Quadrula infucata - (Conrad, 1834)
Sculptured Pigtoe
Taxonomic Status: Accepted
Related ITIS Name(s): Quincuncina infucata (Conrad, 1834) (TSN 80356)
Unique Identifier: ELEMENT_GLOBAL.2.117994
Element Code: IMBIV40020
Informal Taxonomy: Animals, Invertebrates - Mollusks - Freshwater Mussels
 
Kingdom Phylum Class Order Family Genus
Animalia Mollusca Bivalvia Unionoida Unionidae Quadrula
Genus Size: D - Medium to large genus (21+ species)
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Concept Reference
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Concept Reference: Turgeon, D.D., J.F. Quinn, Jr., A.E. Bogan, E.V. Coan, F.G. Hochberg, W.G. Lyons, P.M. Mikkelsen, R.J. Neves, C.F.E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F.G. Thompson, M. Vecchione, and J.D. Williams. 1998. Common and scientific names of aquatic invertebrates from the United States and Canada: Mollusks. 2nd Edition. American Fisheries Society Special Publication 26, Bethesda, Maryland: 526 pp.
Concept Reference Code: B98TUR01EHUS
Name Used in Concept Reference: Quincuncina infucata
Taxonomic Comments: Ortmann and Walker (1922) originally placed burkei and infucata (with mitchelli added later) in the genus Quincuncina based on a zig-zag shell surface sculpture. This shell pattern is found on several other unionid species, however, and is not homologous and does not reflect common ancestry (Lydeard et al., 2000). The shell varies as to the degree of sculpture, with eastern populations being highly sculptured (Suwannee and Ochlockonee river systems) and western populations being nearly or completely devoid of sculpture (Chipola river system). The latter populations begin to resemble some Fusconaia species, particularly some Fusconaia succissa. Recent mitochondrial DNA evidence suggests this species is closely related to, and may possibly require placement in, the genus Quadrula (Serb et al., 2003). Simpson (1900; 1914) placed infucata in the genus Quadrula. Ortmann and Walker (1922) suggested the species belonged in the genus Quincuncina, where it remained until Serb et al. (2003) reported it belongs to Quadrula, based on genetic analyses.
Conservation Status
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NatureServe Status

Global Status: G3
Global Status Last Reviewed: 23Dec2013
Global Status Last Changed: 13Jul2005
Rounded Global Status: G3 - Vulnerable
Reasons: Despite stability or local expansion in one river system (Chipola), population numbers and occupied habitat are beginning to decline in many areas, and many historical occurrences are now extirpated. The species may be on the verge of becoming threatened.
Nation: United States
National Status: N3 (13Jul2005)

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Alabama (S1), Florida (S2S3), Georgia (S3)

Other Statuses

IUCN Red List Category: NT - Near threatened
American Fisheries Society Status: Special Concern (01Jan1993)

NatureServe Global Conservation Status Factors

Range Extent: 1000-5000 square km (about 400-2000 square miles)
Range Extent Comments: The species is endemic to the Apalachicola (including the Apalachicola, Chipola, Chattahoochee, and Flint rivers) and Ochlockonee river systems within the Apalachicolan region of Florida, Alabama, and Georgia. It occurs in mainstem sites as well as several large and some small tributary streams and is among the widest ranging Apalachicolan region endemic mussel species (Clench and Turner, 1956; Brim Box and Williams, 2000; Williams, 2014).

Area of Occupancy: 501-12,500 4-km2 grid cells
Area of Occupancy Comments: Linear occupancy is 200-5000 km.

Number of Occurrences: 1 - 20
Number of Occurrences Comments: Potentially, each unfragmented stretch of river, regardless of number of collection sites, represents a single element occurrence. The species is known from dozens of localities in the Chattahoochee (Georgia, Alabama), Flint (Georgia), Apalachicola (Florida), Chipola (mostly Florida- see van der Schalie, 1940), and Ochlockonee (Florida, Georgia) river drainages. However, most of the Alabama and Georgia records are historic. The only known extant populations in Alabama are in Uchee and Little Uchee creeks, Russell and Lee counties (Williams et al., 2008). In 1999 and 2001, the species was found in 14 sites (360 specimens) and 11 sites (543 specimens), respectively, in surveys of 21 sites (each year) in about a dozen tributary streams of the lower Flint River basin, southwestern Georgia (Golladay et al., 2004). Specimens were also found in a survey of six sites in the Elmodel Wildlife Management Area, Ichawaynochaway Creek, southwestern Georgia (Battle et al., 2003). In the Apalachicola-Chattahoochee-Flint (ACF) basin, it was collected relatively recently at 54 of 324 sites in Alabama, Florida, and Georgia in the mainstem and tributaries of the Flint and Apalachicola Rivers, the mainstem of the Chipola River, and three Chattahoochee River tributaries but not the mainstem (Brim-Box and Williams, 2000).

