Zootaxa,Freshwater leech (Annelida: Hirudinida

Zootaxa 1657: 1–21 (2007) www.mapress.com / zootaxa/

ISSN 1175-5326 (print edition)

Copyright © 2007 · Magnolia Press

ISSN 1175-5334 (online edition)

ZOOTAXA

Freshwater leech (Annelida: Hirudinida) distribution in the Canadian Province of Newfoundland and Labrador and adjacent regions: check-list, new records, new pigmentation forms, and Pleistocene refugia JACQUELINE MADILL1 & PETER HOVINGH2 1

Research Services, Canadian Museum of Nature, P.O. Box 3443, Station “D”, Ottawa, Ontario, K1P 6P4, Canada. E-mail: [email protected] 2 721 Second Avenue, Salt Lake City, Utah 84103, USA. E-mail: [email protected]

Abstract The freshwater leeches (Hirudinida) in the Province of Newfoundland and Labrador were investigated by examining the literature, the Canadian Museum of Nature and the United States National Museum of Natural History records, and a leech survey. New pigmentation forms are described for Erpobdella punctata (Leidy) and Erpobdella obscura (Verrill). This is the first published record for Haemopis lateromaculata Mathers in Canada. Four species (Glossiphonia elegans (Verrill), Helobdella modesta (Verrill), Erpobdella punctata, and Erpobdella obscura) were found in both Labrador and Newfoundland with Erpobdella obscura common in Labrador and the other three species common in Newfoundland. Seven other species of leeches were less abundant in Newfoundland with 6 of these species very restricted in distribution. The abundance of leech species in Newfoundland and the paucity of leech species in Labrador suggested that the island species were present in a Pleistocene refugium associated with Newfoundland or the Grand Banks. Post-Pleistocene barriers to leech mobility are examined, and possible timing of colonization events is proposed in this model. Key words: Annelida, Leeches, Hirudinida, Glossiphoniidae, Haemopidae, Hirudinidae, Erpobdellidae, Newfoundland, Labrador, pigmentation patterns, Pleistocene refugia, new record

Introduction The geographical distribution of fauna and flora on islands has been of fundamental interest in biology. In addition, islands affected by the Pleistocene glaciers provide opportunities to understand colonization under rapidly changing environments. Pertinent to understanding eastern Canadian freshwater fauna in postglacial environments are studies by Dadswell (1974), Schmidt (1986) and Underhill (1986). Island leech (Hirudinida) fauna were noted by Moore (1922) for Île de la Madeleine (2 species), Gates and Moore (1970) for Sable Island (3 species), Richardson (1943) for Prince Edward Island (8 species), Davies (1979) for Île d’Anticosti (4 species), and Pawlowski (1948) for St-Pierre and Miquelon (2 species). The largest of the islands in eastern Canada is Newfoundland (Newfoundland Island) where 8 species of leeches were noted (Pawlowski 1948, Davies 1973, 1979). Several leech studies addressed the problems of colonization of the islands. Davies (1979) suggested a passive route by sea currents for Île d’Anticosti colonization. Blanchard (1896) and Richardson (1943) suggested birds as vectors for 5 of the species, as well as 3 species (Haemopidae and Erpobdellidae) colonizing the island by transportation on debris. Since these papers were published, events including sea-level changes (Fairbanks 1989, Peltier 2002), glacial forebulge (Hetherington et al. 2003), and the timing of recessional stages since the Last Glacial Maximum (LGM) about 18000–20000 radiocarbon years before present, taken as

Accepted by B. Sket: 30 Oct 2007; published: 7 Dec. 2007

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1950 BP (Dyke 2004, Dyke and Prest 1987, Grant 1989) have suggested additional mechanisms that would have allowed the transference of leeches to and from islands. Herein we review leech (Hirudinida) distributions in Newfoundland and Labrador based on: the works of Pawlowski (1948), Davies (1973, 1979), Madill (1985), the Canadian Museum of Nature collection in Ottawa, National Museum of Natural History collection in Washington, and the recent survey of leeches (Hirudinida) in 2004 by PH. We describe new pigmentation patterns for Erpobdella punctata (Leidy) and Erpobdella obscura Verrill and add four additional species (Theromyzon sp., Haemopis grandis (Verrill), Haemopis lateromaculata Mathers, and Erpobdella parva (Moore) to the provincial listing. We review the gastropods, fishes, and mammals of Newfoundland to suggest barriers to colonizing Newfoundland; faunal transport by birds; and finally glacial and postglacial events and their effects on leeches, island refugia, and potential migration.

Methods and materials Taxonomy. Leeches belong to the phylum Annelida, class Oligochaeta, and order Hirudinida (Siddall et al. 2001). Field guides and taxonomic treatises have been traditionally based on morphological analysis (Klemm 1985, Sawyer 1986, Davies and Govedich 2001). Recent molecular studies on leeches add new evidence on the taxonomy of this group. In this paper, the authors are following the most recent classification of leeches after Borda and Siddall (2004b) which synthesizes traditional classification, and comparative molecular data. The families of leeches found in Newfoundland and Labrador have all been recently revised by Siddall et al. (2005) (family Glossiphoniidae), Borda and Siddall (2004a) (families Haemopidae, Hirudinidae, Erpobdellidae), Utevsky and Trontelj (2005) (family Hirudinidae), and Oceguera-Figueroa et al. (2003) (family Erpobdellidae). The two leech specimens previously published as Hirudo medicinalis (Blanchard, 1896) are reported at the genus level since H. medicinalis is now split into 3 species, i.e., H. medicinalis, H. orientalis Utevsky et Trontelj 2005 and H. verbana Carena 1820 (Utevsky and Trontelj 2005). Synonyms are also listed to reflect other names which are still widely in use as listed in Davies and Govedich (2001). Note that according to Siddall et al (2005), Glossiphonia complanata (Linnaeus 1758) and Helobdella stagnalis ((Linnaeus 1758) are no longer considered synonyms in North America for Glossiphonia elegans (Verrill) and Helobdella modesta (Verrill). A major assumption of this paper and in previous surveys is that the species in this study are closely related to the species on the mainland of North America. This is reflected in the choice of taxonomic names, pending verification by genetic analysis. Field survey. The aquatic survey for leeches occurred in August 2004 and was preliminary in nature, using highway access in Newfoundland and in Labrador. Latitude and longitude coordinates were determined by Magellan GPS 4000 Satellite Navigator. Figure 1 shows the survey locations and Table 1 gives the geographical information, along with dates and notes. This survey was completed prior to examining museum collections and literature records. The most accessible aquatic habitats, close to roads and recreation areas were preferentially examined, and also had to be “good” leech habitats, which largely consisted of aquatic environments with emergent and submerged aquatic vegetation, rocks or logs on the near-shore substrate. The survey method included examining the underside of stones and logs in the water and hand-picking leeches from the substrates. Of note, leeches were not sampled on their hosts, such as fishes, amphibians, reptiles, or mammals, by such methods. The leeches were relaxed with 10% ethanol, fixed in 10% formalin overnight, and placed in 70% ethanol. Vouchers were placed with the Canadian Museum of Nature (CMN). The west-east collecting route taken by T. Jaczewski and S. Feliksiar in their August-September 1938 survey of 58 aquatic sites in Newfoundland and St-Pierre and Miquelon (Pawlowski 1948) was largely followed by PH in 2004.

