APPENDIX I-Descriptions of glossiphoniid leeches.


Alboglossiphonia heteroclita (Linnaeus, 1761)

Hirudo heteroclita Linnaeus, 1761

Hirudo hyalina O.F. Müller, 1774

Hirudo swampina Bosc, 1802

Hirudo papillosa Braun, 1805

Hirudo trioculata Carena, 1820

Hirudo arcuata Fabricius, 1826

Clepsine hyalina Moquin-Tandon, 1827

Clepsine carenae Moquin-Tandon, 1827

Glossobdella hyalina de Blainville, 1827

Glossobdella trioculata de Blainville, 1827

Glossobdella carenae de Blainville, 1828

Clepsine carenae de Filippi, 1839

Clepsina hyalina Brightwell, 1842

Glossiphonia heteroclita Moquin-Tandon, 1846

Glossopora hyalina Johnston, 1846

Glossiphonia carenae Moquin-Tandon, 1846

Glossiphonia arcuata Moquin-Tandon, 1846

Clepsine swampina Diesing, 1850

Clepsine papillosa Grube, 1851

Glossiphonia hyalina Thompson, 1856

Clepsine pallida var. a Verrill, 1874

Clepsine heteroclita Whitman, 1878

Clepsine striata Apathy, 1888

Clepsine palonica Lindenfeld and Pietrusuzynski, 1890

Glossiphonia trioculata Blanchard, 1893

Glossosiphonia heteroclita Blanchard, 1894

Glossiphonia heteroclita Castle, 1900

Glossiphonia heteroclita Harding, 1910

Glossiphonia heteroclita Ryerson, 1915

Glossiphonia papillosa Pawlowski, 1936

Glossiphonia hyalina Pawlowski, 1936

Glossiphonia striata Pawlowski, 1936

Glossiphonia swampina Sawyer, 1973

Glossiphonia (Alboglossiphonia) heteroclita Lukin, 1976

This species was originally described in Europe by Linnaeus. Alboglossiphonia heteroclita is rarely found but is distributed widely in the eastern United States and Canada where it feeds mainly on the fluid of mollusks and invertebrates (Elliott and Mann, 1979; Klemm, 1972; 1982; 1991). Alboglossiphonia heteroclita has three pairs of eyes which are grouped in a triangular pattern. The first pair of eyes are always closer together than are the posterior pairs. Gonopores are separated by two annuli (Moore, 1906; Elliott and Mann, 1979; Klemm, 1982; 1991; Sawyer, 1986). The body of this leech is dorsoventrally flattened. Alboglossiphonia heteroclita is usually translucent with little pigmentation (brownish-black chromatophores in sparse clusters). Dorsally, there is often a dark median, longitudinal stripe (sometimes interrupted but without paired stripes) or 4 to 7 mid-dorsal irregular, transverse bars composed of brownish-black pigmentation. No papillae are present (Moore, 1906; Elliott and Mann, 1979; Klemm, 1982; 1991; Sawyer, 1986).


Desmobdella paranensis Oka, 1930

Desmobdella paranensis (Oka, 1930; Oka, 1932; Autrum, 1936)

The genus Desmobdella is monotypic and it is found in South America (Autrum, 1936; Ringuelet, 1944; Sawyer, 1986). Descriptions of Desmobdella species note that one pair of eyes is situated in somite III, the male and female gonopores are separated by one annulus (male: XII a2/(a3 + a4); female: XII (a3 + a4)/a5), and there are six pairs of testes (Autrum, 1936; Ringuelet, 1944). Soós (1969), however, suggested that there are two annuli separating the gonopores. Salivary glands appear as semi-compact masses (Sawyer, 1986). Desmobdella species are six-annulate, but the annuli are poorly separated (Ringuelet, 1944; Sawyer, 1986). Externally, Desmobdella paranensis is unicolored and has no tubercles (Sawyer, 1986). The prey of this presumably non-blood feeding species are unknown (Soós, 1969).


Desserobdella phalera (Graf, 1899)

Clepsine phalera Graf, 1899

Placobdella phalera Graf, 1906

Haementeria (Placobdella) phalera Autrum, 1936

Batracobdella phalera Moore, 1959; Mann, 1961

Actinobdella phalera Sawyer, 1986

Desserobdella phalera Jones and Woo, 1990

This species is widely distributed throughout eastern North America and blood feeds mostly on fishes although it has been reported to feed on invertebrates (Klemm, 1972; 1991; Jones and Woo, 1990). Jones and Woo's (1990) redescription of the primarily quatrannulate D. phalera noted the following characteristics: one pair of large confluent eyes in III and one pair of smaller confluent eyes in II, diffuse salivary gland cells within parenchyma from VIII to XVI, male gonopore in the groove between XI and XII, female gonopore at XII a2/a3, and six pairs of testes located among crop caeca in XIII-XVII (Jones and Woo, 1900; Barta and Sawyer, 1990). Klemm (1985) described this species as having:

Dorsum with three series of usually dark-tipped papillae or white pointed papillae, sometimes with a thick bar encomposing mid dorsal series of papillae and marginal dots; with or without white patches approximately between genital and anal patches; body convex, thick, solidly or lightly pigmented, and opaque.