Population Size: 2500 - 100,000 individuals
Population Size Comments: At the GEXEMPSITE, 68 were found in an hour's time in 1988. Although the species was considered "rather rare" by Clench and Turner (1956), it is sometimes abundant in portions of the Ochlockonee River, Chattahoochee River tributaries, and the Alapaha River mainstem in both Florida and Georgia; populations in several other streams are smaller. Only 7 specimens were collected in a Chipola River survey conducted during World War I, but now the species is common on the lower mainstem above the former Dead Lake area. It is undoubtedy the most common currently recognized Apalachicolan region endemic mussel. In 1999, this species was found to comprise 3.01% (relative abundance) of the 14,873 mussels collected in surveys of 46 sites in 12 tributary streams of the lower Flint River basin, Georgia (Gagnon et al., 2006). In 1999 and 2001, it was found at 14 sites (360 specimens) and 11 sites (543 specimens), respectively, in surveys of 21 sites (each year) in about a dozen tributary streams of the lower Flint River basin, southwestern Georgia (Golladay et al., 2004). In the ACF basin, it was collected relatively recently from 54 of 324 sites (235 live, 122 shells) in Alabama, Florida, and Georgia in the mainstem and tributaries of the Flint and Apalachicola Rivers, the mainstem of the Chipola River, and three Chattahoochee River tributaries but not the mainstem (Brim-Box and Williams, 2000).

Number of Occurrences with Good Viability/Integrity: None to few (0-12)

Overall Threat Impact: High
Overall Threat Impact Comments: Siltation from poorly conducted agricultural and silvicultural activities, localized gravel/sand mining (upper Apalachicola system); localized industrial, municipal pollution; and channel maintenance dredging in the Apalachicola drainage have impacted populations. Additional threats such as watershed development, urban sprawl in greater Atlanta area (upper Apalachicola), and competition from Asian clams potentially aided in the eradication of this once super-abundant species from the Apalachicola River mainstem below Jim Woodruff Dam.

Short-term Trend: Decline of 10-30%
Short-term Trend Comments: Populations appear to be stable to expanding in the lower Chipola River and are still present in some tributaries of the Chattahoochee River, but they are declining or extirpated in certain other systems or mainstems when compared to historic information. The species is very likely extirpated from some sites on the Apalachicola (Brim Box and Williams, 2000), Chattahoochee, and Withlacochee (Suwannee) mainstems, and from the Little River (Ochlockonee) (Golladay et al., 2004). It is also extirpated from Big Creek, Houston Co., Alabama (Mirarchi et al., 2004). The timeframe of declines is uncertain.

Long-term Trend: Decline of <50% to increase of <25%
Long-term Trend Comments: This species appears to have been relatively common prehistorically and historically. Heard (1964) reported it in considerable numbers in the Apalachicola River at Chattahoochee, Florida, but subsequently (Heard, 1975) noted drastic reductions in the river following a heavy invasion of Corbicula fluminea. It is very likely extirpated from sites on the Apalachicola, Chattahoochee, and Withlacochee (Suwannee) mainstems and from the Little River (Ochlockonee) (Golladay et al., 2004). It is also extirpated from Big Creek, Houston Co., Alabama (Mirarchi et al., 2004). The timeframe of declines is uncertain.

Intrinsic Vulnerability: Unknown
Intrinsic Vulnerability Comments: Freshwater mussels are inherently vulnerable to threats from siltation, pollution, eutrophication, channelization, impoundment, collection, drought and water withdrawal, competiton from invasive non-native mussels, and changes to larval host fish populations.

Environmental Specificity: Moderate. Generalist or community with some key requirements scarce.
Environmental Specificity Comments: This species may be somewhat more tolerant of habitat modifications than most regionally endemic and channel-dwelling species. Nonetheless, it has the typical liabilities of filter-feeders (e.g., to siltation, pollution, and eutrophication).

