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FIGURE 5. Placobdella akahkway, n in Placobdella akahkway n. sp. (Clitellata: Hirudinea: Glossiphoniidae), a new leech species from central Canada

FIGURE 5. Placobdella akahkway, n. sp. internal features showing the proboscis (pr), compact salivary glands (sg), bacteriomes (bac) and the esophagus (es). Arrowheads point to some of the diffuse salivary tissues, which are outlined in the schematic drawing and discussed further in the text.

FIGURE 3. Placobdella akahkway, n in Placobdella akahkway n. sp. (Clitellata: Hirudinea: Glossiphoniidae), a new leech species from central Canada

FIGURE 3. Placobdella akahkway, n. sp. Diagram showing segmentation and gross papillation pattern of the holotype specimen. Detailed papillation pattern from somites XVI–XVIII shown as an inset to the right.

FIGURE 4. Placobdella akahkway, n in Placobdella akahkway n. sp. (Clitellata: Hirudinea: Glossiphoniidae), a new leech species from central Canada

FIGURE 4. Placobdella akahkway, n. sp. external features. (A) eyespots (ey); (B) anus (an) and pre-anal papillae; (C) mouth pore (mp); (D) male (mg) and female (fg) gonopores.

FIGURE 2. Placobdella akahkway, n in Placobdella akahkway n. sp. (Clitellata: Hirudinea: Glossiphoniidae), a new leech species from central Canada

FIGURE 2. Placobdella akahkway, n. sp. dorsal (left) and ventral (right) views. (A) Holotype, ROMIZI11495; (B) Paratype, ROMIZI11464, with young attached to the venter; (C) Paratype, ROMIZI11494; (D) Paratype, ROMIZI11542; (E) Paratype, ROMIZI11247.

FIGURE 1 in Placobdella akahkway n. sp. (Clitellata: Hirudinea: Glossiphoniidae), a new leech species from central Canada

FIGURE 1. Map of Canada showing the collection sites for specimens of Placobdella akahkway n. sp. in solid green circles. The star indicates the type locality.

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FIGURES 1–6 in Discovery and natural history of the mussel leech Batracobdella kasmiana (Oka, 1910) (Hirudinida: Glossiphoniidae) in Russia

FIGURES 1–6. The mussel leech Batracobdella kasmiana (Oka, 1910). Fig. 1. The distribution range of B. kasmiana and sites in which leeches were searched but were not found. Red circles represent our new records from the Russian Far East: (R01) Komarovka River, Razdolnaya (Suifen) River basin, (R02) Soldatskoe Lake, Razdolnaya (Suifen) River basin, and (R03) Gladkaja River. Blue circles represent published records from Honshu, Japan: (J01) Komatsu, Biwa Lake, (J02) Hii River, (J03) Kasumigaura Lake, (J04) Owari, (J05) Bizen, and (J06) Kyoto (data sources: Oka 1917; Yamauchi et al. 2008; Itoh 2012; Shimazu 2014). Blue cross-hatching represents the approximate range in China in accordance with boundaries of province-level administrative divisions (Yang 1996). Black empty circles represent sites studied by us in which parasitic leeches were not recorded (E01–E12). Map: Mikhail Y. Gofarov; Fig. 2. Live B. kasmiana specimens (n = 55) in the internal cavity of Sinanodonta sp., Gladkaja River (specimen no. 1). Numerous small black inclusions are parasitic mites (Acari: Unionicolidae). Scale bar: 10 mm. Photo: Ilya V. Vikhrev; Figs. 3–4. Localization of live B. kasmiana specimens on foot (3) and mantle (4) of Sinanodonta sp., Gladkaja River (specimen no. 1). Numerous small black inclusions are parasitic mites (Acari: Unionicolidae). Scale bars: 5 mm. Photos: Ilya V. Vikhrev; Fig. 5. Morphological variability of adult B. kasmiana specimens from Margaritifera dahurica, Komarovka River (d—dorsal view, v—ventral view). Scale bar: 2 mm. Photos: Olga V. Aksenova; Fig. 6. Eyes of adult B. kasmiana specimens (dorsal view), Primorsky Krai, Russian Far East. Scale bar: 0.5 mm. Photos: Olga V. Aksenova.

FIGURE 13 in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 13. Large salivary cells. Translucent, preserved specimens of H. lineata showing internal morphological features of this species: A, large unicellular salivary cells (HL-11A), dorsal view; B, presumptive trematode metacercariae (HL-8B) which can be confused with salivary cells, ventral view. Abbreviations: a, atrium; cc, crop caecum; e, eye; i, intestine; pr, proboscis; me, metacercaria; pc, posterior crop caecum; r, rectum; sc, salivary cell; sb, salivary bundle; t, testis. Scale bar: 1.0 mm.