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MADILL & HOVINGH

TABLE 1. Locations of surveyed aquatic sites. Latitude

Longitude

Date

Notes

580 39.80'W

Newfoundland 1 2

480 16.23'N 0

48 29.08'N

31-Jul-2004

Northern Feeder drainage; pond,Hwy 1N junction Hwy 404

0

31-Jul-2004

Little Barachois Brook drainage: Barachois Pond;Hwy 1

0

58 15.59'W

3

49 27.27'N

56 10.32'W

01-Aug-2004

Berry Brook drainage, lake; SW Springdale: W. Hwy 1

4

490 24.09'N

560 04.90'W

01-Aug-2004

South Brook drainage, South Pond: S Springdale,W. Hwy 1

5

490 03.86'N

560 04.94'W

01-Aug-2004

Exploits River drainage, Joes Lake:10.5 km N. Junction Hwy 370 and Hwy 1

6

480 56.48'N

550 44.97'W

01-Aug-2004

Exploits River drainage;22 km E. Badger; 5.1 km W Grand Falls, Hwy 1

7

480 56.59'N

550 49.66'W

10-Aug-2004

Exploits River drainage, E.Aspen Brook; 16.1 km E. Badger junction 370

8

490 06.95'N

550 04.95'W

01-Aug-2004

Indian Arm Brook, Junction Lake: 1.6 km E. Hwy 340:Notre Dame Junction

9

490 05.44'N

530 48.42'W

10

0

0

48 29.49'N

03-Aug-2004

8 km NE Indian Bay Brook

0

04-Aug-2004

Sandy Pond, Terra Nova National Park: recreation area

0

59 01.22'W

11

46 43.25'N

53 26.50'W

05-Aug-2004

Avalon Peninsula:pond, bridge:W.Trepassey

12

460 44.24'N

530 19.17'W

05-Aug-2004

Avalon Peninsula:pond. Hwy 10 at outflow, E. Trepassey

13

460 47.79'N

530 04.18'W

14

0

0

47 10.08'N

7

km

W.

Northwest

Brook

05-Aug-2004

Avalon Peninsula:pond, N. Hwy 10; SW Chance Cove P.P.

0

05-Aug-2004

Avalon Peninsula:Lake LaManche P.P.

0

52 53.56'W

15

46 58.65'N

55 17.35'W

08-Aug-2004

Burin Peninsula: 8 km NE Little St.Lawarence, upstream from tides

16

460 57.51'N

550 30.20'W

08-Aug-2004

Burin Peninsula:pond: 1.2 km E Lawn

17

500 30.50'N

570 25.74'W

12-Aug-2004

Northern Peninsula:pond at Hwy 430; 4 km S. River of Ponds

0

18

51 05.75'N

56 04.14'W

13-Aug-2004

Northern Peninsula, Roses Brook;downstream lake from Hwy 432 bridge

19

510 31.72'N

550 41.61'W

13-Aug-2004

Northern Peninsula: Pistolet Bay P.P. swimming lake

0

0

Labrador 20

510 44.32'N

560 33.17'W

15-Aug-2004

Hwy 510, 12 km W Red Bay

21

510 47.12'N

560 24.21'W

22

0

52 09.87'N

15-Aug-2004

Hwy 510, 5 km N. Red Bay

0

15-Aug-2004

Hwy 510, 60 km N. Red Bay, at culvert

0

56 05.43'W

23

53 09.89'N

57 31.87'W

16-Aug-2004

Paradise River drainage;11 km on Hwy 516 from Hwy 520 junction

24

530 23.57'N

570 18.37'W

16-Aug-2004

34 km on Hwy 516 from Hwy 520 junction 0.6 km S. Paradise River

25

530 24.91'N

600 24.16'W

17-Aug-2004

lake at Gosling Park; Goose Bay

0

26

53 05.16'N

61 09.46'W

17-Aug-2004

Churchill River drainage;10 km E. Pans River Hwy 500; 71 km E. Goose Bay

27

530 02.70'N

610 29.14'W

17-Aug-2004

Churchill River drainage;18 km E Bobs Brook: 98 km E Goose Bay Hwy 500

28

530 31.85'N

640 00.76'W

18-Aug-2004

Churchill River drainage, lake at Churchill Falls

29

530 02.20'N

660 20.36'W

18-Aug-2004

Ashuanpi River drainage; 9.6 km W. railroad:lake transversed by Hwy 500

0

0

NEWFOUNDLAND LEECHES

Zootaxa 1657 © 2007 Magnolia Press ·

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Previous records. Leeches from Newfoundland and Labrador were examined from the Annelida Collection of the Canadian Museum of Nature (CMNA) and United States National Museum of Natural History (USNM) (Table 2). Identification of the leech species were confirmed at these museums. Latitude and longitude, if recorded, is listed in Table 2. Of note, Haemopis grandis (USNM38612) and Erpobdella parva (USNM38582) are listed for Fox Island. The location shown on Figure 3D is Fox Island, east of Boat Harbor near the Burin Peninsula. This island is the only Fox Island on the Official Newfoundland and Labrador Province highway map. However, 20 Fox Islands are listed for Newfoundland and Labrador in the Gazeteer of Canada (Dept. Energy Mines & Resources 1968). TABLE 2. Locations of museum records from Newfoundland and Labrador. Species

Catalogue No.

