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C.B. Eads, J.E. Price, and J.F. Levine
22001155 SOUTHEASTERN NATURALIST Vo1l4.( 114):,8 N5–o9. 71
Fish Hosts of Four Freshwater Mussel Species in the Broad
River, South Carolina
Chris B. Eads1, Jennifer E. Price2, and Jay F. Levine1,*
Abstract - Freshwater mussel distributions are heavily reliant upon the range and movement
of host fishes and are subject to range restrictions when fish migration is blocked. The
Columbia Dam on the Broad River in Columbia, SC, has been a barrier to the migration of
anadromous species and other fish in the river since 1824. As a result, 5 freshwater mussel
species are restricted to reaches of the river downstream of the dam. In 2006, a fish passage
was created to facilitate fish movement between stream reaches above and below the dam.
Fish hosts that use the passage could facilitate the recolonization of reaches above the dam
by freshwater mussels. We conducted laboratory trials to determine the fish hosts of 4 of
the species limited to reaches below the dam. The most suitable hosts for Lampsilis cariosa
(Yellow Lampmussel) were Morone chrysops (White Bass), Morone saxatilis (Striped
Bass), and Pomoxis nigromaculatus (Black Crappie), whereas Micropterus salmoides
(Largemouth Bass) and Micropterus dolomieu (Smallmouth Bass) transformed fewer juvenile
mussels. Lampsilis siliquoidea (Fatmucket) hosts were Largemouth Bass and Perca
flavescens (Yellow Perch). Yellow Perch, Largemouth Bass, Lepomis macrochirus (Bluegill),
and Lepomis gibossus (Pumpkinseed) were the best hosts for Ligumia nasuta (Eastern
Pondmussel). We suspected that Elliptio roanokensis (Roanoke Slabshell) used anadromous
fishes as hosts because its distribution is limited to mainstem rivers below the downstreammost
dam. We confirmed that 2 Clupeidae, Dorosoma cepedianum (Gizzard Shad) and
Alosa aestivalis (Blueback Herring), and 1 Moronidae, Morone americana (White Perch),
are hosts for Roanoke Slabshell. Many of the host-fish species identified in this study are
highly mobile, and we expect the range of these mussels to eventually expand upstream of
the Columbia Dam as fish make use of the new passage.
Introduction
Freshwater mussels (Unionidae) have a unique lifecycle in which their larvae
(glochidia) must attach to a host (usually a fish) to metamorphose into their next
life stage (McMahon 1991). This requirement makes freshwater mussels especially
susceptible to environmental alterations because their habitat must support the adult
mussels, which are sessile, benthic filter-feeders, as well as their free-swimming
fish hosts to successfully reproduce. Fish are the mollusks’ primary means of dispersal
and facilitate colonization of new habitats by freshwater mussels (Strayer
2008). When physical barriers such as dams block fish migrations, upstream mussel
dispersal is also blocked (Brainwood et al. 2008, Dean et al. 2002, Fritts et al. 2012,
Locke et al. 2003, Watters 1996).
1North Carolina State University, College of Veterinary Medicine, Raleigh, NC 27607.
2John Tyler Community College, 800 Charter Colony Parkway, Midlothian, VA 23114-
4383. *Corresponding author - jay_levine@ncsu.edu.
Manuscript Editor: Arthur Bogan
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On the Broad River in Columbia, SC, a diversion dam and its associated Columbia
Canal were completed in 1824 (National Register Information System
2014). Since that time, the dam has blocked freshwater mussel and fish migration
in the river causing changes in mussel assemblages upstream and downstream of
the dam (Price et al. 2009). Price and coworkers (2009) found 8 species of mussels
immediately downstream of the dam, including Lampsilis cariosa (Say) (Yellow
Lampmussel), Lampsilis radiata (Gmelin) (Eastern Lampmussel), Ligumia nasuta
(Say) (Eastern Pondmussel), Villosa delumbis (Conrad) Eastern Creekshell, Elliptio
roanokensis (I. Lea) (Roanoke Slabshell), Elliptio complanata (Lightfoot) (Eastern
Elliptio), Elliptio congaraea (I. Lea) (Carolina Slabshell), and Elliptio angustata (I.
