Freshwater Sponges (Porifera: Spongillida) of Tennessee

Home , Ephydatia, Ephydatia fluviatilis, Ephydatia muelleri, Heteromeyenia, Radiospongilla, Spongilla

Authors: John Copeland, Stan Kunigelis, Jesse Tussing, Tucker Jett, and Chase Rich Source: The American Midland Naturalist, 181(2) : 310-326 Published By: University of Notre Dame URL: https://doi.org/10.1674/0003-0031-181.2.310

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Downloaded From: https://bioone.org/journals/The-American-Midland-Naturalist on 18 Sep 2019 Terms of Use: https://bioone.org/terms-of-use Access provided by United States Fish & Wildlife Service National Conservation Training Center Am. Midl. Nat. (2019) 181:310–326 Notes and Discussion Piece Freshwater Sponges (Porifera: Spongillida) of Tennessee

ABSTRACT.—Freshwater sponges (Porifera: Spongillida) are an understudied fauna. Many U.S. state and federal conservation agencies lack fundamental information such as species lists and distribution data. Such information is necessary for management of aquatic resources and maintaining biotic diversity. During the months of June, July, and August of 2013, 2014 and 2015 a survey was conducted in Tennessee to document species of freshwater sponges and obtain preliminary distribution data. Selected rivers and streams were sampled at two to five sites. A total of 520 sponge samples were collected. Spicules and gemmules were viewed and photographed using Scanning Electron Microscopy. Thirteen or 39% of the known species of freshwater sponges occurring in the North America have been collected from Tennessee’s rivers and streams. Eunapius fragilis, Spongilla lacustris, and Trochospongilla horrida were the most abundant and widely distributed species. Corvospongilla becki and Heteromeyenia latitenta were the least frequently encountered sponges.

INTRODUCTION Freshwater accounts for only 0.01% of the World’s water and about 0.8% of the Earth’s surface yet it supports at least 100,000 species (Dudgeon et al., 2005). Unfortunately, freshwater biodiversity has experienced a continuous decline over the past several decades. Dudgeon et al. (2005) reported losses of freshwater biodiversity are far greater than those of the most affected terrestrial ecosystems. Although sponges are relatively common inhabitants of freshwaters, they are understudied in the United States. Information needed for management and conservation is lacking. Watermolen (2008) reported the U. S. Fish and Wildlife Service noted the limitations of existing knowledge in its mid- 19700s review of freshwater sponges. In addition Watermolen (2008) noted conservationists have done little since that initial assessment to monitor or further evaluate the status of freshwater sponges. Manconi and Pronzato (2016a) reported the study of freshwater sponges (Spongillida) is a neglected field of study and the major risk of extinction is the decline of experts in their taxonomy and natural history. Over 8000 sponge species are recognized, of which less than 250 are freshwater sponges. All recent freshwater sponges belong to the phylum Porifera, class Demospongiae, order Haplosclerida, and suborder Spongillida. Six families are currently recognized, of which three Metaniidae, Potamolepidae, and Spongillidae occur in the United States. Fourteen genera and 33 species have been documented from the Nearctic Biogeographical Realm (Manconi and Pronzato, 2016b). Tennessee is recognized as one of the most aquatic biological diverse states within the United States (Stein, 2002). Tennessee’s aquatic diversity is due to the state’s geologic and hydrographic diversity (Etnier and Starnes, 1993). Six physiographic provinces and five major river basins (Fig. 1) are found within Tennessee. While much is known concerning the fishes (Etnier and Starnes, 1993) and freshwater mussels (Parmalee and Bogan, 1998) of Tennessee, little has been documented concerning freshwater sponges. Six reports concerning freshwater sponges in Tennessee have been published. Hoff (1943) documented four species from the Reelfoot Lake region: Eunapius fragilis (Leidy, 1851), Heteromeyenia tubisperma (Potts, 1881), Racekiela ryderi (Potts, 1882), and Radiospongilla crateriformis (Potts, 1882). Parchment (1966) collected Spongilla lacustris (L., 1759) from Stones River. Kunigelis and Copeland (2014) reported finding Ephydatia fluviatilis (L., 1759), Eunapius fragilis, Racekiela ryderi, Radiospongilla cerebellata (Bowerbank, 1863), Spongilla lacustris, and Trochospongilla horrida (Weltner, 1893) from eastern Tennessee. Copeland et al., (2015a) added Ephydatia muelleri (Lieberkuhn,¨ 1856) to the list of sponges from eastern Tennessee. Cherokeesia armata Copeland, Pronzato and Manconi, 2015 was recently discovered and described (Copeland et al., 2015b). To develop a better understanding of sponge diversity and species distributions a state-wide survey of selected rivers and streams was undertaken. Due to the uniqueness of the river drainages of Tennessee we expected to discover a diverse freshwater sponge fauna. 310

FIG. 1.—Five major river basins of Tennessee (A) Mississippi, (B) Tennessee, (C) Cumberland, (D) Barren, (E) Conasauga from Kernodle (1972)