Jones and Woo (1990) further noted lateral, metameric pigmentation centered on the neural annulus of each segment and radiating papillae on the rim of the caudal sucker.


Desserobdella picta (Verrill, 1872)

Clepsine picta Verrill, 1872

Placobdella picta Moore, 1906; Ryerson, 1915

Glossiphonia picta Moore, 1923

Haementeria (Placobdella) picta Autrum, 1936

Batrachobdella picta Richardson, 1949; Moore, 1952; Meyer and Moore, 1954; Beck, 1954

Batracobdella picta Mann, 1961

Desserobdella picta Barta and Sawyer, 1990

Desserobdella picta is found in northern and central North America and it feeds exclusively on amphibians. (Klemm, 1972; 1991; Barta and Sawyer, 1990; Watermolen, 1996). This species has been implicated as a vector of trypanosomes and of apicomplexan blood parasites of anurans (Desser et al., 1990; Reilly and Woo, 1982). Barta and Sawyer's (1990) redescription of this species noted two pairs of eyes; the anterior pair in II and the larger posterior pair in III. Other important traits included: salivary cells distributed loosely in the parenchyma, male gonopore located in the middle of XI/XII, female gonopore at XII a2/a3, and six pairs of testes located among crop caeca in XIII-XVII (Barta and Sawyer, 1990). Externally, this leech is greenish-brown, and finely variegated with orange chromatophores. There is a mid dorsal line composed of dark chromatophores, and a pair of dorsolateral rows of white dots on a2 on each side of this line (Barta and Sawyer, 1990). A white transverse stripe or ring is located in the neck region of this leech.


Glossiphonia complanata (Linnaeus, 1758)

Hirudo sexoculata Bergmann, 1757

Hirudo complanata Linnaeus, 1758

Hirudo lateribus attenuatis Hill, 1759

Hirudo crenata Kirby, 1794

Hirudo crinata Pennant, 1816

Glossiphonia tuberculata Johnson, 1816

Glossiphonia complanata Johnson, 1816

Glossopora tuberculata Johnson, 1817

Erpobdella complanata de Blainville, 1818

Gossopora complanata Fleming, 1822

Clepsine complanata Savigny, 1822

Sanguisuga complanata Bruguiere, 1824

Glossobdella complanata de Blainville, 1828

Erpobdella crenata Templeton, 1837

Glossipora tuberculata Thompson, 1841

Glossiphonia sexoculata Moquin-Tandon, 1846

Glossiphonia cimiciformis Baird, 1869

Clepsine mollisima Grube, 1871

Clepsine pallida var.b Verrill, 1872

Clepsine elegans Verrill, 1872

Clepsine patelliformis Nicholson, 1873

Clepsine elegans Verrill, 1874

Clepsine pallida Verrill, 1875

Clepsine sex-puncto-lineata Sager, 1878

Clepsine sabariensis Orley, 1886

Clepsine concolor Apathy, 1888

Clepsine sexoculata Apathy, 1888

Glossiphonia pallida Vaillant, 1892

Glossiphonia complanata Blanchard, 1892 and 1894

Glossiphonia mollisisima Moore, 1900

Glossiphonia elegans Castle, 1900

Glossiphonia complanata Moore, 1906

Glossiphonia concolor Johannson, 1909

Glossiphonia complanata Harding, 1910

Glossosiphonia elegans Pinto, 1923

Glossosiphonia mollissina Pinto, 1923

Glossiphonia paludosa Ussing, 1929

Glossiphonia complanata Mason, et al., 1970

Glossiphoia (Glossiphonia) complanata Lukin, 1976

Glossiphonia complanata was originally described in Europe by Linnaeus. A similar or identical species was described in North America over a hundred years later by Verrill. This species is common in the northern half of the United States and Canada and it is well known in Europe (Miller, 1929; Klemm, 1991). Glossiphonia complanata is known as the "snail leech" because it principally feeds on the fluid of small snails and worms (Miller, 1929). The ground color of this leech is brownish green and the dorsal and ventral surfaces are marked by a pair of very conspicuous longitudinal lines reaching from the eyes to the posterior end. Papillae are elevated and yellowish and interrupt the paramedial lines. There is also a row of yellow spots along each margin. The male and female gonopores are separated by two annuli. Three pairs of eyes are situated close together in two rows parallel with the paramedial lines (Moore, 1906; Miller, 1929; Elliott and Mann, 1979; Klemm, 1982; 1991; Sawyer, 1986).


Haementeria ghilianii (de Filippi, 1849)

Haementeria ghilianii de Filippi, 1849

Liostomum ghilianii Blanchard, 1899; Weber, 1915

Haementeria (Haementeria) ghilianii Autrum, 1936

Found in South America (as are most Haementeria species), this large species uses its proboscis to blood-feed from mammals and reptiles (Sawyer, et al., 1981; 1982; 1991). Salivary cells are compact and are known to secrete hemetin (Sawyer, et al., 1982). One pair of eyes are in somite III, two annuli separate the gonopores (male: XI/XII; female: XII a2/(b5+b6)), and there are five pairs of testes (Ringuelet, 1944; 1980). Externally, H. ghilianii is a uniform olive-green color with no longitudinal pigment pattern and with numerous papillae on each annulus (Ringuelet, 1944; 1980; Sawyer, 1986).


