at FL 263, U. S. 27 and U. S. 90. September 1965. Robert Dobson. July 17, 1977. Daniel J. Bereza, Marc Imlay and M. Vidrine.


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Lampsilis floridensis 50 U. hoesei 35

U. abnormipes 5

U. vamana 1

U. sakantaka 1

Villosa lienosa 49 U. tupara 1


Elliptoideus sloatianus 69 U. tupara 1

Amphinaias infucata 170 U. sakantaka 170

U. vamana 4

Elliptio pullata 61 U. tupara 1


U. abnormipes 56

Toxolasma paulum 18 U. latipalpa 14

Utterbackia peggyae 11 U. hoesei 2

Villosa amygdala 16 U. poundsi 6

Uniomerus carolinianus 1 none

Pleurobema pyriforme 8 none

Medionidus simpsonianus 22 none

Hamiota subangulata 24 none

This large, shallow river had a sand bottom and a tremendous number of mussels. The station provided a good look at the diversity of mussels and mites in rivers of the eastern Florida panhandle. Dan, Marc and I spent a long afternoon sampling this river and marveling over the tremendous diversity in panhandle Florida. Many of the mussels ended up in large samples with their flesh packed in dry ice to be used by Davis and Fuller (1981) in developing their phylogenetic analyses of the North American mussels.

20. Florida. Sumpter and Hernando Counties. Little Withlacooche River at U. S. 301. July 8, 1977. M. Vidrine. March 29, 1980. Thomas Dale Bishop, Karen Dupont, Darryl Clark, Mark LaSalle and M. Vidrine.


Elliptio jayensis 20 U. aculeata 2

U. abnormipes 2

Elliptio icterina/occulta 15 U. alleni 3

Lampsilis floridensis 11 U. hoesei 10

U. abnormipes 1

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U. aculeata 2


U. abnormipes 5

Utterbackia peggyae 1 U. poundsi 1

Uniomerus carolinianus 21 U. lasallei 14


Pomacea paludosa 2 none

This small river in peninsula Florida’s western drainages hosted a rich mussel population with a diverse mite assemblage. It was a slow flowing, sandy stream that provided a good introduction to the mussels and mites of the peninsula. My classmates and I sampled this stream in order to understand how very small streams have highly diverse faunas—I can say that I was highly impressed not only with the mussels but also with the mites. In the 1980s, Barbara Downes would find problems with my discerning U. poundsi from Villosa amygdala and U. lasallei from Uniomerus carolinianus and deduce host-induced morphology as part of her ongoing dissertation problem. In the early 1990s, Dale Edwards, Ron Dimock, Jim Cordes and I would return to panhandle Florida and resample her stations in order to analyze these mites in laboratory studies. Edwards and Labhart (1998) would later clarify this conundrum and clearly verify the species of mites as biochemically different and thus not a result of host-induced morphology as proposed.

21. Florida. Okeechobee and Highlands Counties. Kissimmee River at U. S. 98. July 9, 1977. M. Vidrine.


Elliptio jayensis 66 U. alleni 54

U. causeyae 50

U. abnormipes 5


U. abnormipes 4

N. ingens 4

Toxolasma paulum 1 none

This sandy river drains into the Lake Okeechobee and into the Everglades. While its mussel diversity appeared limited, the mussels were present in astronomical numbers; however, the individual mussels were small in size. The station provided a great example of the diversity of mussels and mites

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in lower peninsular Florida. As I crawled into this river, I could feel hundreds of mussels embedded in the banks. I could easily pick 5 mussels out of the substrate with each hand. I could not imagine the potential thousands of small mussels in this river. After an hour of searching, I was so overwhelmed by the sheer number of mussels and their dizzying but subtle differences that I packed up and moved on to Indian Prairie Canal that encircles Lake Okeechobee, only to find a similar situation.

22. Canada. Ontario. Dufferin County. Amaranth Twp. Willow Brook 2.2 miles east-northeast of Grand Valley. August 5, 1971. B. T. Kidd.