Latitude

Longitude Collector

Date

Location

Newfoundland: Glossiphonia elegans

CMNA 19870467

470 34'N

520 44'W

F. Johansen 15-Aug-1922

St. John's, Burton's Pond

Helobdella modesta

CMNA 19810620

500 42.9'N

570 22.4'W

P. Frank, I. Sutherland

Point Riche Peninsula, Port au Choix Bay, South of bay, outlet stream of pond

Helobdella modesta

CMNA 19870485

470 34'N

520 44'W


St. John’s, Burton’s Pond, stones

Helobdella modesta

CMNA 19870486

470 34'N

520 40'W


St. John’s, Signal Hill, pond

Helobdella modesta

CMNA 19870487

470 35'N

520 45'W


pond in field, north of St. John’s

Helobdella modesta

CMNA 19870494

480 30'N

540 15'W

G. Sharp

3-Jun-1967

Terra Nova National Park, Bonne Bay

Helobdella modesta

CMNA 19870500

480 49'N

570 59'W

E.L. Bousfield

11-Jul-1954

below Pinchgut L., Harry’s River, small lake, marl, wood debris

Haemopis lat- CMNA 1982eromaculata 0654

470 03'N

540 07'W

C. McGrath 15-Nov-1981

Placentia Bay, Patrick's Cove, Patricia Bay

Haemopis lat- CMNA 1982eromaculata 0378

460 49'N

540 05'W

D.G.Walsch 8-Nov-1985

Placentia Bay, Point Lance

R. Bere

(Burin Peninsula, East of Boat Harbor), Fox Island, Rocky Shore

8-Aug-1979

Haemopis grandis

USMN38612

Erpobdella punctata

CMNA 19780344

Erpobdella parva

USNM38582

Erpobdella obscura

CMNA 19005994

490 00'N

570 20'W

F. Johensen 31-Aug-1922

Grand Lake, creek on west side of lake

Glossiphonia elegans

CMNA 19880126

540 33'N

660 30'W

D.R. Oliver 1-Sep-1957

Labrador, Astray Lake, 12 m

Erpobdella obscura

CMNA 19880127

540 33'N

660 30'W

D.R. Oliver 10-Jul-1957

Labrador, Astray Lake, 12 m

480 30'N

530 05'W

3-Aug-1927

12-Jul-1976 A.H. Clarke, J. J. Clarke

Port Union Pond

R. Bere

(Burin Peninsula, East of Boat Harbor), Fox Island, Rocky Shore

3-Aug-1927

Labrador:


MADILL & HOVINGH

Maps. The Grand Banks of Newfoundland were mapped to depict post-glacial events using marine maps from Fisheries and Oceans, Canada (Map 4020, Strait of Belle Isle 1: 150,000; Map 4025, Cap Whittle to Havre-Saint-Pierre and Île d’Anticosti 1: 300,000; Map 4026, Havre-Saint-Pierre and Cap des Rosier to Pointe des Monts 1: 300,000), and U.S. Department of Defense, Defense Mapping Agency Hydrographic / Topographic Center (Map 14024, Island of Newfoundland; Map 14010, Gulf of St. Lawrence; Map 14002, Northumberland Strait; Map 14009, Cape Race to Cape Sable; Map 14003, Cape Henry to Cape Race). Maps showing times of glacial recession were adapted from Dyke (2004) and Occhietti et al. (2004).

FIGURE 1. Surveyed sites in Newfoundland and Labrador. Circles, 2004 survey sites (numbered); Squares, literature locations of Pawlowski (1948) and in part, Davies (1973). The site numbers correspond to Table 1.



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Photo manipulation. Slide transparencies taken by Peter Frank of live leeches from Grand River, Ontario in 1985 were scanned in Adobe Photoshop. Digital images of preserved specimens in the Canadian Museum of Nature Annelida collection were captured using ACT1, using a Nikon EXE1200F digital camera mounted on an Olympus SZX1212 stereomicroscope.

Results Leeches from Newfoundland and Labrador and description of some unique pigmentation forms, are listed below. Leeches from the 2005 survey are illustrated in Figure 2. Sources for common names are personal correspondence with K.A. Coates and M. Wetzel in 1988, and M. Siddall (2006).

Phylum Annelida Order Hirudinida Family Glossiphoniidae Alboglossiphonia heteroclita (Linnaeus 1761) Common name: White leech General distribution: Holarctic (Figure 3A). Newfoundland: Literature: Pawlowski (1948) at Deer Lake.

Glossiphonia elegans (Verrill 1872) Common names: Brook leech or snail leech General distribution: Holarctic (Figures 2A, 4A, Tables 1,2). Newfoundland: Survey: Site 1 (CMNA 2006–0002), Site 10 (CMNA 2006–0005), Site 14 (CMNA 2006–0008), Site 17 (CMNA 2006–0012), and Site 18 (CMNA 2006–0013); Museum specimens: CMNA 1987–0467; Literature: Pawlowski (1948) as Glossiphonia complanata at Deer Lake; Pickavance (1971) as Glossiphonia sp. in Rennies River, St. John’s; Davies (1973) as G. complanata at Rocky Harbour. Labrador: Museum specimens: CMNA 1988–0126. The record by Pickavance is questionable since Glossiphonia heteroclita was a name sometimes used for Alboglossiphonia heteroclita.

Helobdella modesta (Verrill 1872) Common name: Scutate snail leech General distribution: Holarctic (Figures 2B, 4B, Tables 1, 2). Newfoundland: Survey: Site 1 (CMNA 2006–0001), Site 10 (CMNA 2006–0004), Site 12 (CMNA 2006–0006), Site 14 (CMNA 2006–0007), Site 15 (CMNA 2006–0009), Site 16 (CMNA 2006–0010), Site 17 (CMNA 2006–0011), Site 19 (CMNA 2006–0014). Museum specimens: CMNA 1981–0620, CMNA 1987– 0485, CMNA 1987–0486, CMNA 1987–0487, CMNA 1987–0494, CMNA 1987–0500; Literature: Pawlowski (1948) as Helobdella stagnalis at St George’s (two stations), Deer Lake, Grand Falls; Gates and


MADILL & HOVINGH

Moore (1970) as H. stagnalis at Lily Pond; Pickavance (1971) as Helobdella sp. at Rennies River, St. John’s; Davies (1973) as H. stagnalis at Rocky Harbour and at Bonne Bay, Big Pond, and Gull Lake [There is a Big Bonne Bay Pond NNE of Deer Lake (49º 30' N, 570 55' W), Gull Lake (49º 39' N, 56º 57' W), and Big Pond just NE of Colinet and S of Ocean Pond (47º 31' N, 53º 28' W). Labrador: Survey: Site 24 (CMNA 2006–0015). St-Pierre and Miquelon: Literature: Pawlowski (1948) as Helobdella sp.: St-Pierre.