METHODS During June, July, and August of 2013, 2014, and 2015 selected rivers and streams were surveyed to document the freshwater sponges of Tennessee and obtain distribution data. Criteria for determining waterways to be sampled were aquatic biological diversity, physiographic location, channelization, funding, and time available to researchers. No rivers and streams altered by channelization were sampled by suggestion of Tennessee Wildlife Resources Agency. Rivers and streams were surveyed by viewing appropriate hard substrates (i.e., rocks and logs) for sponges while wading. Depending upon stream length and access, two to five collection sites were established. A minimum of one human-hour of search time was spent at each collection site. Five hundred and twenty sponge samples were collected. Information collected for each sponge included: river or stream name, river drainage name, latitude and longitude of collection site, and county name. Latitude and longitude were obtained using a Garmin GPSmap 76CSx. Sponges were viewed using a 10 X head-band magnifier for the presence of gemmules. If gemmules were found a section of the sponge was collected. Sponges were preserved in 70% ethanol until processed for light microscopy (LM) and Scanning Electron Microscopy (SEM). To obtain clean spicule preparations for SEM observation and LM slides excised sponge was dissolved in test tubes containing 65% nitric acid. Once dissolved the remaining spicules were centrifuged to create a slug. Slugs were rinsed and centrifuged three times in distilled water, followed by a final rinse and spin in 70% ethanol. Spicules were pipetted on to glass slides for LM analysis and onto stubs for SEM analysis. Following the procedure of Manconi and Pronzato (2000), a glass substratum was placed under the spicules providing for a black background in the SEM photographs. Disassociated spicules, entire gemmules, and gemmule cross-sections were sputter- coated with gold and observed by Leo 982 and Hitachi TM 3000 SEM. Sponge identification was made using the keys of Manconi and Pronzato (2016b), Reiswig et al. (2010) and descriptions of Penney and Racek (1968). To determine range, mean, and standard deviation for spicular lengths and rotule diameters 30 to 80 of each type of spicule (megascleres, microscleres, and gemmuloscleres) were measured using the measurement program of the Hitachi TM 3000. Ten gemmules of each species were measured for diameter.

RESULTS AND DISCUSSION Thirteen described species of freshwater sponges, one of the family Potamolepidae and twelve of Spongillidae were collected. Scanning electron microscopy photos of gemmules and spicules for each species are provided (Figs. 2–8). Three species, Eunapius fragilis, Spongilla lacustris, and Trochospongilla horrida were frequently encountered. Corvospongilla becki, and Heteromeyenia latitenta may be the rarest sponges in Tennessee. Distribution information and spicular measurements for Tennessee specimens are provided. Species are listed alphabetically by family. Size ranges for spicules are provided, with means and standard deviations presented within parentheses. Distributional data for each species is presented (Figs. 9–13, Table 1).

FIG.2.—SEMphotos(A) Cherokeesia armata, megasclere, (B) Cherokeesia armata,strongyle gemmuloscleres, (C) Cherokeesia armata, protruding oxea gemmulosclere, (D) Cherokeesia armata, gemmule, (E) Corvospongilla becki, gemmuloscleres, (F) Corvospongilla becki, megasclere, (G) Corvospongilla becki, microsclere, (H) Corvospongilla becki, gemmule

POTAMOLEPIDAE BRIEN, 1967

During the early days of the survey a unique sponge was discovered (Copeland et al., 2015b). Through an international effort this sponge was described and named Cherokeesia armata Copeland, Pronzato, and Manconi 2015 (Copeland et al., 2015b). Until the finding of C. armata all known living members of the family Potamolepidae had a southern hemisphere distribution primarily in tropical Africa and South America. Prior to the finding of C. armata two fossils, one from Japan (Matsuoka and Masuda, 2000) and the other from Canada (Pisera et al., 2013), represented this family in the northern hemisphere. Gemmules of Potamolepidae are poorly adapted for overland dispersal (Manconi and Pronzato, 2015). These gemmules are usually sessile, without pneumatic layer and a thin gemmular theca (Manconi and Pronzato, 2015). Gemmules of this family are apparently adapted to persist in situ in extreme environments encountered in tropical ecosystems (i.e. extremely low water levels to extensive flooding and heavy silting) (Manconi and Pronzato, 2015). Potamolepidae is comprised of seven genera and 33 species.

FIG. 3.—SEM photos (A) Ephydatia fluviatilis, gemmulosclere, (B) Ephydatia fluviatilis, megasclere, (C) Ephydatia fluviatilis, gemmule, (D) Ephydatia muelleri, gemmulosclere, (E) Ephydatia muelleri, smooth megasclere, (F) Ephydatia muelleri, spiny megasclere, (G) Ephydatia muelleri, gemmule

Cherokeesia armata.—This sponge, representing a new genus for the family Potamolepidae, was discovered in the summer of 2013. It is currently the only living member of the family Potamolepidae known to occur not only in the Nearctic Biogeographical Realm but in the entire northern hemisphere (Copeland et al., 2015b). Currently its known distribution is limited to Tennessee (Fig. 9, Table 1). The initial discoveries of this sponge occurred in the Nolichucky and Hiwassee Rivers of eastern Tennessee. Additional collections have expanded the range into middle Tennessee. During 2014 C. armata was collected from the Buffalo and Sequatchie rivers, of the Tennessee River drainage, and from Red River, a tributary of the Cumberland River (Table 1). Eighty of each type of spicule were measured (Fig. 2). Spicules: megascleres stout, spiny oxeas, 161.8–228 (199.5 6 15.8) lm in length; microscleres absent; gemmuloscleres are of two types: strongyle-like 17–29.3 (22.4 6 2.5) lm in length, stout, entirely smooth, from extremely bent C-shaped to bean shaped to ring-shaped to button shaped and spiny to smooth oxeas 173.3–258 (219.7 6 16.7) lm in length. Gemmules 591–957 (749 6 84) lm in diameter.