Lea) (Carolina Lance). The same study detected only 4 species in the Broad River
upstream of the dam: Eastern Creekshell, Eastern Elliptio, Carolina Lance, and
Uniomerus carolinianus (Bosc) (Florida Pondhorn). In 2006, the city of Columbia,
SC, constructed a fishway on the dam to facilitate the passage of anadromous fish
such as Alosa sapidissima (Wilson) (American Shad) and Morone saxatilis (Walbaum)
(Striped Bass). Freshwater mussel populations should have also benefited
from this fishway, which may have facilitated dispersal upstream on fish hosts.
However, little is known about the host fishes of mussels limited to reaches downstream
of the dam. Tedla and Fernando (1969) and Wick (2006) studied the hosts
of Yellow Lampmussel and Eastern Lampmussel at the northern part of the species’
range, but hosts for the remaining species are unknown.
The Yellow Lampmussel ranges from the Ogeechee River Basin in Georgia
north to Nova Scotia and Cape Breton Island and westward into the St. Lawrence
River Basin to the lower Ottawa and Madawaska River Basins (Johnson 1970).
Morone americana (Gmelin) (White Perch) and Perca flavescens (Mitchill) (Yellow
Perch) were previously shown to serve as hosts for Yellow Lampmussels in
the laboratory (Wick 2006), and wild White Perch were also found to be a common
host in Maine (Kneeland and Rhymer 2008). Eastern Lampmussels are found from
the Pee Dee River in South Carolina, north to the St. Lawrence River Basin and
west to Lake Ontario (Johnson 1970). Yellow Perch serve as hosts for the Eastern
Lampmussel in the wild. Eastern Pondmussel occurs from the Savannah River in
South Carolina, north to the St. Lawrence River basin and westward through the
Mohawk River, Erie Canal, and Lake Erie (Johnson 1970). The host fish of this species
was unknown prior to our study (Bogan and Alderman 2004). The range of the
Roanoke Slabshell has been reported to extend from the Savannah River in Georgia,
to several watersheds in Virginia (Williams et al. 1993). Nothing was known about
the host requirements of Roanoke Slabshell, but because it is generally limited to
mainstem rivers below the downstream-most dam on a system, we believed it might
use an anadromous fish as its host.
Dams reduce hydrologic and habitat variability (McManamay et al. 2012) and
can geographically isolate fish and mussel populations (Brainwood et al. 2008,
Dean et al. 2002, Fritts et al. 2012, Locke et al. 2003, Watters 1996). Impoundments
are detrimental to many mussel species (Blalock and Sickel 1996, Vaughan and
Taylor 1999), and altered flow regimes and habitat can impact mussel populations
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2015 Vol. 14, No. 1
downstream of dams (Vaughan and Taylor 1999). Dam removal and construction
of fish passages can potentially mitigate these alternations to stream habitat and
fauna (Lenhart 2010). However, the process of dam removal can be detrimental
to remaining mussel populations (Sethi et. al. 2004), and may benefit or harm fish
populations, thus affecting mussel species that rely on them as hosts (Sethi et. al.
2004). The hosts of many imperiled mussel species are unknown or poorly described
(Watters 1996). As part of a project to evaluate the fish passage constructed
in Columbia, SC, and its potential benefits to freshwater mussels, we conducted
laboratory host-trials on 4 mussel species currently found only downstream of
the dam—Yellow Lampmussel, Eastern Lampmussel, Eastern Pondmussel, and
Roanoke Slabshell)—to determine which fish species in the Broad River would
potentially serve as hosts.