FIGURE 2. Photographs of selected Newfoundland and Labrador leeches. D was collected by Cecelia McGrath in 1982. A, Dorsal view of Glossiphonia elegans (Verrill), site 17; B, Dorsal view of Helobdella modesta (Verrill), site 12; C, Dorsal view of Theromyzon sp., site 6; D, Lateral view of Haemopis lateromaculata Mathers, showing the plain ventrum separated from the speckled dorsum by a lighter pattern on the side, CMNA 1982–0654; E, Dorsal view of Haemopis marmorata (Say), site 12; F, Closeup of anal protrusion of Haemopis marmorata, Site 12; G, Erpobdella punctata (Leidy) (normal pigmentation), site 7; H, Erpobdella obscura (Verrill) [normal pigmentation], site 21. Scale bars are in cm unless otherwise indicated. NEWFOUNDLAND LEECHES


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FIGURE 3. Distribution of rare leeches. Circles, 2004 survey; squares, literature locations (Pawlowski 1948, Davies 1973); triangles, museum specimens. A, Alboglossiphonia heteroclita (Linnaeus); B, Theromyzon sp.; C, Haemopis marmorata (Say): circles and squares; Haemopis lateromaculata Mathers: triangles; D, Haemopis grandis (Verrill) and Erpobdella parva (Moore): both museum specimens found at the same site (Fox Island- see text) .

Theromyzon sp. Common names: Waterfowl leech or duck leech General distribution: Holarctic (Figures 2C, 3B, Table 1).


MADILL & HOVINGH

Newfoundland: Museum specimen: Site 6 (CMNA 2006–0003). The species is undetermined since the gonopore separation could not be deciphered. Maculate pigmentation, especially evident around the dorsal edge. Previously unreported for Newfoundland and Labrador. This taxon is unusually rare for a region with an abundance of waterfowl.

Family Haemopidae Haemopis grandis (Verrill 1874) Synonym: Mollibdella grandis (Verrill 1874)

General distribution: Eastern North America (Figure 3D, Table 2). Newfoundland: Museum specimens: USNM 38612: Fox Island (1927, station 15, accession number 259718, collector Ruby Bere). (See note about Fox Island in Previous Records section.) Notes on the species.This species is identified by the characteristic pharyngeal musculature associated with the lack of teeth and jaws (Richardson 1969). A specimen was dissected to examine the internal organs. Haemopis grandis is previously unreported in Newfoundland and Labrador.

Haemopis lateromaculata Mathers 1963 Synonym: Percymoorensis lateromaculata (Mathers 1963)

General distribution: Nearctic (Figures 2D, 3C, Table 2). Newfoundland: Museum specimens (identified by JM): CMNA 1982–0654: Patrick Cove (Placentia Bay); CMNA 1985–0378: Point Lance at St. Mary’s Bay. Both records are on the Avalon Peninsula. Previously unreported for Canada, these records are the first outside of Iowa and Minnesota (Mathers 1963). Notes on the species. The unusual conditions under which Haemopis lateromaculata was found follows: On November 15, 1981, Cecelia McGrath in Patrick Cove noticed large, black worms heading across her lawn. R. Morris, Agriculture Canada, described the animals as “5–7 cm long, olive dorsally with small, black irregular flecks,…ventrally black, sucker large, and light, yellow, longitudinal stripes.” On November 8, 1985, D.G. Walsch found another H. lateromaculata in a freshwater well in Point Lance. All species of Haemopidae are amphibious, i.e., they can swim and crawl in aquatic environments, and when conditions are right, they can travel overland. The skin of leeches is moist and absorbent, and needs wet conditions such as dew-drenched surroundings, or rainfall to avoid desiccation. In both records, H. lateromaculata was collected in the late fall. These Haemopis most likely were looking for suitable winter quarters just as Moore (1923) described for another bloodsucker species, Macrobdella decora. Ms. McGrath claimed that these animals had never been near her property previously and must have traveled a fair distance overland because their original freshwater habitat was nowhere in sight. Dorsally, Haemopis lateromaculata resembles the most common species of horse-leech (Haemopis marmorata (Say)) since its dorsum is dark and mottled. However, the two lateral yellow stripes, and the uniformly plain ventrum are diagnostic of living H. lateromaculata. Preserved specimens can also be identified by dissection and microscope examination. The male and female gonopores are 5 annuli apart, and in the furrows between the annuli. According to Mathers (1963) and Hovingh (pers. observation 2006), H. lateromaculata can be distinguished from H. marmorata by comparing the relative size, shape, and orientation of internal organs. For example, H. lateromaculata has a U-shaped male penis sheath (Mathers 1963) and H. marmorata has a longer J-shaped penis sheath (Klemm 1985).



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Haemopis marmorata (Say 1824) Synonym: Percymoorensis marmorata (Say 1824)

Common name: American horse-leech General distribution: Nearctic (Figures 2E, 3C, Table 1). Newfoundland: Survey: site 17 (CMNA 2006–0024), identified by the presence of teeth and the color of the ventrum which is characteristically mottled or spotted but lighter than the dorsum. No lateral stripe. Stomach contents of CMNA 2006–0024 included 1 juvenile erpobdellid leech, and Cladocera (Bosmina sp.); Literature: Blanchard (1896) 3 specimens from Newfoundland; Pawlowski (1948) at Corner Brook and Deer Lake; Gates and Moore (1970) from Lily Pond. St-Pierre and Miquelon Islands: Literature: Blanchard (1896): St-Pierre (21 specimens); Miquelon (34 specimens). Notes on the species. Pawlowski (1948) discusses Blanchard’s (1896) report of Haemopis collected by Leguillou in 1838 from an unknown location on Newfoundland, as well as a collection by Dr. Kermorganat in St-Pierre and Miquelon. Blanchard identified these specimens as the Eurasian Haemopis sanguisuga Linnaeus but Pawlowski identified Blanchard’s specimens as typical North American Haemopis marmorata. Blanchard (1896) pointed out that prolapsus of the rectum in the Newfoundland specimen was never before noticed in Europe. Pawlowski (1948) agrees, and notes that this condition occurs rarely but only in North America. The closeup of specimen CMNA 2006–0024 illustrates this condition (Figure 2F). JM also re-examined Blanchard’s specimens from the Musée National d’Histoire Naturelle in Paris in 1982 which have become dessicated and hard, and unsuitable for further comment.