SPONGILLIDAE GRAY, 1867

Spongillidae is a cosmopolitan family which contains more genera and species than any other freshwater sponge family (Manconi and Pronzato, 2015). Spongillidae gemmules have a complex design with gemmuloscleres arranged tangentially to radially to create an armor-like coating which protects totipotent cells. Several species have a pneumatic layer which allows for downstream gemmule dispersal.

FIG. 4.—SEM photos (A) Eunapius fragilis, gemmuloscleres, (B) Eunapius fragilis, megasclere, (C) Eunapius fragilis, gemmule, (D) Radiospongilla crateriformis, gemmulosclere, (E) Radiospongilla crateriformis, megasclere, (F) Radiospongilla crateriformis, gemmule

This type of gemmule seems to be perfectly designed for overland dispersal (Manconi and Pronzato, 2015). Corvospongilla becki Poirrier, 1978.—The distribution of this sponge within the United States is poorly known. It is reported from the type locality of Duck Lake in the Atachafalaya Basin, St. Martin Parish, Louisiana (Poirrier, 1978), the Cahaba River and Shades Creek of Alabama (Mobley, 2010), and from Canada (McGill University, 2017). We found two specimens of this sponge in the Duck River at Henry Horton State Park (Fig. 9, Table 1). The Duck River, a tributary of the Tennessee River, is one of the most biologically diverse rivers within the United States (Palmer, 2008). Seventy-five of each type of spicule were measured (Fig. 2). Spicules: megascleres are spiny, stout oxeas to strongyles 125–206.7 (173.2 6 20.2) lm in length; microscleres are micropseudobirotules 12.9–54.5 (33.2 6 7.1) lmin length with rotules composed of hooks 5.1–15 (10.9 6 2.1) lm in diameter; gemmuloscleres are strongyles of two sizes: large gemmuloscleres 70.5–139.3 (97.2 6 17.3) lm; and small gemmuloscleres 26.4–64.2 (40.9 6 7.2) lm in length, straight to slightly curved and covered in spines. Gemmules 635– 1007 (880 6 140) lm in diameter. Ephydatia fluviatilis.—Ephydatia fluviatilis is a cosmopolitan species of the northern hemisphere. This sponge has been reported from North America, Europe, India, and China. We collected this sponge from the Holston River in upper east Tennessee, Indian Creek a tributary of the Powell River in Claiborne County, and French Broad River at Seven Islands Bird State Park. The three waterways are drainages within the Ridge and Valley Physiographic Province and are components of the Tennessee River drainage (Fig. 9, Table 1). Fifty-three of each type of spicule were measured (Fig. 3). Spicules: megascleres are smooth to very sparsely spined oxeas 234–332 (296.4 6 22.4) lm in length; microscleres are absent; Gemmuloscleres are birotules 18.8–26.1 (22.5 6 1.8) lm in length with rotules 14–23.2 (18.7 6 1.8) lm in diameter. Gemmuloscleres may have no spines or as many as four on their shafts. The largest of these gemmulosclere measurements would be near the low end of ranges presented by Penney and Racek (1968) and Ricciardi and Reiswig (1993). Gemmules 322–425 (386 6 35) lm in diameter. Ephydatia muelleri.—This is a northern hemisphere sponge which has a wide distribution in North America and Europe (Penney, 1960). Its known distribution in Tennessee is presented in Figure 10 and Table 1. According to Reiswig et al. (2010), it shows a preference for temperate regions. Seventy-five of

FIG. 5.—SEM photos (A) Heteromeyenia tubisperma, gemmuloscleres, (B) Heteromeyenia tubisperma, microsclere, (C) Heteromeyenia tubisperma, megasclere, (D) Heteromeyenia tubisperma, gemmule, (E) Heteromeyenia latitenta, gemmulosclere, (F) Heteromeyenia latitenta, microsclere, (G) Heteromeyenia latitenta, macrosclere, (H) Heteromeyenia latitenta, gemmule

each type of spicule were measured (Fig. 3). Spicules: megascleres are completely smooth to spiny oxeas 182.6–300.7 (246.2 6 30) lm in length; microscleres are absent; gemmuloscleres are birotules with or without spines on shaft 8.3–15.3 (11.2 6 1.2) lm in length with rotules 10.8–23 (16.9 6 2.1) lmin diameter. Gemmules 349–460 (400 6 39) lm in diameter. Eunapius fragilis.—This sponge may be globally distributed, as it has been reported from Canada, Germany, India, Ireland, Mexico, Russia, South Africa, and United States (Van Soest, et al, 2018). Eunapius fragilis was first reported in Tennessee from the Reelfoot Lake region by Hoff (1943). This species is abundant and widely distributed across Tennessee (Fig. 10, Table 1). Fifty-eight of each type of spicule were measured (Fig. 4). Spicules: megascleres are smooth straight to slightly curved oxeas 154– 251 (199 6 19.8) lm in length; microscleres are absent; gemmuloscleres are straight to slightly curved spiny strongyles 35–113 (68.2 6 14.8) lm in length. Gemmules 337–481 (376 6 54) lm in diameter. Heteromeyenia latitenta (Potts, 1881).—Heteromeyenia latitenta has previously been reported only from the northeastern United States (Reiswig et al., 2010). We collected this sponge from the Pigeon River (Fig. 10, Table 1) where it was the second most abundant sponge collected. The Pigeon River has a long history of dioxin pollution. However, by the mid 1990s recovery efforts on the Pigeon were so successful that a program to re- introduce extirpated species of fish and mollusks was initiated (University of Tennessee Department of Forestry, Wildlife & Fisheries, 2004). Fifty-three of each type of spicule were measured (Fig. 5). Spicules: megascleres are straight oxeas 282–377 (334.5 6 23.1) lm in length;

