Modern Records of the Pink Heelsplitter Mussel,
Potamilus alatus (Say, 1817), in the Ottawa River
Drainage, Québec and Ontario, Canada
Isabelle Picard, Jean-François Desroches, Frederick W. Schueler,
and André L. Martel
Northeastern Naturalist, Volume 16, Issue 3 (2009): 355–364
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2009 NORTHEASTERN NATURALIST 16(3):355–364
Modern Records of the Pink Heelsplitter Mussel,
Potamilus alatus (Say, 1817), in the Ottawa River
Drainage, Québec and Ontario, Canada
Isabelle Picard1, Jean-François Desroches2, Frederick W. Schueler3,
and André L. Martel4*
Abstract - Potamilus alatus (Pink Heelsplitter) is a rare freshwater mussel in the
Ottawa River drainage (Ontario/Québec, Canada), at the northeastern limit of its distribution.
There are few historical records, and one old specimen from an uncertain
locality. The discovery of the Pink Heelsplitter in the Ottawa River drainage dates
from 1863. A few specimens were reported up to 1901, but it was August 2001 before
another specimen was found in the Ottawa River, as an empty shell at Upper Duck
Island, near Ottawa. From 2001 to 2005, the authors found this freshwater mussel at
four localities along the Ottawa River, and two in the tributary South Nation River.
Records include 4 living specimens and 12 empty shells, of which 8 were in fresh
condition. The Pink Heelsplitter seems to persist sparsely in the Ottawa River, but it
may have been extirpated from one of its tributaries (South Nation River) before its
discovery due to mortality associated with Dreissena polymorpha (Zebra Mussel).
Introduction
Potamilus alatus (Say) (Pink Heelpsplitter) is a freshwater mussel (Mollusca:
Unionidae) considered to be uncommon or rare in Canada, where it
has been reported from the provinces of Manitoba, Ontario, and Québec, in
large, slow rivers. Fish-linkage studies indicate that in rivers in the United
States it uses the Aplodinotus grunniens Rafinesque (Freshwater Drum)
as host for the dispersal and development of its morphologically unusual
glochidia larvae (Clarke 1981, Watters 1994). It is distributed throughout
the Mississippi River drainage and in the St. Lawrence River drainage
from Lake Huron to Lake Champlain (Parmelee and Bogan 1998), with the
Ottawa River drainage (Fig. 1) being the northeastern extremity of its distributional
range.
With a large thin shell and a prominent postero-dorsal “wing,” the Pink
Heelsplitter became an ideal substrate for attachment by the introduced
Dreissena polymorpha (Pallas) (Zebra Mussel). Mortality associated with
Zebra Mussels has nearly extirpated Potamilus alatus from the Laurentian
Great Lakes (Gillis and Mackie 1994, Nalepa et al. 1996). Lake Erie, whose
11050-1 Saint-Marc, Sherbrooke, PQ, Canada, J1K 2T7. 2Cégep de Sherbrooke,
Techniques de Bioécologie, 475 du Parc, Sherbrooke, PQ, Canada, J1K 4K1. 3Bishops
Mills Natural History Centre, RR2, Bishops Mills, ON, Canada, K0G 1T0.
4Life Sciences, Malacology, Canadian Museum of Nature, PO Box 3443, Station D,
Ottawa, ON, Canada, K1P 6P4. *Corresponding author - amartel@mus-nature.ca.
356 Northeastern Naturalist Vol. 16, No. 3
populations of unionid mussels have basically been wiped out by the Zebra
Mussel, may previously have contained the bulk of Canada’s Pink Heelsplitter
populations. The Pink Heelsplitter is not yet listed as a “species at risk”
by the Committee on Endangered Wildlife in Canada (COSEWIC 2009), but
in a recent report on the national status of unionid mussels, Metcalfe-Smith
and Cudmore-Vokey (2004) evaluated the status of the Pink Heelsplitter as
“sensitive” in Canada. Currently this mussel is listed as “sensitive” in the
province of Ontario and “at risk” in the province of Québec (Metcalfe-Smith
and Cudmore-Vokey 2004).