Family Hirudinidae Hirudo sp. Common Name: European medicinal leech General distribution: Eurasian Newfoundland: Literature: Blanchard (1896) reports two specimens in the Paris Musée Nationale d’Histoire Naturelle from Newfoundland, location and date missing, and suggests that this species was introduced intentionally by man. No known resident population has been found since 1896 (Davies 1973, Madill 1985). The specimens reside in vial 149 at the Paris Musée Nationale d’Histoire Naturelle.

Family Erpobdellidae Erpobdella parva (Moore 1912) Synonym: Dina parva Moore 1912

General distribution: Nearctic (Figure 3D, Table 2). Newfoundland: Museum specimen: USNM 38582: Fox Island, Rocky Shore (1927, station number 15, accession number 259718, collector Ruby Bere). (See note about Fox Island in Previous Records section.) Identified by the presence of two pairs of labial eyes and the gonopores are separated by 4 or 3 ½ annuli.


MADILL & HOVINGH

Erpobdella punctata (Leidy 1870) Common name: Tiger leech General distribution: Nearctic (Figures 2G, 4C, 5, Tables 1, 2). Newfoundland: Survey: Site 1 (CMNA 2006–0025), Site 2 (CMNA 2006–0026), Site 7 (CMNA 2006– 0027), Site 10 (CMNA 2006–0028), Site 12 (CMNA 2006–0029), Site 13 (CMNA 2006–0030), Site 14 (CMNA 2006–0031), and Site 15 (CMNA 2006–0032); Museum specimen: CMNA 1978–0344; Literature: Pawlowski (1948) at Corner Brook, Deer Lake, and Terra Nova; Gates and Moore (1970) at Lily Pond; Pickavance (1971) as Erpobdella sp. in Rennies River, St. John’s; Davies (1973) at Rocky Harbour. Labrador: Survey: Site 24 (CMNA 2006–0033); St. Pierre and Miquelon: Literature: Pawlowski (1948) at two stations. Notes on the species. Figure 5 shows the pigmentation patterns of E. punctata in Newfoundland. Specimens of E. punctata from Sites 14 (Avalon Peninsula) and 15 (Burin Peninsula) had a distinct pigmentation pattern not previously noted in the Great Basin (Hovingh 2004) or illustrated by Klemm (1985: Fig.7.97). Some specimens (Figure 5C) possessed an inner para-medial row which combine across the median and provide an appearance of a single broad dorsal-median strip (the “mid-dorsal form”). The variations at Sites 1 and 2 (Figure 5F) were less developed across the medium. Klemm (1985: Figure 7.97) shows another variation having inner and outer para-medial rows united with a lighter mid-dorsal region. Sometimes the entire annulus is pigmented across the medium showing a dorsum that is very dark and with no mid-dorsal region. At site 7 (Figure 5B), specimens most closely resembled the ‘normal’ variety. Specimens at other sites were intermediary between the two forms (Figure 5A, D, E).

Erpobdella fervida (Verrill 1781) Synonym: Mooreobdella fervida (Verrill 1871)

General distribution: Eastern North America. Newfoundland: Literature: Davies (1973) at Gull Pond. The Official Highway Map of Newfoundland and Labrador shows five Gull Ponds on Newfoundland: in N-central between Badger and South Brook, and W of Kings Point; in NE W of Glovertown, and N of Gambo; and in the SE in St. Joseph.

Erpobdella obscura (Verrill 1872) Synonym: Nephelopsis obscura Verrill 1872

Common name: Bait leech General distribution: Nearctic (Figures 2H, 4D, 6, 7, Tables 1, 2). Newfoundland: Survey: Site 7 (CMNA 2006–0023), Site 18 (CMNA 2006–0022); Museum specimen: CMNA 1900–5994; Literature: Pawlowski (1948) at Grand Falls. Labrador: Survey: Site 20 (CMNA 2006–0021), Site 21 (CMNA 2006–0020), Site 22 (CMNA 2006– 0019), Site 23 (CMNA 2006–0018), Site 28 (CMNA 2006–0017), and Site 29 (CMNA 2006–0016); Museum specimen: CMNA 1988–0127 (Figure 4D). Notes on the species. Verrill’s (1872) description of Erpobdella obscura strongly resembles some of the Newfoundland and Labrador specimens but there is also a previously unpublished color variation. Klemm (1985) describes the normal appearance of Erpobdella obscura (Figures 6F, 7B, E, I, J) as “color variable, dorsum greenish-brown, covered with sparse scattered black or light colored blotches, interlacing or irregular NEWFOUNDLAND LEECHES


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spots, or plain (uniform), no striping…” In Newfoundland and Labrador, we found Erpobdella obscura, with a pattern which we call ‘ringed’ because of horizontal dark pigment in the furrows between the annuli (Figure 7A, D, F–H). These are remarkably similar to other ‘ringed’ specimens collected by J. Metcalfe-Smith in the Grand River, Ontario (north of Kitchener) at Winterbourne, Glen Morris, Upper Bellwood, and Elora Gorge (Figure 6, Table 3).

FIGURE 4. Distribution of the four widespread leech species. Circles, 2004 survey; squares, literature locations (Pawlowski 1948, Davies 1973); triangles, museum specimens. A, Glossiphonia elegans (Verrill); B, Helobdella modesta (Verrill); C, Erpobdella punctata (Leidy); D, Erpobdella obscura (Verrill).


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FIGURE 5. Pigmentation patterns in Newfoundland and Labrador Erpobdella punctata (Leidy) by PH. A–B, Site 7, C Sites 14–15, D, Sites 12, 13, 24; E, Site 10; F, Sites 1–2.