FIG. 6.—SEM photos (A) Racekiela ryderi, gemmulosclere, (B) Racekiela ryderi, gemmulosclere, (C) Racekiela ryderi, megasclere, (D) Racekiela ryderi, gemmule, (E) Radiospongilla cerebellata, gemmulosclere, (F) Radiospongilla cerebellata, megasclere, (G) Radiospongilla cerebellata, gemmule

microscleres are slender spiny oxeas 10.2–14.5 (12.5 6 1.1) in length; gemmuloscleres are birotules 44.8–69.2 (55.8 6 7.4) lm in length with rotules 16.8–28.1 (21 6 2) lm in diameter. Gemmules 420– 483 (450 6 23) lm in diameter. Heteromeyenia tubisperma.—This sponge is primarily known from the eastern United States (Reiswig et al., 2010). Its distribution in Tennessee is presented in Figure 11 and Table 1. Heteromeyenia tubisperma was first reported in Tennessee by Hoff (1943) from the Reelfoot Lake region. Fifty of each type of spicule were measured (Fig. 5). Spicules: megascleres are oxeas 216–322 (288 6 19.6) lm in length; microscleres are oxeas 66–134 (108.7 6 14.4) lm in length; gemmuloscleres are birotules 45–69 (54.8 6 6.5) lminlength with rotules 14–26 (20.5 6 2.6) lm in diameter. Gemmules 483–535 (512 6 16) lm in diameter. Racekiella ryderi.—This sponge is currently known from the eastern half of North America and Ireland (Reiswig et al., 2010; Cocchiglia et al., 2013). However, Evans and Kitting (2010) reported finding this sponge in the California Delta. Its Tennessee distribution is presented in Figure 11 and Table 1. Racekiella ryderi produces two distinctly different gemmuloscleres. The smaller gemmuloscleres have flattened rotules and shafts with a few spines. Larger gemmuloscleres have several recurred spines on the shaft and recurved hooks on the ends. Thirty of each type of spicule were measured (Fig. 6). Spicules: megascleres are oxeas 197–297 (246 6 20.2) lm in length; microscleres are absent; gemmuloscleres are birotules of two lengths: smaller gemmuloscleres 28.9–45.4 (35.5 6 54.9) lmin

FIG. 7.—SEM photos (A) Spongilla lacustris, gemmulosclere, (B) Spongilla lacustris, microscleres, (C) Spongilla lacustris,megaclere,(D) Spongilla lacustris,gemmule,(E) Trochospongilla horrida, gemmulosclere, (F) Trochospongilla horrida, megasclere, (G) Trochospongilla horrida, gemmule

FIG. 8.—SEM photos (A) Trochospongilla leidii, gemmulosclere, (B) Trochospongilla leidii, megasclere, (C) Trochospongilla leidii, gemmule

FIG. 9.—Distributions of freshwater sponge species in Tennessee (A) Cherokeesia armata (B) Corvospongilla becki (C) Ephydatia fluviatilis

length with rotules 17.3–24.8 (20.5 6 1.6) lm in diameter; larger gemmuloscleres 44.4–64 (57 6 5) lm in length with numerous recurve spines on shaft and with a ring of recurved hooks at shaft ends 13–17.8 (15.2 6 1.2) lm in diameter. Gemmules 320–372 (343 6 18) lm in diameter. Radiospongilla cerebellata.—Some confusion exists concerning this sponge in the United States. The World Porifera Database distribution map for R. cerebellata does not show this species occurring in the western hemisphere (Van Soest et al., 2018) and Manconi and Pronzato (2016b) doubt its presence in the Nearctic. However, it has been reported from Texas (Poirrier, 1972) in the Cahaba River and in Shades Creek in Alabama (Mobley, 2010) and from eastern Tennessee by Kunigelis and Copeland (2014). Its Tennessee distribution is presented in Figure 11 and Table 1. Phenotypically this sponge appears to be R. cerebellata but DNA sequencing is needed for resolution. Fifty-five of each type of spicule were measured (Fig. 6). Spicules: megascleres are smooth oxeas 194–385 (307 6 47.6) lm in length; microscleres are absent; gemmuloscleres are curved spiny strongyles 65–94 (78.3 6 6.6) lm in length. Gemmules 451–583 (506 6 49) lm in diameter. Radiospongilla crateriformis.—This sponge was first reported in Tennessee by Hoff (1943) from the Reelfoot Lake region. Its Tennessee distribution is presented in Figure 12 and Table 1. This species has a Western Hemisphere distribution and has been reported from Argentina, Brazil, and United States (Van Soest et al., 2018). Fifty-five of each type of spicule were measured (Fig 4). Spicules: megascleres are slender slightly curved oxeas having small spines except near the tips 207–274 (240.7 6 16.1) lmin