Other NatureServe Conservation Status Information

Inventory Needs: Determine status of extant populations. Resurvey historic sites and search for new ones by conducting intensive surveys in unsampled portions of the Flint, Chattahoochee, and Ochlockonee rivers.

Protection Needs: Maintain high water and benthic habitat (substrate) qualities, as well as adequate flow regimes, throughout all occupied river systems. This may be partially accomplished via establishment of buffers and streamside management zones for all agricultural, silvicultural, mining, and developmental activities; protection of floodplain forests and adjoining upland habitat is paramount. Best management practices to follow include employing forestry practices that cause minimal soil erosion; preventing access of livestock to natural surface waters and drains; situating roads at least 0.25 mi. (0.4 km) from heads of all tributaries, even more on steep slopes; using silt fencing and vegetation to control runoff and siltation at all stream crossings, especially during construction and maintenance; using and maintaining sewer systems rather than septic tanks and stream-dumping for management of wastewater; and avoiding use of agricultural pesticides on porous soils near streams. Prevent damming, dredging, and pollution throughout drainages, but especially near recorded sites. Remove existing dams, but with great care to limit downstream sedimentation. Limit withdrawal of surface and subterranean waters as necessary to maintain normal stream flows, especially during drought. Prevent or limit establishment of invasive species (including zebra mussel, Dreissena polymorpha) to the extent possible. Where appropriate, protect populations through acquisitions and easements over streamside lands by working with government agencies and conservation organizations.

Distribution
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Global Range: (1000-5000 square km (about 400-2000 square miles)) The species is endemic to the Apalachicola (including the Apalachicola, Chipola, Chattahoochee, and Flint rivers) and Ochlockonee river systems within the Apalachicolan region of Florida, Alabama, and Georgia. It occurs in mainstem sites as well as several large and some small tributary streams and is among the widest ranging Apalachicolan region endemic mussel species (Clench and Turner, 1956; Brim Box and Williams, 2000; Williams, 2014).

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map
Endemism: endemic to a single nation

U.S. & Canada State/Province Distribution
United States AL, FL, GA

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
GA Baker (13007), Calhoun (13037), Clay (13061)*, Clayton (13063), Cobb (13067)*, Coweta (13077), Crawford (13079), Crisp (13081), Decatur (13087), Dooly (13093), Dougherty (13095), Early (13099), Fayette (13113), Fulton (13121)*, Grady (13131), Harris (13145)*, Heard (13149)*, Lee (13177), Macon (13193), Marion (13197)*, Meriwether (13199), Miller (13201), Mitchell (13205), Muscogee (13215)*, Peach (13225), Pike (13231), Randolph (13243), Spalding (13255), Sumter (13261), Talbot (13263), Taylor (13269), Terrell (13273), Thomas (13275), Troup (13285)*, Webster (13307), Worth (13321)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
03 Upper Suwannee (03110201), withlacoochee (03110203)+, Lower Suwannee (03110205), Upper Ochlockonee (03120002)+, Lower Ochlockonee (03120003)+, Upper Chattahoochee (03130001)+, Middle Chattahoochee-Lake Harding (03130002)+, Middle Chattahoochee-Walter F. George Reservoir (03130003)+, Lower Chattahoochee (03130004)+, Upper Flint (03130005)+, Middle Flint (03130006)+, Kinchafoonee-Muckalee (03130007)+, Lower Flint (03130008)+, Ichawaynochaway (03130009)+, Spring (03130010)+, Apalachicola (03130011)+, Chipola (03130012)
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
U.S. Distribution by Watershed (based on multiple information sources) Help
Ecology & Life History
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Basic Description: A small, subcircular, inflated, generally sculptured freshwater mussel.
General Description: See Clench and Turner (1956).
Diagnostic Characteristics: Small, subcircular/subquadrate outline, with heavy to nearly obliterated sculpture, heavy-shelled, with heavy teeth.
Reproduction Comments: Reproductive strategies and glochidial hosts are unknown, but it is possibly tachytictic (short-term brooder), as are members of the somewhat closely related genus Fusconaia.
Habitat Type: Freshwater
Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: N
Mobility and Migration Comments: Adults are essentially sessile. About the only voluntary movement they make is to burrow deeper into the substrate although some passive movement downstream may occur during high flows. Dispersal occurs while the glochidia are encysted on their host (probably a fish).
Riverine Habitat(s): BIG RIVER, CREEK, Low gradient, MEDIUM RIVER, Moderate gradient, Riffle
Special Habitat Factors: Benthic
Habitat Comments: This species is found "In muddy sand and sand in moderate current" (Heard, 1979). Generally it is found in shallow, sand bottomed river pools and rocky areas with swift current including channels with stable substrates of medium-sized creeks to large rivers. In the ACF basin, 45% of recently collected specimens were found at sites that contained a sand and limestone substrate, with 24% at sites with a predominantly sand and detritus substrate (Brim Box and Williams, 2000).
Adult Food Habits: Detritivore
Immature Food Habits: Parasitic
Food Comments: Presumably fine particulate organic matter, primarily detritus, and/or zooplankton, and/or phytoplankton (Fuller, 1974). Larvae (glochidia) of freshwater mussels generally are parasitic on fish and there may be a specificity among some species.
Length: 5 centimeters
Economic Attributes Not yet assessed
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Management Summary
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Biological Research Needs: Determine life history (including fish host species), reproductive biology, fecundity, sizes and viabilities of extant populations, and microhabitat requirements, as well as sensitivity to silt, excessive nutrients, and pollutants. Examine competitive interactions with Corbicula.
Population/Occurrence Delineation
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Group Name: Freshwater Mussels