The species has been known in the Ottawa River drainage from a few historical
records, including the Ottawa River proper near Ottawa (LaRocque
1938; Latchford 1882a, 1882b, 1889, 1906), the Ottawa River at the mouth
of Rouge River (Whiteaves 1863), the Quyon River (tributary to the Ottawa
upstream of Ottawa, below the present Chats Falls Dam; Latchford 1904),
as well as one record around Montreal, without details on locality (Clarke
1981). No specimens have survived from these records, and no specimens
from Québec are found in the collections of the Canadian Museum of Nature
(CMN) or the Redpath Museum (verified in 2001–2002); one exception is a
specimen collected in 1901 by H.M. Ami at an unknown location (Big[s?]
over Spring) along the Ottawa River and deposited at the CMN molluscs
collection (CMN-Moll cat. num. 10157). In his 1880–1881 survey of the
unionids of the Ottawa area, Latchford (1882b) concluded: “As I have not
met with it in my many excursions, I think it must be rare in this vicinity, or
at least be restricted to a small area.”
Database searches of various major molluscs collections in the US,
including that of Illinois Natural History Survey (INHS), Ohio State University
(OSU), University of Michigan Museum of Zoology (UMMZ),
National Museum of Natural History (NMNH; Smithsonian Institution),
and Academy of Natural Sciences of Philadelphia (ANSP), revealed no
additional historical or recent records of the Pink Heelsplitter from the
Ottawa River drainage.
Figure 1. Map showing the Ottawa River, between Quyon and Montréal, where Pink
Potamilus alatus (Heelsplitter) has been reported.
2009 I. Picard, J.-F. Desroches, F.W. Schueler, and A.L. Martel 357
In 1995, when F.W. Schueler began resurveying the Ottawa drainage’s
unionids, the Pink Heelsplitter was the only historically known mussel
species that was not rediscovered in the first years of field work (Schueler
1996a, 1996b, 1997). This paper reports on the subsequent rediscovery of
the Pink Heelsplitter in the Ottawa River, including (i) the first record since
Ami's 1901 specimen, (ii) first specimens from the province of Québec,
and (iii) first records from the South Nation River. Specimens are held in
Isabelle Picard’s collection (IP numbers), the collection of the BMNHC
(EOBM[MOLL]), or at CMN.
Methods
Beginning in 1993, we independently surveyed unionids along the Ottawa
River and its tributaries (Fig. 1) by examining the nearshore shallow
water littoral zone. One or more observers spent 30 to 120 minutes at each
site looking for live animals or shells while walking or wading along shore.
Localities were recorded by GPS or on regional navigation or topographical
maps. A bottom viewer was occasionally used for observing the river bottom.
At each site, we recorded habitat, relative water levels, all unionids found,
and the presence or absence of Zebra Mussels, as well as other environmental
variables required by our individual protocols. Snorkeling and SCUBA
diving were also involved in many surveys during 1993–2007, especially in
the Ottawa River proper and several of its main tributaries, such as the Mississippi
River, the Rideau River and the Gatineau River. Surveys conducted
using snorkeling or SCUBA diving last 30 to 120 min and involved two
people who looked for and collected or photographed live animals or empty
unionid shells.
In this paper a “fresh” empty (dead) shell refers to a specimen whose
valves displayed little or no sign of periostracum erosion, an unbroken or undamaged
and sharp outer shell margin, as well as a shiny non-eroded nacre.
Results
No Potamilus alatus were found in 12 surveys by F.W. Schueler (FWS)
of the St Lawrence River above Montreal, 10 I. Picard and J.-F. Desroches
(IP & J-FD ) surveys of tributaries of the lower Ottawa River, 199 surveys
(FWS, 66; ALM, 133) of the Rideau River, 85 surveys (F.W. Schueler, 76;
A.L. Martel, 9) of the Mississippi River, 9 surveys (IP & J-FD, 5; ALM, 4)
of tributaries of the Gatineau River, 8 ALM surveys of the Gatineau River
proper, or 38 surveys (FWS, 23; IP & J-FD, 7; ALM, 8) of the Ottawa
River above the Chats Falls dam.
In 15 surveys by IP & J-FD of rivers around the islands of Montreal,
and 92 shore and snorkeling/SCUBA diving surveys of the lower Ottawa
River (IP & J-FD, 10; FWS, 65; ALM, 17), and 38 FWS surveys of the
main-channel South Nation River (below Spencerville), we found a very
small number of Pink Heelsplitter in 7 surveys at 6 sites (Fig. 1), and were
358 Northeastern Naturalist Vol. 16, No. 3
given one shell collected by others. The rarity of Pink Heelsplitter across
the Ottawa River drainage was made obvious by its occurrence in only 7 out
of 506 (1.4 %) surveys (1993–2007). In the reaches of the rivers where we
found Pink Heelsplitter, it occurred in 7 out of 145 (4.8 %) surveys.