Methods
In the spring of 2007, we collected gravid Yellow Lampmussel, Eastern Lampmussel,
Eastern Pondshell, and Roanoke Slabshell from the Broad River near
Columbia, SC. We collected most host fish by boat electrofishing in the Congaree
River or backpack-electrofishing in Crane Creek near Columbia, SC; Anguilla
rostrata (Lesueur) (American Eel) were collected by boat electrofishing in the
canal between Lake Moultrie and the Santee River in Berkeley County, SC. We
purchased Alosa aestivalis (Mitchill) (Blueback Herring) from a bait store in Columbia,
SC, and all Yellow Perch tested were collected by angling in Jordan Lake
(Cape Fear River Basin) in Chatham County, NC. We transported all fish to the
Aquatic Epidemiology and Conservation Laboratory at North Carolina State University
in Raleigh, NC, in aerated coolers of ambient water and slowly acclimated
them to laboratory conditions. Because some of the fish species tested for Eastern
Pondmussel and Roanoke Slabshell were difficult to maintain in laboratory conditions,
we infested host fish with those species within 5 days of bringing the fish into
the laboratory. Additionally, the timing of brood release from short-term brooders
such as Roanoke Slabshell was more difficult to control compared to long-term
brooders, so we had to initiate that trial upon glochidial release from available females.
Fish exposed to Yellow Lampmussel and Eastern Lampmussel were held for
6 weeks prior to the initiation of the host trial to ensure that any attached glochidia
from the wild would transform prior to the laboratory exposure.
We extracted glochidia from Yellow Lampmussel (1 individual), Eastern Lampmussel
(2 individuals), and Eastern Pondmussel (2 individuals) by flushing their
marsupia with a water-filled syringe. Gravid Roanoke Slabshells were held individually
in 8-liter containers of laboratory water at 20–22 °C and monitored daily
for released glochidia. The Roanoke Slabshell trial was initiated when a single
female released a brood that consisted primarily of fertilized and actively snapping
glochidia. Glochidia were loose and had little to no associated mucous. For
Eastern Pondmussel and Roanoke Slabshell trials, we anesthetized all fish ≥20 cm
total length with tricaine methanosulfate (MS-222), and pipetted glochidia onto
left and right gill arches until adequate attachment was observed after rinsing the
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gills gently with conditioned water. Direct application was necessary because there
were not sufficient glochidia available to suspend them at adequate concentrations
in a tank large enough to comfortably hold and infest all the larger fish tested for
these species. We infected other fish by placing them together for 10–30 minutes
in a vigorously aerated bath of glochidia (2000–4000 glochidia/liter) from the
species being tested. We monitored the fish for glochidial attachment, after which
we separated all fish by species, transferred them to individual aquaria of various
sizes maintained at 20–24 °C. All fish were held individually except for Blueback
Herring, which were held as a group of 25 fish in a single oval 370-liter tank; 12
of those individuals survived to the end of the trial. We routinely monitored the
tanks for transformed juvenile mussels by siphoning the bottom of the tank through
a 105-μm sieve. Time and personnel constraints prevented us from monitoring
sloughed glochidia. We identified juvenile mussels either by foot movement or the
presence of adductor muscles. We ended the trial when we no longer found juvenile
mussels in the aquaria.
We tested 11 fish species representing 4 families as potential hosts for Yellow
Lampmussel (Table 1) and 10 species from the same 4 families as potential hosts for
Eastern Lampmussel (Table 2). Because Lampsilis spp. use mantle lures for host-fish
attraction (Haag and Warren 1999, Zanatta and Murphy 2006), we focused primarily
on more piscivorous fish as test subjects for these mussel species. We tested 21 fish
species representing 7 families as hosts for Eastern Pondshell (Table 3) and 20 species
from 8 different families as hosts for Roanoke Slabshell (Tables 4, 5).
Results
We recovered Yellow Lampmussel juveniles from 7 of the 11 fish species tested,
but there was substantial variation among fish species in the number of juveniles
Table 1. Results of fish-host trial with Lampsilis cariosa.
Mean # juveniles/
Common name Scientific name # fish tested fish (range)
Anguillidae
American Eel Anguilla rostrata 2 0.0
Centrarchidae
Redbreast Sunfish Lepomis auritus 2 0.0
Bluegill Lepomis macrochirus 2 1.0 (0–2)
Redear Sunfish Lepomis microlophus (Günther) 2 0.0
Smallmouth Bass Micropterus dolomieu 2 60.5 (57–64)
Largemouth Bass Micropterus salmoides 3 156.7 (0–423)
Black Crappie Pomoxis nigromaculatus 1 816.0
Moronidae
White Perch Morone americana 2 Died
White Bass Morone chrysops 1 1276.0
Striped Bass Morone saxatilis 1 4079.0
Percidae
Yellow Perch Perca flavescens 2 0.5 (0–1)
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Table 2. Results of fish-host-trial with Lampsilis radiata.