Haementeria gracilis (Weyenbergh, 1883)

Hybobdella gracilis Weyenbergh, 1883

Blennobdella depressa E.M. Blanchard, 1849

Nephelis santiaguenis Weyenbergh, 1883

Liostomum brasiliense Weber, 1915

Liostomum gracilis Weber, 1915

Haementeria brasiliensis Pinto, 1923

Liostomum gracilis Pinto, 1923

Haementeria bonaerensis Mac Donagh, 1928

Haementeria (Placobdella) brasiliensis Autrum, 1936

Haementeria (Placobdella?) cordovensis Autrum, 1936

Haementeria (Placobdella?) bonaerensis Autrum, 1936

Haementeria bonaerensis Cordero, 1937

Haementeria cfr. brasiliensis Cordero, 1937

Haementeria gracilis Cordero, 1941; Ringuelet, 1945; 1948; 1968

This South American species with compact salivary cells blood-feeds from amphibians and birds (Sawyer, 1986). Externally, Haementeria gracilis has a dark, mid-dorsal, uninterrupted stripe. Three para-medial dark, irregular pigmented stripes are located on each side of the middle stripe. The venter also has dark paramedian stripes (Sawyer, 1986). Distinct dorsal tubercles are present paramedially on annulus a2 and laterally on annulus a3 (Ringuelet, 1944; Sawyer, 1986). As in H. ghilianii, one pair of eyes are in somite III, two annuli separate the gonopores (male: XI/XII; female: XII a2/(b5+b6)), and there are five pairs of testes (Ringuelet, 1944; 1980).


Haementeria molesta (Cordero, 1934)

Placobdella molesta Cordero, 1934

Haementeria molesta Ringuelet, 1980

Ringuelet (1980) redescribed this South American species in the genus Haementeria, removing it from genus Placobdella. One pair of eyes is in III, gonopores are separated by two annuli (male: XI/XII; female: XII (b1+b2)/a2), and salivary cells are compact. Haementeria molesta blood-feeds from amphibian hosts (Ringuelet, 1980; Sawyer, 1986). This species is uniformly green in color. Faint, longitudinal lines are present on the venter and a thin medial and three paired equidistant longitudinal lines are on the dorsal surface (Ringuelet, 1980; Sawyer, 1986). Papillae also are present dorsally (Ringuelet, 1980).


Haementeria tuberculifera (Grube, 1871)

Clepsine tuberculifera Grube, 1871

Hybobdella doringii Weyenbergh, 1879

Hylobdella doringi Blanchard, 1888

Glossiphonia tuberculifera Vaillant, 1890

Haementeria officinalis Blanchard, 1896

Liostomum coccineum Weber, 1915

Liostomum helleri Weber, 1915

Haementeria officinalis Pinto, 1923

Haementeria helleri Pinto, 1923; Augener, 1930

Haementeria (Placobdella) tuberculifera 1936

Haementeria cfr. helleri Cordero, 1937

Haementeria tuberculifera Cordero, 1941; Ringuelet 1944; 1945; 1948; 1968

As for the other Haementeria species in this study, H. tuberculifera is South American with compact salivary glands (Ringuelet, 1944; Sawyer, 1986). Reptiles, especially turtles, serve as hosts. There is one pair of eyes, and two annuli separate the male and female gonopores (Ringuelet, 1944; Sawyer, 1986). External morphology of this species consists of five longitudinal series of tubercules on the dorsum (Ringuelet, 1944). The supramarginal group on a1, large paramedials on a2, and the medial tubercle on a3 form a "V" shape on the dorsum (Sawyer, 1986).


Helobdella elongata (Castle, 1900)

Clepsine nepheloidea Graf, 1899

Glossiphonia elongata Castle, 1900

Glossiphonia nepheloidea Moore, 1906

Glossosiphonia elongata Pinto, 1923

Helobdella nepheloidea Moore, 1924

Helobdella elongata Autrum, 1936

Glossiphonia nepheloidae Miller, 1937

Helobdella elongata is an unpigmented, apapillate, pale-yellow or translucent species found in Canada and the United States (Castle, 1900; Miller, 1929; Sawyer and Shelley, 1976; Klemm, 1982). This species is slender and elongate and is earthworm-like in its behavior. It writhes and twists when disturbed as opposed to curling up into a ball like most glossiphoniids (Castle, 1900; Miller, 1929). Helobdella elongata uses its proboscis to fluid feed from aquatic annelids and insects, worms, and mollusks (Miller, 1929; Klemm, 1972). One pair of eyes is located between III and IV; one annulus separates the male gonopore (a1/a2 of XII) from the female gonopore (a2/a3 of XII); six pairs of testes are arranged intersegmentally in somites (XIII/XIV) - (XVIII/XIX); and the salivary glands are diffuse (Castle, 1900; Ringuelet, 1944; Klemm, 1982; Sawyer, 1986).


