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Hirudiniformes |
| 1 | External copulatory gland pores on venter, located about 10-11 annuli posterior to the male gonopore | 2* |
| - No copulatory gland pores 10-11 annuli behind male gonopore | 5* |
| 2 | Gonopores separated by 2-2.5 annuli | 3* |
| -Gonopores separated by more than four annuli, usually five | 4* |
| 3 | Dorsum with about 21 metameric red or orange dots at mid-line:
With 24 copulatory gland pores (two rows of two groups, containing six gland pores each); known only from eastern Massachusetts (Smith 1977) | Macrobdella sestertia Whitman, 1886 |
| -Dorsum lacking red or orange dots:
With eight copulatory gland pores (two rows of four); widely distributed throughout southeastern states. | Macrobdella ditetra Moore, 1953 |
| 4 | Four copulatory gland pores (two rows of two):
Widely distributed throughout northern states and Canada. | Macrobdella decora (Say, 1824) |
| - Six copulatory gland pores (three transverse rows of two each):
This form occurs at the southern periphery of the range of its very closely allied species Macrobdella decora, near the Missouri-Kansas border; Parallel modifications of the copulatory pores occur at the extreme southern Appalachian end of the range of M decora (Sawyer and Pass 1972). | Macrobdella diplotertia Meyer, 1975 |
| 5 | Glandular area around gonopores, the latter obscured by deep copulatory depressions:
Dorsum with a yelllow or brown stripe; gonopores separated by 3-4 annuli but difficult to see; widely distributed in southern states; A related form, the genotype Philobdella floridana (Verrill 1874), is inadequately distinguished from P gracilis, with which it is probably conspecific; type locality; Lake Okeechobee, Florida | Philobdella gracilis Moore, 1901 |
| - No glandular area around gonopores; gonopores distinct and separated by five or more annuli | 6* |
| 6 | Gonopores separated by five (or 5.5) annuli
Female gonopres small and not nipple-like | 7* |
| - Gonopores separated by seven (or 6.5) annuli:
Female gonopore large, conical, nipple-like in adults, flattened in juveniles | Haemopis septagon Sawyer and Shelley, 1976 |
| 7 | 27 distinct annuli from oral cavity to annulus (XIb6) bearing male gonopores, annuli VIIa3 and VIIIa1 completely subdivided ventrally | Haemopis terrestris (Forbes, 1890) |
| -25 distinct annuli from oral cavity to annulus (XIb6) bearing male gonopores, annuli VIIa3 and VIIIa1 not subdivided ventrally | 8* |
| 8 | With jaws and teeth | 9* |
| - Lacking teeth and distinct jaws | 11* |
| 9 | Dorsal surface with median dark stripe:
Body firm in life; juveniles with metameric dark transverse bands. | Haemopis kingi Mathers, 1954 |
| - No mid-dorsal dark stripe
Irregularly pigmented or unicolour; body limp in life. | 10* |
| 10 | Flexion of penis sheath at XII-XIII:
Caudal sucker large, about 0.75 width of body; recorded only from Iowa and Minnesota. | Haemopis lateromaculata Mathers, 1963 |
| -Flexion of penis sheath at XVII (XVI-XVIII):
Caudal sucker relatively small, about 0.5 width of body; very common and widely distributed throughout northern United States and Canada; two colour morphs; (i) olive green with heavy mottling dorsally and ventrally; (ii) uniform slate grey with a few irregular black blotches, resembling H grandis | Haemopis marmorata (Say, 1824) |
| 11 | Margin of oral sucker thick, rounded, aperture transverse, the lower margin anteriorly convex
Lower surface of velum closely and finely papillate; pharynx with 15 longitudinal folds; flexion of penis sheath at XVI. | Haemopis plumbea Moore, 1912 |
| -Margin of oral sucker thin, aperture elongate and arched, the lower margin sharply concave:
Lower surface of velum smooth; pharynx with 12 longitudinal folds; flexion of penish sheath at XVI | Haemopis grandis (Verrill, 1874) |
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Erpobdelliformes |
| 1 | Ejaculatory duct with a preatrial loop extending anteriorly to ganglion XI | 2* |
| -Ejaculatory duct without a preatrial loop extending to ganglion XI | 8* |
| 2 | All mid-body annuli distinctly subdivided:
Atrial horns spirally coiled like a ram's horn; distributed only in the northern tier of states and Canada. | Erpobdella obscura Verrill, 1872 |
| -All mid-body annuli not subdivided