Use Class: Not applicable
Minimum Criteria for an Occurrence: Occurrences are based on some evidence of historical or current presence of single or multiple specimens, including live specimens or recently dead shells (i.e., soft tissue still attached and/or nacre still glossy and iridescent without signs of external weathering or staining), at a given location with potentially recurring existence. Weathered shells constitute a historic occurrence. Evidence is derived from reliable published observation or collection data; unpublished, though documented (i.e. government or agency reports, web sites, etc.) observation or collection data; or museum specimen information.
Mapping Guidance: Based on the separation distances outlined herein, for freshwater mussels in STANDING WATER (or backwater areas of flowing water such as oxbows and sloughs), all standing water bodies with either (1) greater than 2 km linear distance of unsuitable habitat between (i.e. lotic connections), or (2) more than 10 km of apparently unoccupied though suitable habitat (including lentic shoreline, linear distance across water bodies, and lentic water bodies with proper lotic connections), are considered separate element occurrences. Only the largest standing water bodies (with 20 km linear shoreline or greater) may have greater than one element occurrence within each. Multiple collection or observation locations in one lake, for example, would only constitute multiple occurrences in the largest lakes, and only then if there was some likelihood that unsurveyed areas between collections did not contain the element.

For freshwater mussels in FLOWING WATER conditions, occurrences are separated by a distance of more than 2 stream km of unsuitable habitat, or a distance of more than 10 stream km of apparently unoccupied though suitable habitat. Standing water between occurrences is considered suitable habitat when calculating separation distance for flowing water mussel species unless dispersal barriers (see Separation Barriers) are in place.

Several mussel species in North America occur in both standing and flowing water (see Specs Notes). Calculation of separation distance and determination of separation barriers for these taxa should take into account the environment in which the element was collected. Juvenile mussels do not follow this pattern and juveniles are typically missed by most standard sampling methods (Hastie and Cosgrove, 2002; Neves and Widlak, 1987), therefore juvenile movement is not considered when calculating separation distance.