FIG. 10.—Distributions of freshwater sponge species in Tennessee (A) Ephydatia mulleri (B) Eunapius fragilis (C) Heteromeyenia latitenta

length; microscleres are absent; gemmuloscleres are pseudorotulates with recurved spines 52–77 (64.7 6 5.7) lminlengthwithpseudorotules7.6–14.4(116 1.4) lmindiameter.Shaftsof gemmuloscleres with spines. Gemmules 347–462 (404 6 39) lm in diameter. Spongilla lacustris.—First reported in Tennessee from Stones River by Parchment (1966). Spongilla lacustris is widely distributed in the Northern Hemisphere. This species is common and widely distributed in Tennessee (Fig. 12, Table 1). Fifty of each type of spicule were measured (Fig. 7). Spicules: megascleres are smooth oxeas 165–280 (213.2 6 25.1) lm in length; microscleres are small, slightly curved acanthoxeas 41.4–79.9 (59.5 6 8.4) lm in length; gemmuloscleres are usually curved acanthoxeas or acanthostrongyls 66–142 (97.9 6 12.9) lm in length. Gemmules 343–447 (384 6 35) lm in diameter. Trochospongilla horrida.—This sponge has a northern hemisphere distribution and has been documented from Europe, China, and United States (Van Soest et al., 2018). First reported in Tennessee by Kunigelis and Copeland (2014), this sponge is widely distributed and abundant in Tennessee (Fig. 12, Table 1). Fifty-five of each type of spicule were measured (Fig. 7). Spicules: megascleres are straight to slightly curved acanthoxeas 145–210 (188.4 6 12.6) lm in length; microscleres are absent; gemmuloscleres are small birotules 6.6–9.5 (7.97 6 0.68) lm in length having rotules of slightly different diameters: small rotules 10.3–13.2 (11.8 6 0.6) lm in diameter; large rotules 12.3–15.6 (14.5 6 0.7) lm in diameter. Gemmules 314–372 (344 6 19) lm in diameter. This range for

FIG. 11.—Distributions of freshwater sponge species in Tennessee (A) Heteromeyenia tubisperma (B) Racekiella ryderi (C) Radiospongilla cerebellata

gemmule diameter is much smaller than the 475–540 lm range reported for T. horrida by Penney and Racek (1968) and Manconi and Pronzato (2016b). Trochospongilla leidii (Bowerbank, 1863).—Trochospongilla leidii has been collected from limited regions of the eastern United States (Reiswig et al., 2010) and the Panama Canal (Jones and Rutzler,¨ 1975). We found it only in the Cumberland River drainage within the Nashville Basin Physiographic Province (Fig. 13, Table1). Fifty-five of each type of spicule were measured (Fig. 8). Spicules: megascleres are smooth, straight to slightly curved, oxeas 105–166 (138 6 12.3) lm in length; microscleres are absent; gemmuloscleres are birotules 8.1–11.7 (9.7 6 0.8) lm in length with rotules of equal in size 13.1–17.8 (15.1 6 1) lm in diameter. Gemmules 359–453 (408 6 28) lm in diameter. Of the 33 species of freshwater sponges found in North America at least 13 occur in Tennessee waters. This accounts for 39% of the total North American freshwater sponge fauna. Additional sponge surveys of Tennessee’s lakes, rivers, and streams should be undertaken. We did not collect from Tennessee Valley Authority or U.S. Army Corp of Engineer Reservoirs. Reservoirs are widely distributed within the Tennessee and Cumberland Rivers and provide suitable sponge habitat. It is highly probable additional

FIG. 12.—Distributions of freshwater sponge species in Tennessee (A) Radiospongilla crateriformis (B) Spongilla lacustris (C) Trochospongilla horrida

FIG. 13.—Distribution of Trochospongilla leidii in Tennessee

TABLE 1.—Freshwater Sponge (Spongillida) of Tennessee distribution by species and river basin