Atrial horns simple, not spirally coiled | 3* |
| 3 | Gonopores separated by two annuli, male gonopores in furrow | 4* |
| -Gonopores separated by 2.5 or more annuli, male usually on annulus and not in furrow | 5* |
| 4 | Atrial horns anteriorly directed, with tendency toward the medial:
With two or four longitudinal rows of black pigment concentrations; male gonopore remarkably large in adults; a very common species throughout United States, Canada, and into Mexico; E montezuma from Arizona has posterior crop caeca (Davies, Singhal, and Blinn, 1985). | Erpobdella punctata (Leidy, 1870) |
| - Atrial horns directed anteriolaterally, no tendency toward the medial:
A poorly charaterized Californian species; in spite of its name some, especially young, individuals possess three pairs of eyes (Gadler and Olson, 1985); This has no bearing on the validity of its name. | Erpobdella anoculata (Moore, 1898) |
| 5 | Eyes three pairs:
Male pore very large, on annulus; a coastal plain form of E punctata punctata, from Carolinas | Erpobdella coastalis Sawyer and Shelley, 1976 |
| 6 | Eyes four pairs:
Male pore only moderately large; from northern states and Canada. | 7* |
| 7 | Dorsum with variable dark mid-dorsal stripe:
Dorsum greenish, heavily mottled; gonopores separated by 3.5-4 annuli. | Erpobdella dubia (Moore and Meyer, 1951), n comb. |
| - No mid-dorsal stripe:
Dorsum unpigmented or colour uniformly smokey grey; gonopores separated by 2.5-3.5 annuli. | Erpobdella parva (Moore, 1912), |
| 8 | Gonopores separated by two annuli | 9* |
| - Gonopores separated by three or four annuli | 10* |
| 9 | Atrial horns directed medially:
Body cream-coloured with scattered small black chromatophores irregularly dispersed, not metameric; especially pigmented around mouth; mouth large, elliptical, with thickened anterior lips; typically found in coastal plain of Carolinas | Erpobdella melanostoma Sawyer and Shelley, 1976 |
| - Atrial horns directed anteriorly or laterally, not distinctly medially:
Body uniformly smokey grey, lacking pigment entirely or sometimes with minute black pigment; occurs in northern states and Canada; A closely related form, E bucera Moore 1949, is localized in south-eastern Michigan; atrial horns projecting more laterally than E fervida | Erpobdella fervida (Verrill, 1874) |
| 10 | Gonopores separated by three annnuli: | Erpobdella microstoma (Moore, 1901) |
| - Gonopores separated by four annuli: | Erpobdella tetragon Sawyer and Shelley, 1976 |
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Glossiphoniidae |
| 1 | Eyes four pairs (Theromyzon)
Three nominal species recognized here based on number of annuli between gonopores; in need of anatomical examination of the vaginal region of the female system (see Livanow 1902). | 2* |
| - Eyes one, two or three pairs | 4* |
| 2 | Gonopores separated by two annuli: | Theromyzon bifarum |
| - Gonopores separated by three or four annuli | 3* |
| 3 | Gonopores separated by three annuli:
The common species of the western United States and Canada. | Theromyzon trizonare |
| -Gonopores separated by four annuli:
Common in Europe but its presence in North America requires re-evaluation based on criteria other than gonopores. | Theromyzon tessulatum (OF Muller, 1774) |
| 4 | Mid-body segments biannulate:
Caudal sucker large, conspicuous; known from mountains of North and South Carolina; permanent parasites on salamanders. | Placobdella biannulata (Moore, 1900) |
| - Mid-body segments tri-annulate | 5* |
| 5 | Eyes three distinct pairs
Placobdella hollensis has accessory eyes and not included here. | 6* |
| -Eyes one or two pairs | 8* |
| 6 | Gonopores share a common pore:
Three pairs of eyes, with a significantly smaller space between the eyes of the first pair than that between pairs 2 and 3; size small, less than 15 mm at rest; egg sacs attached directly to venter; colour whitish; six pairs crop caeca; A closely related form, A swampina (Bosc, 1802), lives in coastal Carolinas (Sawyer and Shelley 1976). | Alboglossiphonia heteroclita (Linnaeus, 1761) |
| - Gonopores separated by two annuli
Three pairs of eyes, all about equidistant from each other; size moderate, about 25 mm; egg sacs attached to substrate; darkly pigmented, with some dorsal papillae; seven pairs crop caeca; two conspicuous paramedian lines of brown pigmentation. | 7* |