Separation Barriers: Separation barriers within standing water bodies are based solely on separation distance (see Separation Distance-suitable, below). Separation barriers between standing water bodies and within flowing water systems include lack of lotic connections, natural barriers such as upland habitat, absence of appropriate species specific fish hosts, water depth greater than 10 meters (Cvancara, 1972; Moyle and Bacon, 1969) or anthropogenic barriers to water flow such as dams or other impoundments and high waterfalls.
Separation Distance for Unsuitable Habitat: 2 km
Separation Distance for Suitable Habitat: 10 km
Alternate Separation Procedure: None
Separation Justification: Adult freshwater mussels are largely sedentary spending their entire lives very near to the place where they first successfully settled (Coker et al., 1921; Watters, 1992). Strayer (1999) demonstrated in field trials that mussels in streams occur chiefly in flow refuges, or relatively stable areas that displayed little movement of particles during flood events. Flow refuges conceivably allow relatively immobile mussels to remain in the same general location throughout their entire lives. Movement occurs with the impetus of some stimulus (nearby water disturbance, physical removal from the water such as during collection, exposure conditions during low water, seasonal temperature change or associated diurnal cycles) and during spawning. Movement is confined to either vertical movement burrowing deeper into sediments though rarely completely beneath the surface, or horizontal movement in a distinct path often away from the area of stimulus. Vertical movement is generally seasonal with rapid descent into the sediment in autumn and gradual reappearance at the surface during spring (Amyot and Downing, 1991; 1997). Horizontal movement is generally on the order of a few meters at most and is associated with day length and during times of spawning (Amyot and Downing, 1997). Such locomotion plays little, if any, part in the distribution of freshwater mussels as these limited movements are not dispersal mechanisms. Dispersal patterns are largely speculative but have been attributed to stream size and surface geology (Strayer, 1983; Strayer and Ralley, 1993; van der Schalie, 1938), utilization of flow refuges during flood stages (Strayer, 1999), and patterns of host fish distribution during spawning periods (Haag and Warren, 1998; Watters, 1992). Lee and DeAngelis (1997) modeled the dispersal of freshwater into unoccupied habitats as a traveling wave front with a velocity ranging from 0.87 to 2.47 km/year (depending on mussel life span) with increase in glochidial attachment rate to fish having no effect on wave velocity.

Nearly all mussels require a host or hosts during the parasitic larval portion of their life cycle. Hosts are usually fish, but a few exceptional species utilize amphibians as hosts (Van Snik Gray et al., 2002; Howard, 1915) or may metamorphose without a host (Allen, 1924; Barfield et al., 1998; Lefevre and Curtis, 1911; 1912). Haag and Warren (1998) found that densities of host generalist mussels (using a variety of hosts from many different families) and displaying host specialists (using a small number of hosts usually in the same family but mussel females have behavioral modifications to attract hosts to the gravid female) were independent of the densities of their hosts. Densities of non-displaying host specialist mussels (using a small number of hosts usually in the same family but without host-attracting behavior) were correlated positively with densities of their hosts. Upstream dispersal of host fish for non-displaying host specialist mussels could, theoretically, transport mussel larvae (glochidia) over long distances through unsuitable habitat, but it is unlikely that this occurs very often. D. Strayer (personal communication) suggested a distance of at least 10 km, but a greater distance between occurrences may be necessary to constitute genetic separation of populations. As such, separation distance is based on a set, though arbitrary, distance between two known points of occurrence.

Date: 18Oct2004
Author: Cordeiro, J.
Notes: Contact Jay Cordeiro (jay_cordeiro@natureserve.org) for a complete list of freshwater mussel taxa sorted by flow regime.
Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 08Jan2014
NatureServe Conservation Status Factors Author: Jackson, D. R. (2013); Cordeiro, J. (2009); Butler, R. S. (1992)
Element Ecology & Life History Edition Date: 06Mar2007
Element Ecology & Life History Author(s): Cordeiro, J. (2007); BUTLER, R.S. (1992)

Zoological data developed by NatureServe and its network of natural heritage programs (see Local Programs) and other contributors and cooperators (see Sources).

References
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  • Battle, J., S.W. Golladay, and A.R. Bambarger. 2003. Mussel conservation in the Chickasawhatchee and Elmodel Wildlife Management Areas: methods for a relocation study. Pages 860-863 in K.J. Hatcher (ed.) Proceedings of the Georgia Water Resources Conference, Institute of Ecology, The University of Georgia, Athens, Georgia.

  • Fuller, S.L.H. 1974. Chapter 8: Clams and mussels (Mollusca: Bivalvia). Pages 215-273 in: C.W. Hart, Jr. and S.L.H. Fuller (eds.) Pollution Ecology of Freshwater Invertebrates. Academic Press: New York. 389 pp.

  • Gagnon, P., W. Michener, M. Freeman, and J. Brim Box. 2006. Unionid habitat and assemblage composition in coastal plain tributaries of Flint River (Georgia). Southeastern Naturalist, 5(1): 31-52.

  • Golladay, S.W., P. Gagnon, M. Kearns, J.M. Battle, and D.W. Hicks. 2004. Response of freshwater mussel assemblages (Bivalvia: Unionidae) to a record drought in the Gulf Coastal Plain of southwestern Georgia. Journal of the North American Benthological Society, 23(3): 494-506.