# juveniles /
Common name Scientific name # fish tested fish (range)
Anguillidae
American Eel Anguilla rostrata 1 0.0
Centrarchidae
Redbreast Sunfish Lepomis auritus 2 0.0
Bluegill Lepomis macrochirus 2 1.0
Redear Sunfish Lepomis microlophus 3 0.0
Smallmouth Bass Micropterus dolomieu 1 0.0
Largemouth Bass Micropterus salmoides 2 415.5 (314–517)
Black Crappie Pomoxis nigromaculatus 1 Died
Moronidae
White Perch Morone americana 2 Died
Striped Bass Morone saxatilis 2 1.0 (0–2)
Percidae
Yellow Perch Perca flavescens 2 333.0 (242–424)
Table 3. Results of fish-host trial with Ligumia nasuta.
# juveniles /
Common name Scientific name # fish tested fish (range)
Anguillidae
American Eel Anguilla rostrata 2 0.0
Catostomidae
Quillback Carpiodes cyprinus (Lesueur) 1 0.0
Northern Hogsucker Hypentelium nigricans (Lesueur) 1 0.0
Spotted Sucker Minytrema melanops (Rafinesque) 1 0.0
Shorthead Redhorse Moxostoma macrolepidotum (Lesueur) 1 0.0
Centrarchidae
Redbreast Sunfish Lepomis auritus 5 9.0 (1–28)
Pumpkinseed Lepomis gibbosus 1 78.0
Bluegill Lepomis macrochirus 3 172.3 (91–335)
Redear Sunfish Lepomis microlophus (Günther) 2 2.0 (0–4)
Largemouth Bass Micropterus salmoides 1 91.0
Cyprinidae
Whitefin Shiner Cyprinella nivea (Cope) 3 0.0
Spottail Shiner Notropis hudsonius (Clinton) 4 0.0
Coastal Shiner Notropis petersoni (Fowler) 3 0.0
Ictaluridae
Flat Bullhead Ameiurus platycephalus (Girard) 1 0.0
Channel Catfish Ictalurus punctatus (Rafinesque) 1 0.0
Flathead Catfish Pylodictis olivaris (Rafinesque) 1 0.0
Moronidae
White Perch Morone americana 1 0.0
Striped Bass Morone saxatilis 1 0.0
Percidae
Tessellated Darter Etheostoma olmstedi (Storer) 3 0.0
Yellow Perch Perca flavescens 2 301.0 (258–344)
Piedmont Darter Percina crassa (Jordan and Brayton) 2 0.0
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detected (Table 1). Morone chrysops (Rafinesque) (White Bass) and Striped Bass
yielded the highest number of juveniles. Juvenile mussels were obtained from 5
other species. Transformation was lower and variable on centrachid species, with
Pomoxis nigromaculatus (Lesueur) (Black Crappie) and Micropterus sp. harboring
the most juveniles. Transformation on Lepomis sp. and Yellow perch was low
and inconsistent (Table 1). Juvenile mussels remained attached to viable hosts for
14–22 days with a peak drop-off between days 16 and 18 for all host species.
We recovered Eastern Lampmussel juveniles from 4 of the 10 fish species tested
(Table 2). Micropterus salmoides (Lacépède) (Largemouth Bass) and Yellow Perch
produced the most juveniles. Of the centrachids evaluated, we recovered only one
juvenile from a Lepomis macrochirus Rafinesque (Bluegill); only 1 of 4 percids
yielded juveniles. Juvenile mussels remained encysted on viable hosts for 14–26
days with no noticeable differences between host species. Peak excystment for all
hosts came between days 18 and 22.
Table 4. Results of fish-host trial with Elliptio roanokensis. All Blueback Herring were maintained
in a single tank.