Species River/Creek Latitude Longitude Basin County

Cherokeesia armata Buffalo River 35826026.500 N87853033.900 W Tennessee Wayne Cherokeesia armata Hiwassee River 35813023.600 N84831040.900 W Tennessee Polk Cherokeesia armata Hiwassee River 35813013.600 N84831007.100 W Tennessee Polk Cherokeesia armata Nolichucky River 36809022.000 N82843027.500 W Tennessee Greene Cherokeesia armata Nolichucky River 36810054.100 N82831045.400 W Tennessee Washington Cherokeesia armata Nolichucky River 36809022.000 N82843032.500 W Tennessee Greene Cherokeesia armata Red River 36833016.800 N87808033.700 W Cumberland Montgomery- Robertson Cherokeesia armata Sequatchie River 35808019.500 N85831002.300 W Tennessee Marion Corvospongilla becki Duck River 35835031.600 N8684100 19.000 W Tennessee Marshall Ephydatia fluviatilis Holston River 36828011.400 N82850003.100 W Tennessee Hawkins Ephydatia fluviatilis Indian Creek 36855019.900 N83836016.800 W Tennessee Claiborne Ephydatia fluviatilis French Broad 35856059.500 N83841044.400 W Tennessee Knox Ephydatia muelleri Elk River 35808019.200 N86822059.700 W Tennessee Lincoln Ephydatia muelleri Clear Creek 36806005.300 N83839043.100 W Tennessee Morgan Eunapius fragilis Buffalo River 35829045.200 N87849058.500 W Tennessee Perry Eunapius fragilis Buffalo River 35826026.500 N87853033.900 W Tennessee Wayne Eunapius fragilis Calfkiller River 35859027.900 N85823042.000 W Cumberland White Eunapius fragilis Clinch River 36834045.700 N84859000.300 W Tennessee Hancock Eunapius fragilis Clinch River 36828028.000 N83818013.200 W Tennessee Hancock Eunapius fragilis Daddy’s Creek 35855008.900 N8485501600 W Tennessee Cumberland Eunapius fragilis Duck River 35836043.000 N86800009.800 W Tennessee Maury Eunapius fragilis Duck River 35845016.500 N87816059.400 W Tennessee Maury Eunapius fragilis Duck River 35837004.400 N86851059.000 W Tennessee Maury Eunapius fragilis Duck River 35841041.400 N87813015.400 W Tennessee Maury Eunapius fragilis East Fork Obey 36836043.000 N85801032.000 W Cumberland Fentress River Eunapius fragilis Elk River 35824055.000 N8780000.9800 W Tennessee Giles Eunapius fragilis Elk River 35808059.100 N87830041.400 W Tennessee Lincoln Eunapius fragilis Harpeth River 39809006.600 N87807009.200 W Cumberland Cheatham Eunapius fragilis Harpeth River 36807025.800 N87805056.500 W Cumberland Davidson Eunapius fragilis Nolichucky River 36810054.100 N82831045.400 W Tennessee Washington Eunapius fragilis Nolichucky River 36809022.000 N82843032.500 W Tennessee Greene Eunapius fragilis Powell River 36833020.700 N83835020.700 W Tennessee Claiborne Eunapius fragilis Red River 36833016.800 N87808033.700 W Cumberland Mont/Rober Eunapius fragilis Red River 36835020.300 N87805019.900 W Cumberland Robertson Eunapius fragilis Sequatchie River 35808019.500 N85831002.300 W Tennessee Marion Eunapius fragilis Sequatchie River 35806036.500 N85831038.200 W Tennessee Marion Eunapius fragilis South Fork 36832052.700 N82836044.000 W Tennessee Sullivan Holston River Heteromeyenia latitenta Pigeon River 36857013.800 N83810040.900 W Tennessee Cocke Heteromeyenia Elk River 35836043.000 N8780000.9800 W Tennessee Giles tubisperma Heteromeyenia Richland Creek 36808043.300 N86851012.500 W Cumberland Davidson tubisperma Heteromeyenia Roaring River 36820032.700 N85825029.800 W Cumberland Overton tubisperma Heteromeyenia Wolf River 36833045.000 N85804031.000 W Cumberland Pickett tubisperma

TABLE 1.—Continued

Species River/Creek Latitude Longitude Basin County

Heteromeyenia Wolf River 36835000.000 N85807004.000 W Cumberland Pickett tubisperma Racekiela ryderi Clear Creek 36806005.300 N84839043.100 W Tennessee Morgan Racekiela ryderi Daddy’s Creek 35855008.900 N8485501600 W Tennessee Cumberland Racekiela ryderi Daddy’s Creek 36804042.900 N84845056.600 W Tennessee Morgan Racekiela ryderi Hiwassee River 35813013.600 N84831007.100 W Tennessee Polk Racekiela ryderi Obed River 36804010.800 N84839043.100 W Tennessee Morgan Racekiela ryderi Obed River 36804044.100 N84845044.100 W Tennessee Morgan Radiospongilla French Broad 35855006.900 N83801041.900 W Tennessee Cocke cerebellata River Radiospongilla French Broad 35855019.000 N82857041.500 W Tennessee Cocke cerebellata River Radiospongilla Nolichucky River 36812040.900 N82839049.600 W Tennessee Greene cerebellata Radiospongilla Nolichucky River 36810012.500 N82832031.100 W Tennessee Washington cerebellata Radiospongilla Pigeon River 36857013.800 N83810040.900 W Tennessee Cocke cerebellata Radiospongilla Abrams Creek 35833019.600 N83859053.800 W Tennessee Blount crateriformis Radiospongilla Elk River 35808059.100 N86830041.400 W Tennessee Lincoln crateriformis Radiospongilla Mill Creek 3585901300 N8684000900 W Cumberland Davidson crateriformis Radiospongilla Nolichucky River 36809022.000 N82843032.500 W Tennessee Greene crateriformis Radiospongilla Nolichucky River 36809022.000 N82843027.500 W Tennessee Greene crateriformis Radiospongilla Sequatchie River 35808019.500 N85831002.300 W Tennessee Marion crateriformis Spongilla lacustris Buffalo River 35827051.900 N87828045.700 W Tennessee Lewis Spongilla lacustris Cane Creek 35864011.800 N85833029.100 W Cumberland Van Buren Spongilla lacustris Clear Creek 36807017.000 N84844045.800 W Tennessee Morgan Spongilla lacustris Cumberland River 36809030.600 N86853031.700 W Cumberland Davidson Spongilla lacustris Duck River 35841041.400 N87813015.400 W Tennessee Maury Spongilla lacustris Little Sequatchie 35809059.700 N85857029.200 W Tennessee Marion River Spongilla lacustris Mill Creek 3585901300 N8684000900 W Cumberland Davidson Spongilla lacustris New River 36838026.100 N84855025.600 W Cumberland Scott Spongilla lacustris Nolichucky River 36810054.100 N82831045.400 W Tennessee Washington Spongilla lacustris Nolichucky River 36809022.000 N82843032.500 W Tennessee Greene Spongilla lacustris Piney Creek 35866014.600 N85838054.200 W Cumberland Van Buren Spongillia lacustris Obed River 36803042.700 N84857038.900 W Tennessee Cumberland Spongilla lacustris Red River 36835020.300 N87805019.900 W Cumberland Robertson Spongilla lacustris Richland Creek 36808043.300 N86851043.300 W Cumberland Davidson Spongilla lacustris South Fork 36832052.700 N82836044.000 W Tennessee Sullivan Holston River Trochospongilla horrida Abrams Creek 35833019.600 N83859053.800 W Tennessee Blount Trochospongilla horrida Buffalo River 35827051.900 N87828045.700 W Tennessee Lewis