TABLE 3. Normal and ‘Ringed’ specimens of Erpobdella obscura (Verrill) from the Grand River, Ontario, collected by J. Metcalfe-Smith, CCIW, and from Newfoundland and Labrador. Location

Latitude

Longitude

Catalogue Number Normal

Catalogue Number ‘Ringed’

Upper Lake Bellwood, Grand River, 430 49.83’N Ontario

800 18.00’W

CMNA 1983–0292 CMNA 1983–0293

CMNA 1984–0689 CMNA 1986–0125

Elora Gorge, Grand River, Ontario

430 39.75’N

800 27.00’W

Winterbourne, Grand River, Ontario 430 33.60’N

800 28.60’W

CMNA 1986–0114

CMNA 1983–0294 CMNA 1986–0117

Glen Morris, Grand River, Ontario

430 16’.70’N

800 20.93’W

CMNA 1984–0693 CMNA 1984–0697 CMNA 1986–0122

CMNA 1983–0295 CMNA 1984–0690 CMNA 1984–0694 CMNA 1984–0695 CMNA 1986–0121

Newfoundland

(see text)

(see text)

CMNA 2006–0022

CMNA 2006–0023

Labrador

(see text)

(see text)

CMNA 2006–0020 CMNA 2006–0019 CMNA 2006–0017

CMNA 2006–0017 CMNA 2006–0016

CMNA 1986–0131

The anatomy of ‘ringed’ forms was the same as normal specimens (Figure 6). Verrill (1874) had described the large mouth (Figure 6A), the presence of 4 pairs of eyes (2 labial and 2 buccal) (Figure 6B), the separation of the gonopores by 2 annuli (Figure 6D), the partial subdivision of some annuli (Figures 6B, D, E, 7G, J), the large, raised anal orifice (Figure 6E), and the spiraling of the ejaculatory ducts of Erpobdella obscura exiting the atrial cornua (Figure 6C, G). The anatomy of both varieties was very similar in these respects. According to Sawyer (1972) the most distinguishable feature of the species is subdivision of the annuli. In Ontario, The normal variety co-existed in the same locations as ‘ringed’ specimens (Table 3). Intermediate forms were also noted. High magnification reveals that normal specimens have anastomosing patches of dark color dispersed all over the dermis, and the furrows of the annuli are colored (Figure 7I, J). The patterns were different in ‘ringed’ specimens, with pigment restricted to the furrows of the annuli, as well as between subdivisions of the annuli (Figure 7G, H). Colors of live ‘ringed’ specimens in Ontario were recorded using the Naturalist’s Colour Guide (Smithe 1975). Background color varies between clay (color 26); smoke gray (color 45); buff (color 124); and tawny olive (color 223D). The rings were either black, plumbeous (color 78) or glaucous (color 79) respectively. The stomach contents of the ‘ringed’ specimens were analyzed by J. Met-



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calfe-Smith in May, and July, 1983, and included mostly insect larvae: chironomids, simulids, diptereans, ephemeropterans, plus 1 ostracod. In addition to the 4 specimens in Newfoundland and 14 specimens in Labrador, another 150 specimens in Ontario were examined (Figures 6, 7). Mean length of preserved ‘ringed’ specimens were consistently longer, as follows: Ontario ‘ringed’ 3.61 cm, normal 2.81 cm; Newfoundland ‘ringed’ 7.41 cm, normal 3.89 cm; Labrador ‘ringed’ 6.68 cm, normal 3.75 cm, and intermediate forms 5.01cm.

FIGURE 6. Erpobdella obscura (Verrill) from Grand river, Ontario. A–E, G: Drawings by Susan Laurie-Bourque of CMNA 1984–0689. F by Peter Frank using Kodachrome film in May 1988. A, Anterior, ventral view; B, Anterior, dorsal view; C, Closeup of seminal vesicle; D, Well-developed male and female gonopores located between annuli; E, Posterior dorsal view; F, Photographs of two preserved leeches comparing the two color variations (ringed form on left and normal form on right); G, Dorsal view of dissected reproductive organs. The right side of the testes and spermduct are omitted on the right side. Broken lines show cut ends of organs. a, atrium; ejd, ejaculatory duct; ep, epididymis; fg, female gonopores; mg, male gonopores; lft, large follicles of testisacs; t, testisac; mft, multifollicular testisacs; nc, nerve chord. Ganglia are numbered in Roman numerals.


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FIGURE 7. Pigmentation patterns of Erpobdella obscura (Verrill). A–F are illustrations by PH from Labrador and Newfoundland: A, Site 28; B, Sites 22, 23; C, Site 21; D, Site 29; E, Site 18; F, Site 7. G–J are microscope photographs of typical examples of ringed and mottled patterns of Erpobdella obscura (Verrill) from Newfoundland and Labrador compared to specimens from the Grand River, Ontario. Scale bars are in mm. G, closeup of midportion of ringed specimen from Newfoundland, CMNA 2006–0023. H, anterior portion of ringed specimen from Newfoundland, CMNA 2006– 0020. I, anterior portion of mottled specimen from Newfoundland, CMNA 2006–0018. J, closeup of midportion of mottled specimen from Newfoundland, CMCA2006–0018.

In Newfoundland and Labrador, some Erpobdella obscura specimens had normal pigmentation patterns as figured by Klemm (1985: Figure 7.103) such as those found in sites 18 and 21 (Figure 7C). Specimens from sites 7 and 29 were unique ‘ringed’ forms (dark interannuli) (Figure 7D, F). Secondary intra-annular segmentation occurred in two to four adjacent annuli per segment which were also very dark, and extended partly into the ventrum. Both ‘ringed’ and normal specimens cohabited at site 28. Intermediate forms occurred at sites 22, 23, and 28 with both spots and rings (Figure 7A, B). The ‘ringed’ form has not been found in the Great Basin (Hovingh 2004) or noted by Klemm (1985). Survey Results. Pawlowski (1948) noted that of the 58 stations surveyed in Newfoundland, leeches were found in only 14 sites (24%). In the 2004 survey which was only concerned with leech distribution, 19 sites in Newfoundland and 10 sites in Labrador were surveyed. Leeches were found at 13 sites (68%) in Newfoundland Island and in 7 sites (70%) in Labrador. The success of the 2004 survey is attributed to selecting sites with abundant rocks and logs and vegetative growth which looked “good” for leech collecting. Five species were common to both surveys, with Alboglossiphonia and Theromyzon found in one of the surveys. No Piscicolidae (fish leeches) were caught in any of the surveys, but there is a chance that piscicolid leeches may exist in the study area since the preferred method of collection for this group is to look for these leeches on their host fishes. A casual mention of Macrobdella decora in Labrador (Moore 1923) is not substantiated in any literature records, nor was this species found in any surveys. Also, the single report of Hirudo (Blanchard 1896) from Newfoundland Island suggests that this species does not reproduce in Newfoundland. Newfoundland and Labrador leeches include the widely distributed and common leeches Glossiphonia elegans, Helobdella modesta, Erpobdella punctata, and Erpobdella obscura. These were the only leeches NEWFOUNDLAND LEECHES