Mean # juveniles/
Common name Scientific name # fish tested fish (range)
Anguillidae
American Eel Anguilla rostrata 1 0.0
Catostomidae
Quillback Carpiodes cyprinus 2 0.0
Northern Hogsucker Hypentelium nigricans 1 Died
Spotted Sucker Minytrema melanops 1 0.0
Notchlip Redhorse Moxostoma collapsum (Cope) 1 0.0
Shorthead Redhorse Moxostoma macrolepidotum 1 0.0
Centrarchidae
Redbreast Sunfish Lepomis auritus 2 0.0
Bluegill Lepomis macrochirus 1 0.0
Redear Sunfish Lepomis microlophus 2 0.0
Largemouth Bass Micropterus salmoides 2 0.0
Smallmouth Bass Micropterus dolomieu 2 0.0
Black Crappie Pomoxis nigromaculatus 1 0.0
Clupeidae
Blueback Herring Alosa aestivalis 10 30.4
Gizzard Shad Dorosoma cepedianum 2 22.0 (20–24)
Cyprinidae
Whitefin Shiner Cyprinella nivea 3 0.0
Ictaluridae
Channel Catfish Ictalurus punctatus 1 0.0
Flathead Catfish Pylodictis olivaris 1 0.0
Moronidae
White Perch Morone americana 2 17.5 (0–35)
Striped Bass Morone saxatilis 1 0.0
Percidae
Yellow Perch Perca flavescens 1 0.0
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We recovered transformed Eastern Pondmussel juveniles from 6 of the 21 fish
species tested (Table 3). Yellow Perch yielded the most juveniles overall. Lepomis
gibbosus (L.) (Pumpkinseed), Bluegill, and Largemouth Bass as well as several
other centrachids were also relatively good hosts. The duration of glochidial attachment
on host fish ranged from 15–24 days. Excystment from Lepomis auritus L.
(Redbreast Sunfish) was completed by day 17 on 4 of the 5 individuals tested, but
other more efficient hosts had peak excystment of juveniles between days 20 and
Table 5. Results of fish-host trial with Elliptio roanokensis.
# juveniles
Common name Scientific name Replicate produced
Anguillidae
American Eel Anguilla rostrata A 0
Catostomidae
Quillback Carpiodes cyprinus A 0
Quillback Carpiodes cyprinus B 0
Northern Hogsucker Hypentelium nigricans A Died
Spotted Sucker Minytrema melanops A 0
Notchlip Redhorse Moxostoma collapsum A 0
Shorthead Redhorse Moxostoma macrolepidotum A 0
Centrarchidae
Redbreast Sunfish Lepomis auritus A 0
Redbreast Sunfish Lepomis auritus B 0
Bluegill Lepomis macrochirus A 0
Redear Sunfish Lepomis microlophus A 0
Redear Sunfish Lepomis microlophus B 0
Largemouth Bass Micropterus salmoides A 0
Largemouth Bass Micropterus salmoides B 0
Smallmouth Bass Micropterus dolomieu A 0
Smallmouth Bass Micropterus dolomieu B 0
Black Crappie Pomoxis nigromaculatus A 0
Clupeidae
Blueback Herring Alosa aestivalis 12 304
Gizzard Shad Dorosoma cepedianum A 24
Gizzard Shad Dorosoma cepedianum B 20
Cyprinidae
Whitefin Shiner Cyprinella nivea A 0
Whitefin Shiner Cyprinella nivea B 0
Whitefin Shiner Cyprinella nivea C 0
Ictaluridae
Channel Catfish Ictalurus punctatus A 0
Flathead Catfish Pylodictis olivaris A 0
Moronidae
White Perch Morone americana A 0
White Perch Morone americana B 35
Striped Bass Morone saxatilis A 0
Percidae
Yellow Perch Perca flavescens A 0
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22. There was no noticeable difference in transformation time between the more
viable host species.
Transformed Roanoke Slabshell juveniles were recovered from only 3 of the 20
fish species tested. Two Clupeidae species, Blueback Herring and Dorosoma cepedianum
(Lesueur) (Gizzard Shad) were hosts, and 1 of the 2 White Perch evaluated
also facilitated transformation (Table 5). Juveniles remained encysted on hosts for
10–16 days with no noticeable difference between host species.