Helobdella fusca (Castle, 1900)

Clepsine papillifera Verrill, 1874

Glossiphonia lineata Moore, 1898

Glossiphonia fusca Castle, 1900

Glossiphonia fusca fusca Moore, 1906

Glossiphonia (Helobdella) fusca Moore, 1922; in part

Helobdella fusca Moore, 1959

Helobdella triserialis Soós, 1969; Davies, 1971 in part

This species is widely distributed in North, Central, and South America, and is common in the Great Lakes region (Klemm, 1991). Helobdella fusca is a non-blood feeder and it is known to feed on aquatic annelids, insects, and molluscs (Klemm, 1972). Like all Helobdella species, H. fusca has one pair of eyes, one annulus between the male and female gonopores, and diffuse salivary glands (Ringuelet, 1944; Sawyer, 1986). Helobdella fusca is a variable species with respect to its external morphology (Miller, 1929; Klemm, 1972; 1982). Moore (1906) noted extreme discrepancies in color pattern and papillation. This species is described as having either a uniform coffee-brown color or dorsal longitudinal whitish stripes alternating with coffee-brown strips and/or lines. Transverse pigmentation on the dorsum is lacking and small whitish or pale spots may or may not be present in the anal region (Klemm, 1986; 1991). Papillae are in series of one to five and they may be small, large, or doubled. Papillae are usually observed in longitudinal lines with a double or fused median series (Miller, 1929). High variability in this species has resulted with indistinguishable specimens being described as Helobdella lineata as well as H. fusca (Soós, 1969; Klemm, 1972).


Helobdella lineata (Verrill, 1874)

Clepsine papillifera var. lineata Verrill, 1874

Glossiphonia lineata Moore, 1898; Moore, 1901;1936; 1959; Cordero, 1937; Ringuelet, 1943, 1944, 1968; Mann, 1961

As with Helobdella fusca, there have been classification problems with Helobdella lineata. Specimens matching the description of H. lineata have been recognized as Helobdella fusca and as Helobdella triserialis (Soós, 1969; Sawyer, 1986). Sawyer (1986) recognizes both of these species as Helobdella triserialis and he describes it as "a notoriously variable species representing a complex of forms." Typically H. triserialis/H. lineata has three series of black-tipped papillae on the dorsum, fours series of metameric white dots on the neural annulus external to the papillae, and no mid-dorsal dots. Atypical forms have been described as having longitudinal stripes or reduced number of papillae (Sawyer, 1986). Ringuelet (1944) recognizes H. lineata as a variant of H. triserialis. Sawyer and Shelley (1972), however, recognize H. lineata as an unique species but they still note that it is highly variable in both papillation and pigmentation (ranging from dark stripes with heavy papillation to translucent-white with light papillation). This species is distributed in North, Central, and South America (as is H. triserialis) (Sawyer, 1986; Klemm, 1991). Common food includes the fluid of aquatic insects and annelids, and molluscs (Klemm, 1972). As with other Helobdella species, eyes are one pair (usually in IV), one annulus separates male and female gonopores, and salivary glands are diffuse (Ringuelet, 1944; Sawyer, 1986).


Helobdella papillata (Castle, 1900)

Clepsine papillifera var. b Verrill, 1872

Helobdella fusca var. papillata Moore, 1906

Helobdella fusca Moore, 1918; in part

Helobdella papillata Moore, 1952

This uncommon species has been collected in northern North America, especially in the Great Lakes region, and it is known to fluid feed mostly from snails (Klemm, 1991). Internal morphology is the same as other Helobdella species: one pair of eyes in IV, one annulus separating the gonopores, and diffuse salivary glands (Ringuelet, 1944; Sawyer, 1986). Classification via morphological characteristics has been difficult. It has been noted that H. lineata and H. papillata resemble H. fusca, but they have all been placed into separate species based on slight morphological character differences (Klemm, 1972). Presently, H. papillata is noted as having a dorsum with 5-9 longitudinal rows of large, whitish, and rounded papillae on each neural annulus. Coloration is yellowish-brown or unpigmented (Klemm, 1972; 1991; Sawyer, 1986).


Helobdella stagnalis (Linnaeus, 1758)