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found in Labrador. Two widespread taxa in North America were found at one location each: Theromyzon, a waterfowl leech, and Erpobdella parva, suggesting a confinement of movement on Newfoundland. Haemopis marmorata was found at 3 sites by surveys and H. grandis and H. lateromaculata were noted only in museum collections. Future Haemopis surveys with baited traps would assist in determining the ranges of these 3 species. Alboglossiphonia heteroclita and Erpobdella fervida were only noted in literature from 1 site each.

FIGURE 8. Postglacial events that explain the presence of leeches in Newfoundland and Labrador, adapted from Dyke (2004) and Occhietti et al. (2004). Panel A . Barrier events: the Laurentian Channel and the northward Esquiman Channel (unlabeled) with its 200 meter depth contour; #1, the Strait of Belle Isle with glacial and marine blockage and possible land bridge; #2, the Jacques Cartier Passage with glacial and marine blockage and possible land bridge; #3, North Shore of the Gulf of St. Lawrence ice sheet blockage; #4, St. Lawrence River glacial blockage. Late Glacial Maximum refugia include A, St Pierre Bank (38–60 m below sea level); B, Green Bank(60–80 m below sea level); and C, Grand Bank (70–75 m below sea level) off the southern coast of Newfoundland. The shaded areas are land when the minimum sea level was at -120 m during the Last Glacial Maximum and the heavy solid line is the 200 m contour of the present sea level. The ice sheets are denoted by a heavy dotted line at 18,000 years BP (not labeled) associated in part with the -200 m contour; the thick dashed line on the Grand Banks is the limit of a 14,000 years BP glacial lobe (unlabeled); the dashed line at 11,000 years BP, and the thin dashed line at years 9000 years BP (both labeled). Light dots denote provincial boundaries. Panel B . Scheme showing hypothetical movement of leeches through time in Newfoundland and Labrador regions: (1) Colonization of Newfoundland from Labrador before 70,000 years BP; (2) colonization of the Grand Banks from Newfoundland after 50,000 years BP; (3) colonization of Newfoundland from the Grand Banks 13,000 to 11,000 years BP; (4) the return colonization of Labrador and the North Shore 11,000 to 10,000 years BP; (5) and the colonization of Île d’Anticosti from the North shore and the colonization of Île de la Madeleine from Prince Edward Island (PE) after 10,000 years BP; and (6) the colonization of Sable Island from Nova Scotia.

Discussion Barriers to fauna movement. 11 species of leeches are present in Newfoundland Island, 4 species in Labrador, while 26 species occur in Québec (Klemm 1985). A reduction in the number of species in Labrador suggests major paleo-barrier events took place when leeches colonized Labrador. Mammals exhibit a different pattern with 32 to 38 species (exclusive of bats and marine mammals) occurring in the Gulf of St. Lawrence (Labrador inclusive), yet only 13 mammals (mostly carnivores) occur on


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Newfoundland Island (Banfield 1974). If rodents alone are considered, only 3 rodents occur on Newfoundland Island while 11 species occur in the adjacent provinces and Labrador. These 3 microtine rodents (beaver Castor canadensis, muskrat Ondatra zibethica, and meadow vole Microtis pennsylvanica) are confined to semiaquatic habitats. Of the 14 species of primary freshwater fish in the freshwater drainages of the Gulf of St. Lawrence, only 1 fish species is found in Newfoundland and 9 species are found in Labrador (Scott and Crossman 1973, Schmidt 1986, and Underhill 1986). Thus, each faunal group has responded differently to paleo-environments. In the colonization of Newfoundland Island, there is a major barrier to movement of mammals and fish from Labrador to Newfoundland Island and another major barrier to movement of leeches from Québec and Newfoundland Island to Labrador. No amphibians or reptiles occurred naturally on Newfoundland Island. Passive movement of fauna by birds. Mobility of aquatic fauna has often been attributed to incidental movement by birds. Active movement through habitat has been postulated for fishes (Hubbs and Miller 1948) and mollusks (Taylor and Bright 1987, Hovingh 1993) although movement of mollusks by birds may be invoked to explain data that is not interpretable by other hypotheses. Siddall et al. (2005) suggested that migrating birds transporting incidental mollusks could also spread species of Helobdella between North and South America while the leeches were feeding internally on soft tissues of mollusks. A distance of 120 km between Newfoundland and Cape Breton Island may exceed the potential capabilities of passive migration of gastropods by birds. Malone (1965) found Promenetus exacuous (Say) could survive five hours out of water and the young could remain attached to a Killdeer (Charadrius vociferous Linnaeus) leg indefinitely. This gastropod occurred in the Maritime Provinces but not in Newfoundland and Labrador (Clark 1981). Boag (1986) found that 18% of Stagnicola elodes (Say) (found in the Maritime Provinces and Newfoundland) survived on a feather after 15 minutes in the air. The measured flight speed of waterfowl is 60 km/h (Tucker and Schmidt-Koenig 1971) or 2 hours to get from Cape Breton to Newfoundland. Gastropod passive mobility by birds can be compared to the blood-sucking Theromyzon which actively seeks out waterfowl and spends 2.5 hours feeding in the nasal passage of birds (Davies and Wilkialis 1981). Each nasal passage may have multiple leeches (Bartonek and Trauger 1975). This would readily allow Theromyzon transport of 150 km, more than necessary to reach Newfoundland across Cabot Strait from the Maritime Provinces. This type of transportation of gastropods and leeches would require the bird acting as a vector to settle in suitable freshwater habitat after the flight. Glacial and post-glacial events and their effects on leeches. The Pleistocene occurrences of glaciers had an immense impact on the flora and fauna of North America (Rogerson 1983, Pielou 1991). During the Last Glacial Maximum, most of Canada was covered by ice (Dyke 2004). The Laurentide Ice Sheet was centered in eastern Canada and the adjacent Appalachian Glacier Complex consisted of many local and converging ice centers in New England, Gaspésie, and Maritime Provinces. Newfoundland had its own ice centers. The presence of freshwater leeches on islands associated with the Gulf of Lawrence can be explained by: (1) their presence on these islands since Illinoian glacier (190,000–130,000 years BP) and survival in refugia; (2) their presence as a result of occupation during the early Wisconsinan glacier (70,000 years BP) and survival in Late Pleistocene refugia; or (3) their presence as a result of postglacial colonization 15,000 to 10,000 years BP. Key events in the Late Pleistocene affecting Newfoundland Island are illustrated in Figure 8A. Major barrier events included differential recession of glaciers (Catto 1998, Dyke 2004, Occhietti et al. 2004, Shaw 2003, Stea et al. 1998) with a prominent blockage of movement into eastern Québec and Labrador (Figure 8A: #3). At 18,000 years BP Fairbanks (1989) and Peltier (2002) suggested that the sea level was 120 m lower than today and the Grand Banks were exposed and connected to Newfoundland (Fig 8A: A,B,C). These coastal areas are proposed as refugia for leeches.