  • Heard, W.H. 1964. Corbicula fluminea in Florida. The Nautilus 77(3):105-107.

  • Heard, W.H. 1975. Determination of the endangered status of freshwater clams of the Gulf and southeastern states. Department of Biological Sciences, Florida State University, Tallahassee, Florida. Final Report prepared for the Office of Endangered Species, Bureau of Sport Fisheries and Wildlife, U.S. Department of Interior, contract number 14-16-000-8905. 31 pp.

  • Heard, W.H. 1979. Identification manual of the fresh water clams of Florida. State of Florida, Department of Environmental Regulation, Technical Series, 4(2): 1-82.

  • Howard, A.D. 1915. Some exceptional cases of breeding among the Unionidae. The Nautilus 29:4-11.

  • Lefevre, G. and W.T. Curtis. 1912. Studies on the reproduction and artificial propogation of fresh-water mussels. Bulletin of the Bureau of Fisheries 30:102-201.

  • Lydeard, C., R. Minton, and J.D. Williams. 2000. Prodigious polyphyly in imperiled freshwater pearly-mussels (Bivalvia: Unionidae): a phylogenetic test of species and generic designations. Pages 145-158 In E.M. Harper, J. D. Taylor, and J. A. Crame (eds.) The Evolutionary Biology of the Bivalvia. Geological Society of London, Special Publications 177.

  • Moyle, P. and J. Bacon. 1969. Distribution and abundance of molluscs in a fresh water environment. Journal of the Minnesota Academy of Science 35(2/3):82-85.

  • Ortmann, A.E. and B. Walker. 1922. On the nomenclature of certain North American naiades. Occasional Papers of the Museum of Zoology, University of Michigan, 112: 1-75.

  • Serb, J.M., J.E. Buhan, and C. Lydeard. 2003. Molecular systematics of the North American freshwater bivalve genus Quadrula (Unionidae: Ambleminae) based on mitochondrial ND1 sequences. Molecular Phylogenetics and Evolution, 28: 1-11.

  • Simpson, C.T. 1900. Synopsis of the naiades, or pearly freshwater mussels. Proceedings of the United States National Museum, 22(1205): 501-1044.

  • Simpson, C.T. 1914. A Descriptive Catalogue of the Naiades or Pearly Fresh-water Mussels. Bryant Walker: Detroit, Michigan. 1540 pp.

  • Strayer, D. 1983. The effects of surface geology and stream size on freshwater mussel (Bivalvia, Unionidae) distribution in southeastern Michigan, U.S.A. Freshwater Biology 13:253-264.

  • Strayer, D.L. 1999a. Use of flow refuges by unionid mussels in rivers. Journal of the North American Benthological Society 18(4):468-476.

  • Strayer, D.L. and J. Ralley. 1993. Microhabitat use by an assemblage of stream-dwelling unionaceans (Bivalvia) including two rare species of Alasmidonta. Journal of the North American Benthological Society 12(3):247-258.

  • Turgeon, D.D., J.F. Quinn, Jr., A.E. Bogan, E.V. Coan, F.G. Hochberg, W.G. Lyons, P.M. Mikkelsen, R.J. Neves, C.F.E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F.G. Thompson, M. Vecchione, and J.D. Williams. 1998. Common and scientific names of aquatic invertebrates from the United States and Canada: Mollusks. 2nd Edition. American Fisheries Society Special Publication 26, Bethesda, Maryland: 526 pp.

  • Van der Schalie, H. 1938a. The naiad fauna of the Huron River in southeastern Michigan. Miscellaneous Publication of the Museum of Zoology, University of Michigan 40:7-78.

  • Watters, G.T. 1992a. Unionids, fishes, and the species-area curve. Journal of Biogeography 19:481-490.

  • Williams, J. D.  2013.  Comments on the status of the bivalve mussels Quadrula infucata and Q. kleiniana.  E-mail to D. R. Jackson, dated 22 December 2013.

  • Williams, J. D.  2013a.  Comments on the status of the bivalve mussels Quadrula infucata and Q. kleiniana.  E-mail to D. R. Jackson, dated 22 December 2013.