TABLE 1.—Continued

Species River/Creek Latitude Longitude Basin County

Trochospongilla horrida Calfkiller River 35859027.900 N85823042.000 W Cumberland White Trochospongilla horrida Clear Creek 36807017.000 N84844045.800 W Tennessee Morgan Trochospongilla horrida Clinch River 36828028.000 N83818013.200 W Tennessee Hancock Trochospongilla horrida Collins River 35837005.700 N85841007.700 W Cumberland Warren Trochospongilla horrida Conasauga River 39800024.000 N84842049.800 W Conasauga Polk Trochospongilla horrida Cumberland River 36809030.600 N86853031.700 W Cumberland Davidson Trochospongilla horrida Duck River 35837004.400 N86851059.000 W Tennessee Maury Trochospongilla horrida Duck River 35835031.600 N8684100 19.000 W Tennessee Marshall Trochospongilla horrida Elk River 35836043.000 N8780000.9800 W Tennessee Giles Trochospongilla horrida Harpeth River 39809006.600 N87807009.200 W Cumberland Cheatham Trochospongilla horrida Little Sequatchie 35809059.700 N85857029.200 W Tennessee Marion River Trochospongilla horrida New River 36838026.100 N84855025.600 W Cumberland Scott Trochospongilla horrida Nolichucky River 36810054.100 N82831045.400 W Tennessee Washington Trochospongilla horrida Obed River 36803042.700 N84857038.900 W Tennessee Cumberland Trochospongilla leidii Cumberland River 36809030.600 N86853031.700 W Cumberland Davidson Trochospongilla leidii Harpeth River 39809006.600 N87807009.200 W Cumberland Cheatham Trochospongilla leidii Harpeth River 36808017.000 N86855044.200 W Cumberland Davidson

species will be found. Much remains to be documented concerning the distribution of sponges within Tennessee. Freshwater sponges represent a neglected and understudied faunal group. Other than species lists, of which most are more than likely incomplete, relatively little information concerning sponges is known for most states. We encourage state wildlife and conservation agencies to initiate surveys for freshwater sponges. We also suggest beginning graduate students interested in natural history studies consider taking on freshwater sponges.

Acknowledgments.—The authors acknowledge the Tennessee Wildlife Resources Agency, Lincoln Memorial University, the Well Being Foundation of Claiborne County, Tennessee for funding and The United States Geological Survey for providing the map used for showing species distributions. We are grateful to Bill Reeves, Tennessee Wildlife Resources Agency, David Withers, Tennessee Department of Environment and Conservation, and Stephanie Williams, Tennessee Department of Environment and Conservation for field assistance. Also, we gratefully acknowledge the expertise and constructive suggestions by the reviewers of this paper.

LITERATURE CITED

COCCHIGLIA, L., M. KELLY-QUINN, AND J. LUCEY. 2013. Classification of freshwater sponge collection at EPA Kilkenny. Environmental Protection Agency, Johnstown Castle, Co. Wexford, Ireland, p.31. COPELAND, J. E., J. A. TUSSING,T.M.JETT, AND S. KUNIGELIS. 2015a. Freshwater Porifera of eastern Tennessee. J. TN Acad. of Sci., 90:23. COPELAND, J., R. PRONZATO, AND R. MANCONI. 2015b. Discovery of living Potamolepidae (Porifera: Spongillina) from Neartic freshwater with description of a new genus. Zootaxa, 3957:37–48. DUDGEON, D., A. H. ARTHINGTON,M.O.GESSNER, Z-I. KAWABATA,J.D.KNOWLER,C.LE´ VEQUEˆ ,R.J.NAIMAN,A- HPRIEUR-RICHARD,D.SOTO, M.L.J. STIASSNY, AND C. A. SULLIVAN. 2005. Freshwater biodiversity: importance, threats, status and conservation challenges. Biol. Rev., 81:163–182. ETNIER,D.A.AND W. C. STARNES. 1993. The fishes of Tennessee. The University of Tennessee Press, Knoxville, TN, p. 681.