Hirudo bioculata Bergmann, 1757

Hirudo stagnalis Linnaeus, 1758

Hirudo bioculata Müller, 1774

Hirudo pulligera Daudin, 1800

Hirudo circulans Sowerby, 1806

Helluo (Hirudo) bioculatus Oken, 1815

Glossiphonia perata Johnson, 1816

Glossopora punctata Johnson, 1817

Erpobdella bioculata de Blainville, 1818

Clepsine bioculata Savigny, 1822

Glossopora bioculata Fleming, 1822

Clepsine sowerbyi Moquin-Tandon, 1827

Hirudo (Glossobdella) pulligera de Blainville, 1827

Glossobdella bioculata de Blainville, 1828

Erpobdella stagnalis Templeton, 1836

Clepsina stagnalis de Filippi, 1837

Glossiphonia circulans Moquin-Tandon, 1846

Glosopora circularis Johnston, 1846

Glossiphonia bioculata Moquin-Tandon, 1846

Glossipora bioculata Thompson, 1856

Clepsine filippi Polonio, 1863

Clepsine modesta Verrill, 1872

Clepsine submodesta Nicholson, 1873

Clepsine viridissima Picaglia, 1877

Clepsine bioculata Levinsen, 1883

Glossiphonia modesta Vaillant, 1890

Glossosiphonia stagnalis Blanchard, 1894

Glossiphonia scutifera Young, 1894

Helobdella stagnalis Blanchard, 1896

Helobdella bioculata Bayer, 1898

Glossiphonia stagnalis Moore, 1898

Glossiphonia stagnalis Castle, 1900

Glossiphonia (Helobdella) stagnalis Moore, 1922

Glossopora punctata Johnson, 1925

Helobdella stagnalis Andre, 1930

Bakedebdella gibbosa Sciacchitana, 1939

Erpobdella stagnalis Oliver, 1958

Helobdella stagnalis Moore, 1952

Helobdella stagnalis is reported from every continent except Australia and it was originally described in Europe by Linnaeus (Elliottt and Mann, 1979; Klemm, 1991). The defining characteristic of this leech is a chitinoid plate and underlying gland on the dorsum of somite VIII (Moore, 1906). The function of this plate is unknown. Eyes are simple and in one pair and widely separated on IV. Gonopores are separated by a single annulus. The skin surface is smooth and without papillae. Coloration can be opaque, pink, gray, brown, or green. This variability depends on the age of the leech and contents of alimentary tract (Moore, 1906; Miller, 1929; Elliottt and Mann, 1979; Klemm, 1982). Helobdella stagnalis feeds from the fluid of invertebrates, aquatic insects, and molluscs (Miller, 1929; Klemm, 1991).


Helobdella transversa Sawyer, 1972

This Helobdella species has one pair of eyes in IV, one annulus separating the male and female gonopores, and diffuse salivary cells (Ringuelet, 1944; Sawyer, 1986). Reported only from southern Michigan, the diet of Helobdella transversa is unknown, but it is thought to fluid feed on aquatic snails and other invertebrates (Sawyer, 1972; Klemm, 1991). This species is noted as having a dorsum with interrupted transverse rusty-brown bands alternating with irregular whitish bands. The whitish bands consist of 8-10 confluent white metameric spots on each neural annulus. No longitudinal pattern is visible (Klemm, 1982; 1991; Sawyer, 1986)


Helobdella triserialis (Blanchard, 1849)

Glossiphonia triserialis Blanchard, 1849

Clepsine triserialis Grube, 1859

Clepsine lineolata Grube, 1871

Clepsine papillifera var. b Verrill, 1872

Clepsine papillifera var. lineata Verrill, 1874

Glossiphonia lineolata Vaillant, 1890

Helobdella triserialis Blanchard, 1896

Glossiphora lineata Moore, 1898

Glossiphonia fusca Castle, 1900; Moore, 1906; Ryerson, 1915

Anoculobdella trituberculata Weber, 1915

Placobdella triserialis Apathy, 1917

Glossiphonia fusca Moore, 1918; in part

Glossiphonia (Helobdella) fusca Moore, 1922; in part

Glossosiphona fusca Pinto, 1923

Helobdella fusca Moore, 1924

Helobdella punctata-lineata Moore, 1939

Helobdella nigricans Ringuelet, 1943

Helobdella striata Ringuelet, 1943

Helobdella triserialis lineata Ringuelet, 1943; 1980

Helobdella unilineata Ringuelet, 1943

Helobdella lineata Soós, 1969; Davies, 1971- in part

Helobdella fusca Soós, 1969- in part

Helobdella trialbolineata Klemm, 1974

Helobdella lineata Klemm, 1977

Helobdella punctata-lineata Moore, 1939; 1959

Helobdella punctatalineata Klemm, 1972b, 1976; 1977

Helobdella punctatolineata Sawyer and Kinard, 1980

This species is variable and polymorphic and it is represented by several complex forms exhibiting different morphological features (Klemm, 1991). Both Helobdella lineata and H. fusca have been considered different forms of this species (Klemm, 1972). Klemm's 1982 description of Helobdella triserialis is characteristic of the variability of this species:

Dorsum with three rows (typically) or fewer of small, black-tipped or uniformly pale white papillae; pigmentation uniform, or arranged in numerous longitudinal light and dark brown, gray, or black lines and/or stripes, or dorsum with three broad, longitudinal white stripes, one median row and one on each side sub marginal, also at posterior end two short white stripes, few or many whitish spots on neural annuli, unaligned or if aligned, then confined to areas lateral to papillae, or color uniform with papillae few, scattered, or in three rows; whitish spots on neural annuli unaligned or aligned.