On 16 August 2001, ALM and Jacqueline Madill (CMN) conducted a
unionid survey on the western shoreline and shallow littoral zone of Upper
Duck Island (45º28'01"N, 75º37'28"W), ON, downstream of Ottawa, where
they found, partly buried in the sand of the shallow littoral zone, an old empty
shell of Pink Heelsplitter (Fig. 2). This specimen, although partly eroded
and represented by a single valve (100 mm), still clearly displayed the characteristic
prominent postero-dorsal “wing” and brightly colored pink nacre.
This specimen is currently being accessioned at the CMN Molluscs collection.
The site where this Potamilus specimen was collected was dominated
by shells of Elliptio complanata Lightfoot (Eastern Elliptio), Lampsilis
radiata (Gmelin) (Eastern Lampmussel), and L. cardium Rafinesque (Plain
Pocketbook), the latter being particularly common.
On 3 October 2001, F.W. Schueler and B. Wigney were surveying unionids
and the extent of Dreissena in the South Nation River, Prescott and
Russell counties, ON, Canada. At the foot of the High Falls, in Casselman,
(45º19'25"N, 75º5'42"W), they found an old, broken 85-mm Pink Heelsplitter
valve in 30–50 cm deep rocky bedrock flats at the foot of the shallow
Figure 2. Photograph of the first modern specimen of the Pink Heelsplitter (Potamilus
alatus), which was collected near the shore of Upper Duck Island, in middle of the
Ottawa River, near the Québec-Ontario boundary, on 16 August 2001. The specimen
measured 100 mm (shell length).
2009 I. Picard, J.-F. Desroches, F.W. Schueler, and A.L. Martel 359
bedrock falls (EOBM[MOLL]00415). Zebra Mussels were scattered or
dense under rocks there, the largest seen was 32.5 mm; other unionids noted
included one each of Lasmigona costata (Rafinesque) (Flutedshell), Eastern
Elliptio, and Eastern Lampmussel.
At Jessups Falls Conservation Area (45º33'33"N, 75º3'45"W; Fig. 1),
near the mouth of the South Nation, on a trampled bedrock shore of the
muddy river, they found an old broken valve of the Pink Heelsplitter
(greatest dimension = 96 mm EOBM[MOLL]00205). Zebra Mussels
were abundant, and no living Unionidae were seen: Leptodea fragilis
(Rafinesque) (Fragile Papershell) was abundant in the shell piles, with
lesser numbers of Pyganodon grandis (Say) (Giant Floater) and Eastern
Elliptio, and single shells of Utterbackia imbecillis (Say) (Paper Pondshell;
not previously reported from the South Nation; F.W. Schueler,
unpubl. data), and Ligumia recta (Lamarck) (Black Sandshell) (not previously
reported from the South Nation).
On 31 July 2002, Tim Haxton and student assistants from the Ontario
Ministry of Natural Resources were seining for fish in the Ottawa River at
the Petrie Islands (approximately 45°29'40"N, 75°30'50"W) and picked up
a fresh 101-mm Pink Heelsplitter empty shell, which was donated to the
BMNHC in 2005 (EOBM[MOLL]00363). The collection was made ≤5 m off
shore in water 2 m deep; the substrate was 20% sand and 80% clay/silt. This
stretch of the river was disturbed by decades of gravel extraction, which has
recently ended. Other collections of unionids from the Petrie Islands include,
in decreasing order of abundance, Eastern Elliptio, Eastern Lampmussel,
and Plain Pocketbook.
On 14 September 2002, a 112-mm freshly dead Pink Heelsplitter shell
(IP-M0201) was found by IP & J-FD in the Ottawa River at Lac des Deux-
Montagnes, west of Montreal Island, Québec, in a bay called Anse-à-l'Orme
(45°27'4''N, 73°56'21''W), at the junction of Pierrefonds and Senneville. The
shell was in shallow water, the water level of the lake being exceptionally low
at this time. The other species of freshwater mussels found at Anse-à-l'Orme
are, in decreasing order: Eastern Lampmussel, Elliptio spp. (mostly Eastern
Elliptio), Fragile Papershell, Plain Pocketbook, and Anodonta implicata Say
(Alewife Floater; first record in Quebec since 1976). Pink Heelsplitter was the
least frequent species. The Zebra Mussel is absent from this locality.