FIGURE 10 in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 10. Evidence that the neck region of H. lineata is capable of extending and swelling (see section on feeding biology). A, proboscis everted; B–D, proboscis at rest, i.e. not everted. A (HL-9D), neck region narrows (white arrow) when proboscis is extended; B (HL-15A), annulation of the neck region at rest is more subdivided than annuli in mid-body region; C (HL-13A), ventral nerve cord (blue) between ganglia VIII–IX and IX–X is characteristically sinuous (black arrow) when individual is at rest. D (HL-13A), single salivary ductules are sinuous (black arrow) within the proboscis when individual is at rest. Abbreviations: co, circumpharyngeal commissure (brain); sd, single salivary ductule (brown); sb, salivary bundle (brown). Scale bar: 10 mm.

FIGURE 5 in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 5. Dissections of individuals of H. lineata from disparate locations in the Albemarle region reveal that certain features of the digestive and reproductive systems are anatomically invariant. This relative lack of internal variability corroborates the tentative view they all represent the same biological species in this region regardless of pigmentation and papillation. A, Pasquotank County (HL-5A, 12.8 mm); B, Wayne County (HL-6A, 18.5 mm); C, Tyrrell County (HL-15A. 15.5 mm). Abbreviations: a, atrium; cc, crop caecum; es, esophagus; ic, intestinal caecum; pc, posterior crop caecum; pr, proboscis; r, rectum; sc, salivary cell; sb, salivary bundle; sv, seminal vesicle; t, testis; tp, tip of proboscis.

FIGURE 7 in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 7. Caudal end of H. lineata showing arrangement of dorsal papillae and the rectum under different conditions of lighting and preservation (5% formalin). A (HL-4), in life, under natural light. B, faded preserved specimen (HL-9B), LED light. C, faded preserved specimen (HL-9F), LED light. PAPILLAE. A, papillae are dark and conspicuous, located on a2 (middle) annulus along the medial stripe and the intermediate position. B, papillae are present but unpigmented (i.e. faded) and barely discernible. C, peculiar columnar link between external papillae and the underlying crop caeca. RECTUM. A, brown foodstuff visible in the crop caeca. Note: the rectum consists of two adjacent compartments. B, the rectum is visible through the translucent body wall as a slightly convoluted tube leading to a relatively large anus. C, the rectum has an apparent anterior third chamber located behind the intestine. Abbreviations: an, anus; pa, papilla; pc, posterior crop caecum; r, rectum. Scale bar: 1.0 mm.

FIGURE 16 in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 16. Reproductive anatomy of H. lineata. A, ventral view of adult (HL-5F) showing six pairs of testes; B, male structures showing atrium and seminal vesicles; C, female structures showing mature ovisac containing ova; D, spermatophore; E, external location of male gonopore at XIa3/XIIa1. Abbreviations: a, atrium; os, ovisac; ov, ovum; sv, seminal vesicle; t, testis. Scale bars: A, 1.0 mm; B–C, 100 μm.

FIGURE 3 in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 3. Three co-existing species of Helobdella indigenous to the Albemarle region of the Outer Banks, North Carolina. A, H. lineata, the subject of this paper, photographed in life, pigment and papillar pattern intact (HL-26); B, H. stagnalis (sensu lato) (Hst-1A); C, H. elongata (HE-2A). Scale bar: 1.0 mm.

FIGURE 2 in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 2. Two types of aquatic habitat in southern Tyrrell County contain snail leeches provisionally identified as Helobdella lineata with different respective pigment patterns, suggesting pigmentation in this species is an adaptation to surroundings rather than inherently taxonomic. A–C, Agricultural dike canal draining low farmland [35.719874–76.107744]. Virtually all leeches collected in this muddy canal have typical metameric white spots, described in the paper as pigment variant #1. D, Sandy shoreline of a swamp with typical lush vegetation and clear water [35.721466–76.10195]. All individuals at this site have longitudinal stripes (described herein as pigment variant #4), i.e., lacking metameric white spots. In spite of differences in pigmentation the two respective habitats are only 0.55 km apart. The canal (A–C) is location of the 12-year population study presented in this paper. Arrows, aluminium leech traps (roughly 14 x 23 cm).