Internally, this species possesses the same characteristics as other Helobdella species: one pair of eyes in IV, one annulus separating gonopores, and diffuse salivary glands (Ringuelet, 1944; Sawyer, 1986). Helobdella triserialis is widely distributed in North, Central, and South America and is known to feed on the fluid of aquatic snails and other invertebrates (Klemm, 1991).


Hemiclepsis marginata (Müller, 1774)

Hirudo marginata Müller, 1774

Hirudo variegata Braun, 1805

Hirudo cephalota Carena, 1821

Haemocharis (?) cephalota Carena, 1821

Hirudo oscillatoria Saint-Amans, 1825

Piscicola marginata Moquin-Tandon, 1827

Piscicola tessellata Moquin-Tandon, 1827

Glossobdella cephalota de Blainville, 1827

Ichthyobdella marginata de Blainville, 1828

Haemocharis marginata de Filippi, 1837

Clepsine marginata Müller, 1844

Glossiphonia marginata Moquin-Tandon, 1846

Clepsine marginata Diesing, 1850

Hirudo flava Dalyell, 1853

Glossiphonia flava Johnston, 1865

Clepsine viridissima Picaglia, 1877

Clepsine marginata Apáthy, 1888

Glossiphonia marginata Blanchard, 1892

Hemiclepis marginiata Blanchard, 1894

This primarily European and Asian species blood feeds from fish or amphibian larvae (Soós, 1969; Elliottt and Mann, 1979; Sawyer, 1986). Eyes are typically in two pairs on III and IV (Elliottt and Mann, 1979; Sawyer, 1986). Two annuli separate the gonopores. The male gonopore is located at X/XI and the female gonopore is located at XI2/XI3 (Autrum, 1936). There are 10 pairs of testes (Soós, 1969). The head of Hemiclepsis marginata is characteristically broadened (Elliottt and Mann, 1979; Sawyer, 1986). Externally, this species is green or pale yellow-brown, usually with seven longitudinal rows of bright yellow spots (Elliott and Mann, 1979).


Marsupiobdella africana (Goddard and Malan, 1912)

Marsupiobdella africana is the only species in this monotypic African genus and is unique in having developed a permanent brood pouch on its ventral surface in which eggs and young develop. This pouch is an open groove involving seven to eight annuli from somites XVII to XVIII (van der Lande and Tinsley, 1976). Eyes are one pair in an unknown somite (Soós, 1969). The male gonopore is situated in the groove between XIa3 and XIIa1 and the female gonopore lies between a2 and a3 of somite XII. Six pairs of testes are arranged in somites XIII through XIX. The proboscis opening lies in the center of the velum situated in the ventral side of the oral sucker, and when the proboscis is retracted it extends from somites VI to XI (van der Lande and Tinsley, 1976). Salivary glands are very diffuse (Soós, 1969). Externally, Marsupiobdella africana is transparent with green and brown pigment-containing cells scattered throughout the tissues (van der Lande and Tinsley, 1976). This monotypic species feeds on blood from various hosts such as frogs and toads (van der Lande and Tinsley, 1976).


Oligobdella biannulata (Moore, 1900)

Microbdella biannulata Moore, 1900

Oligobdella biannulata Moore, 1920

Oligobdella species are the only bi-annulate glossiphoniid leeches. All other glossiphoniids are triannulate. Oligobdella biannulata is the type species of this genus and has one pair of eyes in the posterior half of somite III, one annulus between the male (between XI and XII) and the female (between a1 and a2 on XII) gonopores, and five pairs of testes (XV-XX) (Moore, 1906). The skin of this leech is smooth and lacks papillae. No external pigment patterns are present (Soós, 1959; Sawyer and Shelley, 1976). The color of the dorsum is pale olive-green, brown, or gray and the caudal sucker is large and conspicuous (Moore, 1906; Klemm, 1982; Sawyer, 1986). This species presently is found only in North and South Carolina where it blood feeds on desmognathine salamanders (Sawyer and Shelley, 1976; Klemm, 1982; Sawyer, 1986). Oligobdella biannulata is considered to be a species of some evolutionary importance due to its bi-annulate state (possibly evidence of an intermediate genus between the uni-annulate Oligochaeta and the multi-annulate Hirudinea) (Moore, 1906; Sawyer, 1971; Sawyer and Shelley, 1976).


Placobdella montifera (Moore, 1906)