  • Williams, J. D., A. E. Bogan, and J. T. Garner. 2008. Freshwater Mussels of Alabama and the Mobile Basin in Georgia, Mississippi and Tennessee. University of Alabama Press, Tuscaloosa, Alabama. 908 pp.

  • Williams, J. D., R. S. Butler, G. L. Warren, and N. A. Johnson.  2014.  Freshwater Mussels of Florida.  University of Alabama Press, Tuscaloosa.  498 pp.

  • Williams, J. D., R. S. Butler, G. L. Warren, and N. A. Johnson.  2014a.  Freshwater Mussels of Florida.  University of Alabama Press, Tuscaloosa, Alabama. 498 pp.

  • Williams, J.D., M.L. Warren, Jr., K.S. Cummings, J.L. Harris, and R.J. Neves. 1993b. Conservation status of freshwater mussels of the United States and Canada. Fisheries 18(9): 6-22.

  • Williams, James, Arthur Bogan, and Jeffrey Garner. 2008. Freshwater Mussels of Alabama and the Mobile Basin in Georgia, Mississippi, and Tennessee. University of Alabama Press, Tuscaloosa, AL. pp 908

  • van der Schalie, H. 1940. The naiad fauna of the Chipola River in northwestern Florida. Lloydia 3(3):191-208.

References for Watershed Distribution Map
  • Brim Box, J. and J.D. Williams. 2000. Unionid mollusks of the Apalachicola Basin in Alabama, Florida, and Georgia. Alabama Museum of Natural History Bulletin, 21: 1-143.

  • Clench, W.J. and R.D. Turner. 1956. Freshwater mollusks of Alabama, Georgia, and Florida from the Escambia to the Suwanee River. Bulletin of the Florida State Museum Biological Sciences, 1(3): 97-239.

  • Mirarchi, R.E., J.T. Garner, M.F. Mettee, and P.E. O'Neil. 2004b. Alabama wildlife. Volume 2. Imperiled aquatic mollusks and fishes. University of Alabama Press, Tuscaloosa, Alabama. xii + 255 pp.

  • Williams, J.D., A.E. Bogan, and J.T. Garner. 2008. Freshwater Mussels of Alabama & the Mobile Basin in Georgia, Mississippi & Tennessee. University of Alabama Press: Tuscaloosa, Alabama. 908 pp.

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Ridgely, R.S., T.F. Allnutt, T. Brooks, D.K. McNicol, D.W. Mehlman, B.E. Young, and J.R. Zook. 2003. Digital Distribution Maps of the Birds of the Western Hemisphere, version 1.0. NatureServe, Arlington, Virginia, USA.

Acknowledgement Statement for Bird Range Maps of North America:
"Data provided by NatureServe in collaboration with Robert Ridgely, James Zook, The Nature Conservancy - Migratory Bird Program, Conservation International - CABS, World Wildlife Fund - US, and Environment Canada - WILDSPACE."

Citation for Mammal Range Maps of North America:
Patterson, B.D., G. Ceballos, W. Sechrest, M.F. Tognelli, T. Brooks, L. Luna, P. Ortega, I. Salazar, and B.E. Young. 2003. Digital Distribution Maps of the Mammals of the Western Hemisphere, version 1.0. NatureServe, Arlington, Virginia, USA.

Acknowledgement Statement for Mammal Range Maps of North America:
"Data provided by NatureServe in collaboration with Bruce Patterson, Wes Sechrest, Marcelo Tognelli, Gerardo Ceballos, The Nature Conservancy-Migratory Bird Program, Conservation International-CABS, World Wildlife Fund-US, and Environment Canada-WILDSPACE."

Citation for Amphibian Range Maps of the Western Hemisphere:
IUCN, Conservation International, and NatureServe. 2004. Global Amphibian Assessment. IUCN, Conservation International, and NatureServe, Washington, DC and Arlington, Virginia, USA.

Acknowledgement Statement for Amphibian Range Maps of the Western Hemisphere:
"Data developed as part of the Global Amphibian Assessment and provided by IUCN-World Conservation Union, Conservation International and NatureServe."

NOTE: Full metadata for the Bird Range Maps of North America is available at:
http://www.natureserve.org/library/birdDistributionmapsmetadatav1.pdf.

Full metadata for the Mammal Range Maps of North America is available at:
http://www.natureserve.org/library/mammalsDistributionmetadatav1.pdf.

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