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The Laurentian Channel formed in the mid-Pleistocene (Stea et al. 1998). As the sea levels rose from 13,000–15,000 years BP, only St-Pierre Bank was emergent, allowing the transfer of aquatic fauna to Burin Peninsula (Figure 8B: route 3). Until 10,000 years BP, the Laurentide Ice persisted in the North Shore (Figure 8A: #3) and blocked movement of aquatic fauna between western Québec and Labrador. During minimum sea levels, a land connection between Newfoundland and Labrador was covered with glacial ice (Figure 8A: #1), and as the glaciers receded, the isostatic rebound and the forebulge resulted in aerial exposure of this connection, allowing for fauna exchange between 11,000 and 10,000 years BP (Figure 8B: route 4) (Fairbanks 1989, Grant 1989, Hetherington et al. 2003, Painchaud et al. 1984, Peltier 2002). The freshening of the Gulf of St. Lawrence by removal of the St Lawrence River glacial blockage (#4, Figure 8A) and episodic events (10,600–9200 years BP) associated with Lake Agassiz outlets to the St Lawrence River (Dynesius and Nilsson 1994, Rodrigues and Vilks 1994, Teller et al. 2002) might have allowed movement of leeches if the water was less than 16% sea water (Reynoldson and Davies 1976). Based on this geological and geographical knowledge, we propose that the present occupation of leeches on Newfoundland Island is a result of postglacial colonization from off-shore refugia associated with the Grand Banks and not from the adjacent areas of Québec and Labrador. Figure 8B shows a hypothetical timing of leech movement in the Newfoundland Island region. South of Laurentian Channel and coastal to the Maritime Provinces and New England, the Northeastern refugium of Pleistocene subaerial banks may have been occupied by approximately 20 species of primary freshwater fish (Schmidt 1986), and by mammoths and other fauna on the Georges Bank (Whitmore et al. 1967, Cooke et al. 1993). Leeches (Helobdella modesta, Haemopis marmorata, and Erpobdella punctata) presently found on Sable Island (Gates and Moore 1970), probably occupied the Scotian Shelf. Two species of leeches (H. marmorata and E. punctata) have been found on Île de la Madeleine (Moore 1922). These occurrences could be explained by similar processes as those described for Newfoundland. Of note, ice disappeared from Prince Edward Island about10,000 years BP with land connections to New Brunswick and Nova Scotia until 5000 years BP, allowing 8 species of leeches to colonize that “island”. Analysis of Pleistocene and Holocene events provides an alternative to transportation of fauna by birds and floating debris. The above discussion of barriers, the mobility of fauna by avian vectors, and the post-glacial migration of Newfoundland and Labrador leech species from the Maritime Provinces yields insight into the geographical isolation of Newfoundland, St. Pierre, Miquelon, and Île de la Madeleine Islands. The molecular evidence of Borda and Siddall (2004b), Light and Siddall (1999), and Siddall et al. (2005) has redefined the taxonomy of leeches. Despite a lack of morphological differentiation, Siddall et al (2005) consider the North American populations of Helobdella stagnalis (Linneaus 1758) and Glossiphonia complanata (Linnaeus 1758) to be species separate from their European counterparts based upon genetic analyses. Such species assignments may not be warranted due to the possibility that many population intergrades may exist from western Europe to eastern North America, through eastern Europe, Asia, and western North America, or through Greenland and Iceland. It would be most important to know whether American, European or other populations colonized Newfoundland Island and Labrador. We strongly believe that future research include analysis of mitochondrial DNA of leeches in Iceland, Greenland, and Newfoundland as well as across both the Asian and North American continents.

Conclusions Literature records, museum vouchers, and a survey of the Province of Newfoundland and Labrador leeches confirm the presence of 11 species for this province. In addition, new pigmentation forms of Erpobdella punctata and Erpobdella obscura are described. By examining the faunal barriers among geographic areas, assessing the transport by birds, and examining the environments of the Last Glacial Maximum and the recession of


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ice from that stage, we propose that the leeches of Newfoundland occurred in a Pleistocene refugium on the Grand Banks, recolonized Newfoundland as conditions permitted, and thereafter recolonized Labrador. Our model no doubt will be greatly modified as more leech surveys occur and more geological information is made available.

Acknowledgments We thank Bill Moser (USNM) and Jean-Marc Gagnon (CMN) and greatly appreciate the use of their facilities in examining the leeches, and Donald J. Klemm, (USEPA) for examining numerous leeches in this study. We thank the staff of CMN for their support during the project, especially Joanne Dicosimo, Dr. Mark Graham, Roger D. Bull, Dr. Kathleen Conlan, Peter Frank, Dr. Richard Harington, Ed Hendrycks, Dr. André Martel, Christine McClelland, and Dr. Claude Renaud. We also thank Lucie Jamieson, Susan Laurie-Bourque, Cecelia McGrath, Janice Metcalfe-Smith (Environment Canada), R. Morris (Agriculture Canada), and Dr. Andrew Smith for their contributions. We are grateful to John Gosse, and David Cote (Terra Nova National Park, Environment Canada).

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