EVANS,L.EAND C. L. KITTING. 2010. Documentation and identification of the one known freshwater sponge discovered in the California Delta. Open Mar. Biol. J., 4:82–86. HOFF, C. C. 1943. Some records of sponges, branchiobdellids, and molluscs from the Reelfoot Lake region. J. TN. Acad. Sci., 18:223–227. JONES,M.L.AND K. RU¨ TZER. 1975. Invertebrates of the upper chamber, Gaun´ Locks, Panama Canal, with emphasis on Trochospongilla leidii (Porifera). Mar. Biol., 33:5–66. KERNODLE, J. M. 1972. Tributary river basins in Tennessee. Tennessee Water Resources Miscellanous Publication Number 8 p. 5. KUNIGELIS,S.C.AND J. E. COPELAND. 2014. Identification of isolated and in situ freshwater sponge spicules of eastern Tennessee. Microsc. Microanal., 20 (Suppl. 3):1294–1295. MANCONI,R.AND R. PRONZATO. 2000. Rediscovery of the type material of Spongilla lacustris (L., 1759) in the Linnean herbarium. Ital. J. Zool., 67:89–92. MANCONI,R.AND R. PRONZATO. 2015. Phylum Porifera, p. 133–157. In: Thorp, J. and D. C. Rogers (eds.) Ecology and General Biology: Thorp and Covich’s Freshwater Invertebrates. 4th Edition, Vol. 1. Academic Press, London, UK. MANCONI,R.AND R. PRONZATO. 2016a. How to survive and persist in temporary freshwater? Adaptive traits of sponges (Porifera: Spongillida): A review. Hydrobiologia, 782:11–22. MANCONI,R.AND R. PRONZATO. 2016b. Phylum Porifera, p. 39–83. In: Thorp, J. and D. C. Rogers (eds.): Keys to Nearctic Fauna: Thorp and Covich’s Freshwater Invertebrates. 4th Edition, Vol. 2. Academic Press, London, UK. MATSUOKA,T.AND Y. MASUDA. 2000. A new potamolepid freshwater sponge (Demospongiae) from the Miocene Nakamura Formation central Japan, Paleontol. Res., 4:131–137. MCGILL UNIVERSITY. 2017. Canadian biodiversity. Retrieved May 2, 2017, from thttp:// canadianbiodiversity.mcgill.ca/english/species/sponges/index.htm MOBLEY, A. S. 2010. The bacterial community of a freshwater sponge, Radiospongilla cerebellata:a comparison of terminal restriction fragment length polymorphisms (T-RFLP) and 16S RRNA clone library methods. M. S. Thesis, The University of Alabama at Birmingham. p. 63. PALMER, S. R. 2008. Eco logic...from the Nature Conservancy-Averting a water supply crisis while protecting endangered species: partnerships pay off for Tennessee’s Duck River. J. Amer. Wat. Wks. Assn., 100 (8):40–43. PARCHMENT, J. G. 1966. Notes on the ecology of sponges. J. TN Acad. Sci., 41:65. PARMALEE,P.W.AND A. E. BOGAN. 1998. The freshwater mussels of Tennessee. The University of Tennessee Press, Knoxville, TN, p. 328. PENNEY, J. T. 1960. Distribution and bibliography (1892–1957) of the fresh water sponges. University of South Carolina Publications, Series 3, 3:1–97. PENNEY,J.T.AND A. A. RACEK. 1968. Comprehensive Revision of a Worldwide Collection of Freshwater Sponges (Porifera: Spongillidae). U. S. Nat. Mus. Bull., 272. p. 184. PISERA, A., P. A. SIVER, AND A. P. WOLF. 2013. A first account of freshwater potamolepid sponge (Demospongiae, Spongillina, Potamolepiidae) from the middle Eocene: biogeographic and paleoclimatic implications. J. Paleontol., 87:373–378. POIRRIER, M. A. 1972. Additional records of Texas freshwater sponges (Spongillidae) with the First record of Radiospongilla cerebellata (Bowerbank, 1863) from the Western Hemisphere. The Southwest. Nat., 16:434–435. POIRRIER, M. A. 1978. Corvospongilla becki n. sp., a new fresh-water sponge from Louisiana. Trans. Amer. Micros. Soc., 97:240–243. REISWIG, H. M., T. M. FROST, AND A. RICCIARDI. 2010. Chapter 4 Porifera, p. 91–123. In: Thorp, J. H. and A. P. Covich, (eds.). Ecology and Classification of North American Freshwater Invertebrates. Academic Press, London, UK. RICCIARDI,A.AND H. M. REISWIG. 1993. Freshwater sponges (Porifera, Spongillidae) of Eastern Canada: taxonomy, distribution, and ecology. Can J. Zool., 71:665–682. STEIN, B. A. 2002. States of the Union: ranking America’s biodiversity. NatureServe, Arlington, VA, p. 1– 25.

UNIVERSITY OF TENNESSEE DEPARTMENT OF FORESTRY,WILDLIFE &FISHERIES. 2004. Pigeon River restoration project. Retrieved March 23, 2017 from web.utk.edu/~mjwilson/index.php. VAN SOEST, R. W. M., N. BOURY-ESNAULT,J.N.A.HOOPER,K.RU¨ TZLER,N.J.DE VOOGD,B.ALVAREZ,E.HAJDU, A. B. PISERA,R.MANCONI,C.SCHO¨ NBERG,M.KLAUTAU,B.PICTON,M.KELLY,J.VACELET,M. DOHRMANN, M.-C. DIAZ,P.CA´ RDENAS,J.L.CARBALLO,P.RIOS, AND R. DOWNEY. 2018.World Porifera Database. Accessed at http://www.marinespecies.org/porifera/ on 2018-6-1. WATERMOLEN, D. J. 2008. Catalog of North American state and regional freshwater sponge references. Misc. Pub. PUB-SS-1040, Bureau of Science Services, Wisconsin Dept. of Nat. Res. Madison, WI, p. 1–12.

JOHN COPELAND and STAN KUNIGELIS, Lincoln Memorial University, Harrogate, Tennessee 37752 JESSE TUSSING and TUCKER JETT, United State Department of Agriculture, Carthage, Tennessee 37030, and CHASE RICH, Tennessee Wildlife Resources Agency, Nashville, Tennessee, 37211. Submitted: 7 August 2018; Accepted 20 December 2018.