Discussion
The unique double-edged relationship between freshwater mussels and their fish
hosts necessitates that conservationists carefully consider both the health and well
being of host fish and mussel populations when making conservation decisions.
Dam removal or the creation of a fish passage has intuitive benefits to freshwater
mussel-population health; however, any construction activity within a stream or
riparian areas poses a risk to remaining populations (Sethi et al. 2004, Vaughn and
Taylor 1999). A working knowledge of the fish-hosts required to sustain an individual
species of mussel is needed to understand the potential benefit or hazards posed
by dam removal or fish-passage construction. The Columbia Dam fish passage was
constructed in 2006, and planned assessments of fish and mussel distribution and
diversity prompted these host-fish trials.
The hosts we observed to be most effective for Yellow Lampmussel—White
Bass, Striped Bass, and Black Crappie—had not been previously reported in the
literature. White Perch have been shown to be effective hosts for this species at
the northern end of its range both in the laboratory (Wick 2006) and in the wild
(Kneeland and Rhymer 2008) but did not survive to the end of the encystment period
in our trial. Because its congeners White Bass and Striped Bass were robust
hosts in our trials, we suspect White Perch may also be effective hosts in the Broad
River. In contrast, Wick (2006) found Yellow Perch to be effective hosts for Yellow
Lampmussel in Maine, but the 2 Yellow Perch we tested yielded only a single
juvenile between them. This freshwater mussel exhibits a relatively high degree of
genetic divergence among populations in the formerly glaciated northern end of
its range (Kelly and Rhymer 2005). The variability in host efficacy we observed
between individual Largemouth Bass has been observed in other mussel–host-fish
relationships (Eads et al. 2010, Haag and Warren 1997). Fish develop an immune
response to repeated challenges with glochidia (Rogers and Dimock 2003). We may
have tested an individual fish that had been previously exposed to glochidia and had
become immune (Watters and O’Dee 1996). When possible, to prevent using a fish
that has become immune to glochidial attachment, fish used in laboratory host-trials
should be collected from places where no mussels exist. However, fish should be
collected from the same basin as the mussels whenever possible because fish from
the same basin have been shown to serve as more efficient hosts of some mussel
species (Rogers et al. 2001).
Largemouth Bass and Yellow Perch were strong hosts for Eastern Lampmussel.
Largemouth Bass commonly serve as hosts for mussels of the genus Lampsilis
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2015 Vol. 14, No. 1
(Haag and Warren 1997, Johnson et al. 2012, Keller and Ruessler 1997). Yellow
Perch have been identified as hosts for Eastern Lampmussel in the wild at the
northern end of its range (Kneeland and Rhymer 2008, Tedla and Fernando 1969).
Kneeland and Rhymer (2008) also found glochidia from this species on a single
Pumpkinseed in the wild but could not confirm whether it facilitated transformation.
We did not test Pumpkinseed, but no other Lepomis was a suitable host.
Hanek and Fernando (1978) reported L. radiata glochidia attached to Pumpkinseed
and Ambloplites rupestris (Rafinesque) (Rock Bass) but did not indicate whether
it was L. r. radiata (Gmelin) (Eastern Lampmussel) or L. r. siliquoidea (Barnes)
(Fatmucket), which they considered to be subspecies at the time. Trdan (1981) reviewed
Tedla and Fernando (1969) when he conducted additional life-history work
on the Fatmucket and speculated that “the Yellow Perch is one of only a few species
of fish that can serve as a host” for Eastern Lampmussel.
To our knowledge, little research has been conducted on the early life history of
Eastern Pondmussel (Bogan and Alderman 2004), but we found Yellow Perch to be
a robust host for this species. We suspect that with its tendency to serve as a host
for the mantle-luring Eastern Lampmussel in the wild (Kneeland and Rhymer 2008,
Tedla and Fernando 1969), Yellow Perch would likely also be attracted to the lure
of Eastern Pondmussel. Juveniles also successfully transformed on Bluegill and
several other centrachids. Corey and coworkers (2006) found that Largemouth Bass
and Pumpkinseed attacked the lure of Eastern Pondmussel, but Bluegill and Luxilus
cornutus (Mitchill) (Common Shiner) responded poorly to its lure in research
under controlled conditions. This finding suggests that Bluegills may be poor hosts
in nature even though they are suitable laboratory hosts. However, wild Bluegills
may behave differently and be more attracted to the lure under natural conditions.