Anodontoides ferussacianus 4 U. arcuata 1

Strophitus undulatus 6 U. dimocki 3

U. arcuata 3

Lasmigona compressa 12 U. arcuata 12

U. dimocki 12

U. neocooki 12

During the winter of 1982, I spend several days in the Canadian National Collections under the leadership of Muriel Smith, the curator of the mussel collections. I examined lots of mussels from across the Canadian provinces. This is an example of what I found in containers of mussels from a single station in Ontario. Canadian provinces are rich in mussels belonging to the Anodontini and rich in mites that typically parasitize them. My great surprise came in containers from western Canada and the northwestern United States (Pacific drainages) in the next station. 23. Canada. British Columbia. Vereux Lake 8.5 miles north of Oliver. August 6, 1972. B. T. Kidd.


Anodonta kennerlyi 6 U. conroyi 2

Anodonta nuttalliana 15 U. conroyi 12

Gonidea angulata 4 none

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During my examination of mussel lots from the Canadian National Collections, mussels from British Columbia had mites that changed my way of thinking. Dan Bereza and I had spent the previous winter in Mexico and the previous weeks at the Smithsonian Institution and University of Michigan Museum of Natural Sciences examining mussels for mites. I had some great finds that for the most part agreed with what I had found in my dissertation in 1980. But these collections from western Canada and Washington contained a mite species that I had seen only as a single male specimen that I considered as incidental from Sam Fuller’s collection of Anodonta wahlamatensis in northern California (Unionicola sp. nov. type 4 in Vidrine 1980a). These mites, U. conroyi, broke all of the rules in that it was a member of Unionicolides, a group of mites that were found in the Ambleminae in the United States and not in the Unioninae (Anodontini). In the Michigan collections, I found a close relative of U. conroyi in Anodonta dejecta from western Mexico—so these mites were in Anodonta from Mexico to Canada in the Pacific drainages. I returned after my work in Canada to Philadelphia for a final visit with Dan. I returned to Louisiana to find new love and a new life, which took me away from musseling and into prairies and the classroom, and then I focused on writing about all that I had found and seen.

24. Texas. Brazos, Grimes, Madison, Robertson, Leon and Limestone Counties. Navasota River at 26 stations. May 1973-1975. Thomas Robert Calnan. December 29, 1977. Alan Neumann, Daniel J. Bereza and M. Vidrine.



U. abnormipes 1



U. aculeata 1


U. aculeata 1

Arcidens confragosus 5 U. abnormipes 2

Toxolasma texasiensis 69 U. latipalpa 26

U. abnormipes 1

U. aculeata 1

Cyrtonaias tampicoensis 7 U. hoesei 3

U. abnormipes 2

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U. abnormipes 42

U. aculeata 2


U. hoesei 1





U. tupara 3


Uniomerus tetralasmus 32 U. stricta 32

Villosa lienosa 37 U. hoesei 1

U. abnormipes 18

Leptodea fragilis 19 none

This medium-sized stream with mud, sand and gravel bottoms had a diverse assemblage of mussels and mites. Thomas Robert Calnan studied it extensively for his thesis in the early 1970s. The river appeared near natural, but mussels were not as common in the search in the late 1970s. This river later would seem extraordinary in that the population of Cyrtonaias had U. hoesei and not the typical mites that I found in this genus, U. calnani—a species that I would name for Thomas Robert Calnan. The stream also had a good population of Toxolasma texasiensis with U. (Berezatax) latipalpa.

25. Mexico. Tamaulipas. Small river (locally called Arroyo los Gatos and presumably a headwater of the Casas Viajas River system) at MX 80 in Nuevo Morelos. November 5 and 13, 1978. Daniel J. Bereza, R. Herschler and C. Dunn.