| 7 | Dorsum with conspicuous tubercles on second (2a) and third (a3) annuli of each mid-body segment:
In North America known only from Alaska and British Columbia (Klemm 1981) |
| Glossiphonia verrucata (Fr Muller, 1844) |
| -Dorsum relatively smooth, lacking tubercles on the third (a3) annulus:
Common throughout North America except in southeastern states; this species, originally described as elegans Verrill, 1872, differs from its European namesake by having seven, instead of six, pairs of crop caeca. | Glossiphonia complanata (Linnaeus, 1758). |
| 8 | Anterior sucker distinct from body
Temporary parasites of fish (Shelley and Braswell 1981) | 9* |
| - Anterior sucker not distinct from body | 10* |
| 9 | Dorsum with three prominent ridges or keels: | Placobdella montifera Moore, 1906 |
| - Dorsum smooth, lacking ridges: | Placobdella nuchalis Sawyer and Shelley, 1976 |
| 10 | Eyes distinctly separated, usually by the diameter of one eye or more
Crop caeca six or fewer pairs; no mycetomes; predaceous on invertebrates; eyes single, not compound; gonopores separated by one or no annuli. | 11* |
| -Eyes close together or touching
Crop caeca seven pairs; one pair mycetomes; parasitic on vertebrates; eyes in some species compound; gonopores separated by two annuli. | 17* |
| 11 | Brown plaque (=scute) in mid-dorsal neck region:
Nearly indisitinguishable from the European H stagnalis | Helobdella modesta Verril |
| - No plaque in neck region | 12* |
| 12 | Gonopores share a common pore:
Dorsal and ventral surfaces heavily pigmented with uniform greyish-blue chromatophores and with thin dark paramedial lines extending to neck region; body opaque; dorsum smooth; a northern species widely distributed throughout Canada, and in Michigan | Marvinmeyeria lucida (Moore, 1954) |
| - Gonopores separated by one annulus | 13* |
| 13 | Crop caeca represented only by one short posterior pair ('post-caeca'), to XX/XXI:
Body elongated, subcylindrical; lateral margins of body almost parallel; translucent, unpigmented body; caudal sucker small, terminal. | Helobdella elongata (Castle, 1900) |
| -Crop caeca with six pairs crop caeca
Body leaf-shaped to sublanceolate; caudal sucker ventral | 14* |
| 14 | Dorsum with distinct papillae or tubercles | 15* |
| -Dorsum smooth, lacking papillae or tubercles | 16* |
| 15 | Dorsum roughly and heavily papillated:
Dorsum with many whitish, rounded papillae arranged in 5-9 longitudinal series on each neural (a2) annulus; occurs in Great Lakes region. | Helobdella papillata (Moore, 1906) |
| - Dorsum typically with three series of small black-tipped papillae:
A notoriously variable species representing a complex of forms; typically with four series of metameric white dots on neural annulus external to papillae; no mid-dorsal dots; atypical forms with longitudinal stripes of with reduced number of papillae; Often given the name H triserialis which is strictly endemic to South America. | Helobdella papillata (E Blanchard, 1849) |
| 16 | Dorsum with distinct transverse pigment patterns:
Dorsum with transverse rusty-brown bands alternating with white bands, the latter consisting of 8-10 confluent white dots on neural annulus; no longitudinal pattern; pigment fades in ethanol; known from western Great Lakes states | Helobdella transversa Sawyer, 1972 |
| - Dorsum with longitudinal pigment patterns:
Dorsum with six major longitudinal white stripes alternating with six coffee-brown stripes, including a mid-dorsal band; no transverse pattern; pigment remaining after preservation in ethanol; a variable species inadequately distinguished from Helobdella triserialis complex with which it is closely allied. | Helobdella fusca (Castle, 1900) |
| - Dorsum with longitudinal pigment patterns:
Dorsum with fine coffee-brown stripes, excluding a mid-dorsal band; no transverse pattern | Helobdella lineata (Verril) |
| 17 | Salivary glands diffuse, in anterior two-thirds of body
Caudal sucker typically modified for semipermanent attachment to host (ie large, on peduncle or with rim of specialized papillae); body typically rounded, soft-bodied, unicoloured; semipermanent parasites on fish or amphibians. | 18* |
| - Salivary glands compact, typically two pairs which insert at base of proboscis