The other two species that were attracted to the lure in the study by Corey et al.
(2006) also served as hosts in our laboratory study. Differences in the outcomes of
laboratory and field studies reflect the need for caution in making extrapolations
from laboratory studies. Hosts selected to propagate juveniles in the laboratory due
to success of transformation or availability (e.g., hatchery) may not be sympatric
with native species or play any role in their natural life-history.
As we expected, our results supported the idea that Roanoke Slabshell uses
one or more anadromous clupeids as hosts. Because of their sensitivity to capture,
transport, and holding (Chittenden 1971), we were unable to maintain and test
Alosa sapidissima Wilson (American Shad) or A. mediocris (Mitchill) (Hickory
Shad). However, we confirmed that 2 other Clupeids served as viable hosts in the
laboratory. For a fish species to be an efficient mussel host in the wild, the timing
of their spawning runs would have to coincide with release of Roanoke Slabshell
glochidia. Roanoke Slabshells were found to be gravid from late March to June
in the Broad River, but the majority were gravid in late March and mid-April at
water temperatures of 14–18 °C (Price and Eads 2011). Typical spawning temperatures
reported for Clupeids are 11–14 °C for Hickory Shad (Harris 2010),
12.5–15.5 °C for Alosa pseudoharengus (Wilson) (Alewife) (Bigelow and Schroeder
1953), 14–22°C for Blueback Herring (O’Connell and Angermeier 1999),
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and 13–20 °C for American Shad (Walburg and Nichols 1967). This timing suggests
the later-spawning Blueback Herring and American Shad are likely hosts in
the wild. Sampling for naturally infested fish during spawning runs is needed to
identify their role as E. roanokensis hosts.
White Perch were present above the Columbia Dam prior to the construction
of the fish passage (Bettinger et al. 2003). Only 1 of 2 White Perch evaluated
supported the transformation of juveniles. The other fish was collected from the
Congaree River where Roanoke Slabshell is relatively common, and may have had
prior exposure and was immune to glochidial attachment. This semianadromous
fish species makes late-winter to early spring spawning runs up coastal rivers (Mansueti
1961) and may be most plentiful in coastal rivers when Roanoke Slabshell is
releasing glochidia. Additional studies are needed to further evaluate White Perch
as a potential host for this mussel.
Many of the hosts we identified in this study, especially the Moronidae and Clupeidae,
are highly mobile and will likely benefit from fish passages including the
one on the Columbia Dam (Davis et al. 1970, Mansueti 1961, Smith and Hightower
2013). Increased host-fish movement will likely enable mussel species currently
restricted to reaches below the dam to colonize upstream habitats. Because of its
apparent reliance on anadromous fish, Roanoke Slabshell should especially benefit
from attempts to restore fish migration on coastal rivers through dam removal
and construction of fish passages. Smith (1985) noted that the range expansion of
Anodonta implicata (Say) (Alewife Floater) in the Connecticut River was likely
due to the construction of fish passages which allowed their Clupeid host to move
upstream. Likewise, efforts to expand mussel populations may also be beneficial
to American Shad, the eggs of which were found developing inside 49 out of 757
freshwater mussels examined in the Altamaha River (Wisniewski et al. 2013). A
majority of those eggs were found in Elliptio hopetonensis (I. Lea) (Altamaha
Slabshell), a congener of Roanoke Slabshell. The mantle of the mussels seemed to
provide a clean, well-oxygenated place for eggs to develop where the benthic habitat
was otherwise silt-laden and inhospitable to them. The intertwined ecological
relationship of freshwater mussels and their fish hosts stresses the need to manage
coastal rivers for all native species and for fully functioning ecosystems.
Acknowledgments
We thank the US Fish and Wildlife Service and the South Carolina Department of Natural
Resources for providing funding for this study. We also thank Art Bogan for his helpful
editorial suggestions.
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