Anodontites trapesialis 1 U. entrerrianensis 1

U. neoperforata 1

Friersonia iridella 35 U. hoesei 20

U. serrata 5

Popenaias metallica 29 U. acylindrotarsa 17

U. laurentiana 16

N. ingens 1

Disconaias fimbriata 12 U. hoesei 2

U. serrata 2

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U. aculeata 1

Actinonaias signata 17 U. berezai 1

U. aculeata 1

U. laurentiana 1

In 1982, Daniel Bereza, Robie Hensley, Steve Hensley and I sampled nearly 40 stations in eastern drainages of Mexico from the Rio Grande south to Lake Catemaco in Veracruz. This station is typical of what was found and served as the impetus for the 1982 trip. Many of the mussels from the 1982 remained unidentified with Dan’s untimely death—the mussels are currently housed in the collections of The Academy of Natural Sciences of Philadelphia and in the U. S. National Collections at the Smithsonian Institution. Mexican streams varied from small clear creeks to large rivers—each with dizzying diversity. Dan was trying to relax as many of these mussels as possible for later study and determination, but we sacrificed representative samples in order to obtain mites. I certainly expected a more dizzying array of mites, but a few exciting species kept turning up, namely U. berezai, U. acylindrotarsa and U. hensleyi. We found good populations of species that I had seen in the United States, but we were impressed with the species of Atacella that we were finding in Anodontites trapesialis, our müllerid mussel in North America. January and February in streams in Mexico proved to be very impressive. Mussel beds were scattered but impressive. Large species, Disconaias disca and Cyrtonaias spp., were most impressive, while small species, Popenaias spp, Friersonia spp., Actinonaias spp. and an array of unknowns were scattered about in beds and singly here and there. The most becoming memory of the trip was the impressive collecting permit from the central government—a large embossed and wax-seal signed proclamation—it got us into places that we would not have gotten into and out of places that we have not gotten out of on a routine basis. Many of the mussels were preserved after relaxation with their mites in situ—I did not have the chance to examine them in all these years, and I suspect more mysteries remain hidden among these mussels that were Dan’s pride and joy.

We want to encourage students to: 1. conduct surveys, especially in mussel beds, 2. search preserved mussels for new species and records of

associations, and 3. conduct morphological and molecular analyses of both the mussels

and the mites in an effort to clarify aspects of our current knowledge.

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We hope these stories serve as an impetus for future work. Collectively, these stations should give the readers an overview of the events on my path to understanding these mites and their hosts. Every mussel bed has/had an amazing story to tell. These beds helped to reveal the intricate and diverse associations between mussels and mites that I would come to see as ‘glue’ connecting all of the aquatic fauna to one another. These beds were filled with fish, insects and an endless array of associations—and I was focused on sampling just one tier of these associations. If we cannot begin to understand this diversity, we will never take the first step to preserve it and treasure it. It will all be lost! For the purpose of creating hypotheses and reflecting on the diversity of mussel beds both present and historical, Edwards and Vidrine (2013c) attempted a mental exercise in paleoecology as an endnote. We are adding this endnote as a closing to this chapter. Chapter endnote from Edwards and Vidrine 2013c (paleoecology) “Closing the chapter with a mental exercise might best provide a view of the possibilities nested within our data. The exercise involves forensically reconstructing decimated communities of mussels and mites, with either the mussels or the mites or both largely eliminated. Two good examples worthy of this kind of thought experiment are Bayou Bartholomew, an arm of the Ouachita River in northeastern Louisiana, and Muscle Shoals, the largest mussel bed ever known, in the Tennessee River in Alabama. Both of these mussel beds have been described, and lists of the mussels known to historically occur in these beds are known (see George and Vidrine 1993 and Haag 2012, respectively). Based upon what we know regarding how mussel and mite assemblages are constructed (exemplified in Appendix 5) and what we can infer based upon known host-specificities of the mites and guild structures of mite communities within mussels (information summarized in Chapters 6 and 7), we can reconstruct a hypothetical assemblage for each of these beds by integrating this information. For example, the list of mussels from Bayou Bartholomew can be correlated with the lists of mussels and their mites in other stations in the Ouachita River system in Louisiana and neighboring Arkansas. Records from other drainages, e. g., Strawberry River in Arkansas

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and southern drainages in Louisiana, can be used to tweak the ultimate list of species of mites in the assemblage and their prospective associations. The resultant hypothetical assemblage for the mussel bed would represent our best estimate of the historical structure of the community. Similarly, the list of mussels from the mussel bed in Muscle Shoals can be correlated with lists of known mussel and mite assemblages from the Clinch River, Powell River, Duck River, Stones River and upper Mississippi River drainages. Again the resultant hypothetical assemblage would have some gaps, but it would readily serve as a working model in discussions regarding habitat evaluation and the historical/potential diversity of these habitats. While we do not wish to delve too heavily in such postmortem reconstructions, we are aware that understanding the past and endeavoring to reconstruct the past with specific evidence reveals previous environmental structure and biological diversity and that these are potentially rewarding efforts. Unfortunately, in many parts of the world, too little is known about these assemblages to even risk venturing a guess regarding historical assemblages.”

chapter 5dilapidated bridge locations. and suddenly, i knew the answer. bridges and roads are built...

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