Caudal sucker not remarkable, ventral; body typically flattened, firm-bodied and with distinct pigment patterns; temporary parasites on amphibians and reptiles. | 20* |
| 18 | Anus anteriorly displaced, at XXIII/XXIV:
Permanent parasite in gill-chamber of fish, especially the drum, Aplodinotus grunniens; caudal sucker set off from body by a narrow peduncle. | Placobdella pediculata (Hemingway, 1908), |
| - Anus in normal position on segment XXVII
Caudal sucker relatively enlarged, rim moderately thick, bulbous and set off from body by a short peduncle. | 19* |
| 19 | Rim of caudal sucker with about 30 retractile digitate processes which when everted project finger-like along inner margin of sucker cavity:
Usually found in gill-chambers of fish; mid-body segments three-annulate with tendency toward secondary subdivision; An aberrant form, A annectens Moore 1906, is characterized by about 60 digitiform processes, but has not been found since original description | Actinobdella inequiannulata Moore, 1901 |
| -Rim of caudal sucker lacking digitate processes:
Known only from Ozark hellbender Crypotbranchus in Missouri. | Actinobdella cryptobranchii (Johnson and Klemm, 1977) |
| 20 | Accessory eyes present:
Two pairs of variable concentrations of dark pigment situated two and five annuli behind the single functional pair of eyes, giving the impression of three pairs of eyes | Placobdella hollensis (Whitman, 1892) |
| - No accessory eyes | 21* |
| 21 | Dorsum with conspicuous white genital and anal patches
White ring in neck region | 22* |
| - Dorsum without white patches | 24* |
| 22 | Dorsum smooth, no papillae or tubercles
Body flattened | 23* |
| -Dorsum with three series of usually dark-tipped papillae:
Body convex, thick, opaque; a variable species, incompletely characterized; known hosts: fish (Amin 1981). | Placobdella phalera (Graf, 1899), n comb |
| 23 | Five longitudinal rows of white prominences surrounded by yellowish dots equidistant longitudinally and transversely;
No true dark-tipped papillae; body ovate-lanceolate; known from Michigan. | Placobdella michiganensis (Sawyer, 1972) |
| -Lacking the five longitudinal rows of white prominences:
No true dark-tipped papillae; body lanceolate; lightly pigmented with small brown, cutaneous chromatophores; or pigmented with a broken longitudinal median stripe and two rows of submarginal spots; widespread in south-eastern United States. | Placobdella translucens Sawyer and Shelley, 1976 |
| 24 | Dorsum smooth, with no (or only inconspicuous) papillae | 25* |
| - Dorsum conspicuously papillated, particularly evident at mid-dorsal line | 26* |
| 25 | Dorsum with a vague, dark median stripe:
With two paramedial pairs of yellowish metameric spots; venter not striped; commonly encountered on mating amphibians in early spring, in woods ponds of northern United States and Canada | Placobdella picta (Verrill, 1872), n comb |
| - Dorsum lacking dark median stripe:
Venter with 8-12 bluish-green longitudinal stripes; dorsal pigmentation variable; typically encountered on turtles but often free-living | Placobdella parasitica (Say, 1824) |
| 26 | Dorsum rough, entirely covered with numerous papillae, usually varying in size or randomly arranged:
Dorsum usually with a brown band at mid-dorsal line, interrupted 4-5 times; venter unstriped, with scattered dark chromatophores or plain; typical of the northern United States and Canada |
| Placobdella ornata (Verrill, 1872) |
| -Dorsum less rough, not entirely covered with papillae; larger papillae in 5-7 longitudinal rows | 27* |
| 27 | Venter with about eight bluish longitudinal stripes;
Body ovate-lanceolate; dorsum with a vague, median, longitudinal stripe, contained in a wider bluish stripe; the two pairs of salivary glands enter base of proboscis independently; widely distributed (Davies and Wilkialis 1980; Sawyer and Shelley 1976). | Placobdella papillifera (Verrill, 1872) |
| - Venter plain or with fine black chromatophores:
Body lanceoloate; typically with narrow dark mid-dorsal stripe which is usually continuous; the two pairs of salivary glands enter base of proboscis via a common duct; typically found in southeastern United States, where it replaces P ornata: sometimes parasitic on alligators (Forrester and Sawyer 1974) | Placobdella multilineata Moore, 1953 |