On 28 May 2005, two freshly dead Pink Heelsplitter shells and a single
living individual were found in the Réserve Faunique de Plaisance, at the
Pentecôte Bay of the Ottawa River, PQ (45°36'18''N, 75°3'51''W; IP-M0240,
IP-M0241). At the same place, on 3 June, within 2–3 hours, an additional
four fresh shells were collected (IP-M0248) and two live Pink Heelsplitters
were observed (Fig. 3). Pink Heelsplitters found at this locality ranged from
82 to 114 mm. Other unionids found were, in decreasing order of abundance,
Eastern Lampmussel, Fragile Papershell, Giant Floater, Eastern Elliptio,
and Black Sandshell. The two latter species were less frequent than Pink
Heelsplitter. No Zebra Mussels were observed.
360 Northeastern Naturalist Vol. 16, No. 3
On 26 September 2007, F.W. Schueler and a South Nation Conservation
crew surveyed islands in the Ottawa River at the mouth of the South
Nation River. In the fine-sand/clay bottom near a marshy island in the river
mouth (45.57543°N, 75.10286°W) at 50 cm depth, they found a living 107-
mm Pink Heelsplitter that was extensively crusted with little Dreissena. At
another site nearby (45.57207°N, 75.09270°W) they found an old 113-mm
Pink Heelsplitter shell. Other Unionids found were, in decreasing order of
abundance (all heavily fouled with Dreissena): Eastern Elliptio, Eastern
Lampmussel, Giant Floater, and Fragile Papershell (shell only).
Discussion
These results confirm that the Pink Heelsplitter persists in the Ottawa
River, and that it is present, though rare, between the historically known
locations of Ottawa and Montreal. Main-channel sites above Ottawa to the
historic site at Quyon have not been adequately investigated, and therefore
we do not know if the species inhabits that unionid-rich section of the river.
The South Nation River population may have been extinct before or shortly
after their discovery, because of the invasion of that Ottawa River tributary
by the Zebra Mussel, which began in 1999 (Naomi Langlois-Anderson,
South Nation Conservation Authority, Berwick, ON, Canada, pers. comm.)
Figure 3. Photograph depicting a live specimen of Pink Heelsplitter (Potamilus
alatus), collected at Plaisance, QC, along northern shore of the Ottawa River, on 28
May 2005. This is the first live record of this species for the province of Québec. The
specimen measured 114 mm (shell length).
2009 I. Picard, J.-F. Desroches, F.W. Schueler, and A.L. Martel 361
At the town of Casselman, the river is now much more heavily infested with
Zebra Mussels than it was in 2001 (F.W. Schueler, pers. observ.), though
our understanding of surviving unionid fauna in this deep muddy river is
still preliminary (Michele Rodrick, South Nation Conservation Authority,
Berwick, ON, Canada, pers. comm).
The Pink Heelsplitter is no doubt very rare in Québec, where it has been
given S1 species status (Paquet et al. 2005, Picard 2004), being listed “at risk”
in that province in the report by Metcalfe-Smith and Cudmore-Vokey (2004).
Its official current status in Ontario, where it was one of the first unionids to be
severely impacted by the Zebra Mussel in the Laurentian Great Lakes (Gillis
and Mackie 1994, Haag et al. 1993, Nalepa et al. 1996), is S3.
In most of its distributional range, one of the major threats to the Pink
Heelsplitter is the invasive Zebra Mussel. All unionids are affected by the
presence of Dreissena (Martel et al. 2001, Nalapa et al. 1996), but members
of the (former) subfamilies Anodontinae and Lampsilinae (Pink Heelsplitter
is included in the latter) are more sensitive to Dreissena than the Ambleminae
(Baker and Hornbach 1997, Haag et al. 1993). In Lake St. Clair, Gillis
and Mackie (1994) found a 93% decline of Pink Heelspliter and Fragile
Papershell soon after the Zebra Mussel invasion.