Glossiphonia trisculcata Baird, 1869

Clepsine papillifera var. carinata Verrill, 1874

Hemiclepsis carinata Moore, 1901

Placobdella montifera Moore, 1906

Haementeria (Placobdella) montifera Autrum, 1936

Placobdella montifera Moore, 1952

Placobdella parasitica Amin, 1977

This species is widely distributed in North America but it is not common (Klemm, 1991). Placobdella montifera blood feeds and has been reported to prey upon turtles, fish, frogs, and toads (Klemm, 1972; 1991). This placobdellid species is characterized as having compound eyes (situated on anterior annulus of III where the anterior pair is smaller and either in the hollow of or slightly anterior to the larger pair; if the anterior pair is not visible, eyes resemble Haementeria species), two pairs of diffuse salivary glands, gonopores separated by two annuli (male gonopore: XI/XII; female gonopore: a2/a3 on XII), and six pairs of testisacs situated intersegmentally among the crop caeca from XIII-XIV to XVIII-XIX (Moore, 1906; Miller, 1929; Davies and Wilkialis, 1982; Yang and Davies, 1985; Sawyer and Shelley, 1976; Sawyer, 1986). Placobdella montifera is easily recognized by its widely expanded discoid head and dorsum of three prominent and nearly continuous ridges or keels which may not be discernible in live specimens (Moore, 1906; Miller, 1929; Klemm, 1982; Sawyer, 1986). Coloration is greenish-brown or pale olive-brown with light yellow and green marginal spots (Miller, 1929; Klemm, 1982). A dark brown, interrupted, median dorsal line is usually present and the ventral surface is plain brown (Miller, 1929).


Placobdella ornata (Verrill, 1872)

Clepsine ornata Verrill, 1872

Clepsine ornata var. rugosa Verrill, 1874

Clepsine ornata var. stellata Verrill, 1874

Glossiphonia parasitica var. rugosa Castle, 1900

Placobdella rugosa Moore, 1901

Haementeria (Placobdella) rugosa Autrum, 1936

Placobdella ornata Moore, 1952- Northern variety

Placobdella ornata Moore, 1959

Placobdella multilineata Beck, 1954

Placobdella multilineata Klemm, 1972a

Externally, this species has a dorsal surface that is completely covered with numerous papillae which bear clusters of secondary, minute papillae at the apex. Smaller papillae are randomly arranged and the larger papillae are in five longitudinal metameric rows. A brown, interrupted, mid dorsal band is usually present. Dorsal coloration is a mixture of brown, green, and yellow and the ventral surface is either unstriped with scattered dark chromatophores or is plain (Klemm, 1982; 1991; Sawyer, 1986). This placobdellid species is characterized as having gonopores separated by two annuli (male gonopore: XI/XII; female gonopore is a2/a3 on XII), six pairs of testisacs situated intersegmentally from XIII-XIV to XVIII-XIX among the crop caeca, compound eyes in III, and two pairs of compact salivary glands (Moore, 1906; Miller, 1929; Davies and Wilkialis, 1982; Yang and Davies, 1985; Sawyer and Shelley, 1976; Sawyer, 1986). Placobdella ornata is common and widespread in the northern half of the United States and Canada and there have been reports of it in southern localities (Sawyer, 1986; Klemm, 1991). Turtles are most commonly fed upon, but there have been some reports of blood feeding from amphibians and molluscs (Klemm, 1972; 1991).


Placobdella papillifera (Verrill, 1872)

Clepsine papillifera Verrill, 1872 in part

Placobdella papillifera Moore, 1952; Meyer and Moore, 1954; Sawyer and Shelley, 1976

?Haementeria officinalis Ringuelet, 1976

This species has been reported sporadically from the United States and Canada (Sawyer and Shelley, 1976; Davies and Wilkialis, 1982; Klemm, 1991). Placobdella papillifera blood feeds from turtles and it has been seen feeding from the American alligator (Smith, 1976). Davies and Wilkialis (1982) showed that live ducks, frogs, and fish are not fed on in the laboratory but that fresh heparinized blood from a wide range of vertebrates will be consumed. Placobdella papillifera is characterized as having compound eyes (situated on anterior annulus of III where the anterior pair is smaller and either in the hollow of or slightly anterior to the larger pair), two pairs of compact salivary glands, gonopores separated by two annuli (male gonopore: XI/XII; female gonopore: a2/a3 on XII), and six pairs of testisacs (from XIII-XIV to XVIII-XIX) (Moore, 1906; Miller, 1929; Davies and Wilkialis, 1982; Yang and Davies, 1985; Sawyer and Shelley, 1976; Sawyer, 1986). Base coloration of this species can be greenish-blue with longitudinal striping or obscure yellowish brown produced by alternating flesh-color and olive-green narrow lines. Dorsum is seen with bluish para-medial stripes containing a dark interrupted medial stripe. The dorsum also is darkly pigmented with dark chromatophores and metameric unpigmented marginal patches. The sensory (a2) annulus is the most densely colored (in comparison to a1 and a3). Transverse lines of chromatophores can be seen in each annulus, and additional chromatophore stripes run paramedially down each lateral margin. A series of brown chromatophores run down the median dorsal line and appear continuous in non-elongated individuals. Medial white areas are present in the head and neck region and the caudal sucker has unpigmented areas. Two to eight bluish longitudinal stripes line the venter. Five to seven longitudinal rows of large white-tipped papillae are present dorsally. Papillae situated on a sensory annulus are located on a field of chromatophores and they are sometimes doubled (Sawyer and Shelley, 1976; Davies and Wilkialis, 1982; Klemm, 1982).