FIGURE 6 in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 6. Variation in dorsal pigmentation and papillation in H. lineata. Two individuals collected from the same location. A, (HL-4), lightly pigmented with metameric spots. B, (HL-26), darkly pigmented with metameric spots. By far the majority of individuals throughout the Albemarle region were lightly pigmented with metameric white dots as illustrated in A. Note: pigmentation disappeared quickly after preservation making the specimen translucent. Abbreviations: b, brooding depression on ventral side; e, eye; m, medial stripe; p, posterior crop caecum; rectum; pa, papilla; ws, metameric white spot. Scale bar: 1.0 mm

FIGURE 1 in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 1. The Albemarle region of the Outer Banks in the mid-Atlantic coastal plain of North Carolina, USA. The wetlands area shaded in green is the geographic focus of this study of an indigenous snail leech provisionally identified as Helobdella lineata. Over 400 specimens of this heretofore unidentified species were collected over twelve years of this study. Scale bar: 100 km.

FIGURE 12 in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 12. Terminal structure of the proboscis of H. lineata at increasing magnifications. A (HL-9L), ventral view. Everted proboscis showing unusually large mouth; tip of proboscis with circular array of lobules; the proboscis and its lumen widens near the tip. B, C, salivary ductules and close-up of lobules at the proboscis tip; D, salivary pores with secretions at tip of proboscis. Abbreviations: lu, lumen of proboscis; lt, lobule of tip of proboscis; mo, mouth; os, oral sucker; ot, opening of tip of proboscis; sd, salivary ductule; sp, salivary pores at tip of proboscis. Scale bars: A, 1.0 mm; C, 0.1 mm.

FIGURE 9 in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 9. Taxonomic significance of eyes of Helobdella compared to Placobdella. A, B, Helobdella lineata; C, Placobdella multilineata. A, Distance between eyes is maximal when the proboscis (pr) is everted. B, When the proboscis is retained internally the eyes are closer together. C, In the turtle leech P. multilineata the eyes coalesce physically at the mid-line and are not moveable relative to each other. Furthermore, each apparent 'eye' of P. multilineata on each side is composite, being comprised of a larger anterior eye and a smaller posterior eye. Scale bars: A, B, 0.3 mm; C, 0.1 mm.

FIGURE 8 in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 8. Views of the same individual of H. lineata (HL-26) reared in isolation in UK and gave rise to viable offspring on three occasions without intervening fertilisations. Batch 1(left): egg sacs had been laid on venter on 8 Jan, and photographed five days later prior to hatching (arrow). Batch 2 (right): same individual without subsequent fertilisation deposited a second batch of eggs on 18 Feb; viable embryos (arrow) were photographed 11 days later. Note five distinct pairs of crop caeca in both images but the configurations of the caeca are markedly different. Batch 3 (not photographed) was laid on 28 March. Scale bar: 1.0 mm.

FIGURE 15 in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 15. Extra-oral feeding of H. lineata (continued) A (HL-9A, dorsal view), boluses of liquidised snail have fused into one large bolus internal to the mouth, now incapable of passing out of the mouth. B (close-up of A). Note: compare the large size of the bolus to that of the mouth, and lumen of proboscis. C, Another individual (HL-9G, ventral view) showing continuity of liquidised snail (bm) in the post-proboscis region, much foodstuff having already filled the esophagus and crop. Note: compare swelling in the neck region shown in Fig. 10. Abbreviations: bm, bolus mass; c, crop; cc, crop caecum; es, esophagus; m, mouth; pc, posterior crop caecum; tp, tip of proboscis. Scale bar (A, C): 1.0 mm.

FIGURE 14. Extra-oral feeding. Evidence that H in Biology, morphology and taxonomy of a snail-feeding leech from North Carolina, USA, provisionally identified as Helobdella lineata (Verrill, 1874) (Glossiphoniidae): First evidence for extra-oral digestion in the Hirudinea

FIGURE 14. Extra-oral feeding. Evidence that H. lineata 'swallows' a liquidised bolus of pre-digested snail. A (HL-11D, ventral view), several boluses of liquidised snail aggregated at the proximal end of the proboscis and about to pass through the mouth. B (dorsal view of specimen A), the distal end of the proboscis contains two discrete boluses (labelled, b) near the proboscis tip, en route toward the mouth. C (HL-1B, ventral view), About 10 aggregated boluses having passed through the mouth. Note the exceptional size of the mouth in B and C. Abbreviations: b, bolus; es, esophagus; m, mouth; tp, tip of proboscis. Scale bar: 0.25 mm.