Environment Canada's Lower Great Lakes Unionid Database was queried
(Daryl McGoldrick, Environment Canada, Burlington, ON, Canada,
pers. comm.). It confirmed that the only eastern Lake Ontario records of
the Pink Heelsplitter were from the mouth of Marysville Creek on Hungry
Bay, Bay of Quinte (44°10'N, 77°12'W)—two samples from 1965
and 1995. Strayer and Jirka (1997) doubted a record from Sodus Bay, NY,
across from the Bay of Quinte, but it is possible that the species had at least
these two isolated populations in Lake Ontario, between the main range
of the species and the Ottawa drainage populations. Also, Pink Heelsplitter
has not been reported from the St. Lawrence River or its tributaries in
northern New York State (Strayer and Jirka 1997). If the species was found
in central Lake Ontario only in the Bay of Quinte and Sodus Bay, and if
these populations are now extirpated by the effects of the Zebra Mussel,
then the Ottawa River population is an isolated outlier of increased conservation
significance.
The abundance of the Pink Heelsplitter in the Ottawa River drainage
may be limited by the numbers of the host fish, the Freshwater Drum, present
in the system. This fish is commonly caught by anglers in the Ottawa
River (Brian Coad, CMN, Ottawa, ON, Canada, pers. comm.). The three
records (preserved specimens) of Freshwater Drum deposited in the CMN
fish collection (CMNfi1972-0142.1, 1972-0143.1, 1984-0257.11) and
originating from the Ottawa River have all been collected in the lower Ottawa
River, where historical and recent records of the Pink Heelsplitter also
occur (this study). The Freshwater Drum is also present but not commonly
caught in the South Nation River, where the Pink Heelsplitter has been
recently reported (this study). In the South Nation Conservation's 2001
362 Northeastern Naturalist Vol. 16, No. 3
Near-shore Community Index Netting of the lower section of the South
Nation River, 2 of 366 fish sampled were of this species (Naomi Langlois-
Anderson, 2007 pers. comm.). CMN’s fish collection also contains one
specimen of the Freshwater Drum, which was collected in a major tributary
of the South Nation River, 2 km from the South Nation River proper (Castor
River; CMNfi1975-0953.13). The North American distribution of the
Pink Heelsplitter overlaps, more or less, with that of the Freshwater Drum
(compare distribution maps of Pink Heelsplitter in Parmelee and Bogan
[1998], and that of the Freshwater Drum in Lee et al. [1980]). Details on
the possible relationships between the abundance and distribution of this
fish in the Ottawa River drainage, as factors explaining the current pattern
of abundance and distribution of the Pink Heelsplitter in the same drainage,
remain to be elucidated.
The vast Ottawa River drainage may represent a major refugium for
the Pink Heelsplitter, and other unionids, in Canada. It is believed that
acidic low-calcium water of the Ottawa River would not permit high
densities of Zebra Mussels to exist (Strayer 1991). Nonetheless, larvae
of the Zebra Mussel are flushed into the Ottawa River from the calciumrich
Mississippi, Rideau, and South Nation rivers where this pest mollusc
has caused severe devastation among unionid populations (Martel 1995;
Martel et al. 2001, 2006a). Time will tell if the low-calcium waters of the
Ottawa River will, as hypothesized, prevent the invasive Zebra Mussel
from causing significant damage to the rich unionid populations, including
the rare stocks of Pink Heelsplitter present in the river, particularly
between Ottawa and Montreal.
There is little that unites the character of the sites where the Pink
Heelsplitter was found except that they are large-river sites with soft substrate
and few macrophytes. The numbers and demography of the extant
populations are not known. Further surveys in the same areas and elsewhere
in the Ottawa River would allow a better assessment of the species
status (Schueler 1999).
Acknowledgments
The authors thank Nicolas Lauzon, Benoît Lafleur, and Bev Wigney for field assistance,
Tim Haxton for sending in shells collected by Ontario Ministry of Natural
Resources field parties, and the South Nation Conservation for support of work in
that drainage. They are also grateful to Janice Metcalfe-Smith and Daryl McGoldrick
for consultation. Environment Canada’s Lower Great Lakes Unionid Database was
helpful in allowing us to access Potamilus records from Ontario. We are indebted to
Jacqueline Madill for her assistance in the field, as well as for producing the distribution
map and for kindly helping with the final preparation of the manuscript. We
also thank Kathy Klein for a critical review of the manuscript. This study was in part
funded by support for the BMNHC by Frank Ross and Donna Richoux and by a CMN
research grant to A.L. Martel.
2009 I. Picard, J.-F. Desroches, F.W. Schueler, and A.L. Martel 363
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