Placobdella parasitica (Say, 1824)

Hirudo parasitica Say, 1824

Clepsine parasitica Diesing, 1850

Clepsine marmarata Sager, 1878

Clepsine chelydrae Whitman, 1889

Clepsine plana Whitman, 1891

Glossiphonia parasitica Moore, 1898

Glossiphonia parasitica var. plana Castle, 1900

Placobdella parasitica Moore, 1901

Glossosiphonia parasitica Pinto, 1923

Haementeria (Placobdella) parasitica Autrum, 1936

Placobdella parasitica Moore, 1952

Placobdella parasitica blood feeds mainly from turtles although it is also reported to feed on aquatic annelids and insects, amphibians, and molluscs (Klemm, 1972; 1991). Of the North American blood sucking leeches, P. parasitica is one of the most common and wide-spread (Miller, 1929; Klemm, 1991). Internal morphology consists of compound eyes situated on the anterior annulus of III (the anterior pair is smaller and either in the hollow of or slightly anterior to the larger pair), salivary glands are compact and in two pairs, gonopores are separated by two annuli (male gonopore: XI/XII; female gonopore: a2/a3 on XII), and six pairs of testisacs are situated intersegmentally from XIII-XIV to XVIII-XIX (Moore, 1906; Miller, 1929; Davies and Wilkialis, 1982; Yang and Davies, 1985; Sawyer and Shelley, 1976; Sawyer, 1986). Externally, P. parasitica is variable and its coloration is rich and striking (Miller, 1929). Dorsally, the underlying color is greenish brown, variously spotted, blotched, and striped bright yellow and orange (middorsally) (Miller, 1929; Klemm, 1982; 1991). There is no dark median stripe (Sawyer, 1986). Marginal spots are regular, of somewhat triangular shape, and extend over two annuli (Miller, 1929). Ventrally there are 8-12 bluish, greenish, or brownish longitudinal stripes or lines. Light yellow areas are confined mostly to a median longitudinal band which may be continuous or interrupted (Miller, 1929; Klemm, 1982).


Placobdella pediculata (Hemingway, 1908)

Haementeria (Placobdella) pediculata Autrum, 1936

Placobdella pediculata Moore, 1952

This species blood feeds only from one fish (Aplodinotus grunniens) (Hemingway, 1908; Klemm, 1982). Placobdella pediculata is uncommon and reported mainly from the Midwest region of the United States (Klemm, 1982). Body color of this leech is brownish or grayish and opaque. The skin is perfectly smooth containing no cutaneous papillae (Hemingway, 1908; Klemm, 1982). The posterior somites attenuate to form a narrow pedicle just in front of the caudal sucker. The stands out freely exposed behind the dorsally high and contractile body (Hemingway, 1908; Klemm, 1982). Placobdella pediculata has diffuse salivary cells, and compound eyes (situated on anterior annulus of III where the anterior pair is smaller) which resemble Haementeria species eyes. Male (between XI and XII) and female (a2/a3 on XII) gonopores are separated by two annuli, and six pairs of testisacs are situated intersegmentally from XIII-XIV to XVIII-XIX (Moore, 1906; Miller, 1929; Davies and Wilkialis, 1982; Yang and Davies, 1985; Sawyer and Shelley, 1976; Sawyer, 1986).


Placobdella transluscens Sawyer and Shelley, 1976

Placobdella transluscens is either lightly pigmented brown with small cutaneous chromatophores uniformly distributed on the dorsum or is pigmented with a broken longitudinal median stripe and two rows of sub marginal spots (Sawyer and Shelley, 1976; Klemm, 1982; Sawyer, 1986). The venter is unpigmented. Other unpigmented regions include a ring around the neck, metameric patches along margins, the anal patch, and three medial patches between the nuchal ring and the anal patch. Dark green chromatophores surround the genital area and outline the gut (Sawyer and Shelly, 1976). The dorsum and the venter are smooth without conspicuous papillae (Sawyer and Shelley, 1976). Gonopores are separated by two annuli (male gonopore is between XI and XII and the female gonopore is between annuli a2 and a3 on XII), eyes are compound (situated on anterior annulus of III where the anterior pair is smaller and either in the hollow of or slightly anterior to the larger pair), salivary glands are diffuse, and six pairs of testisacs are situated intersegmentally from XIII-XIV to XVIII-XIX (Moore, 1906; Miller, 1929; Davies and Wilkialis, 1982; Yang and Davies, 1985; Sawyer and Shelley, 1976; Sawyer, 1986). This species is recognized as being excessively flattened and thin and it is widespread in the south-eastern United States (Klemm, 1982; Sawyer, 1986). Prey are unknown (Sawyer and Shelley, 1976).


Theromyzon biannulatum (Klemm, 1977)

Hemiclepsis occidentalis Moore, 1912

Protoclepsis occidentalis Moore, 1922

Theromyzon occidentalis Autrum, 1936

Theromyzon meyeri Moore, 1959; Sawyer, 1972

Theromyzon maculosum Klemm, 1972; Davies, 1971, 1973

Theromyzon biannulatum Klemm, 1977

This species blood feeds from the nasal passages and conjunctiva of the eyes of waterfowl and is known only from the central and eastern United States (Klemm, 1982; 1991). Gonopores are separated by two annuli and eyes are in four pairs on II, III, IV, V (Klemm, 1982, Sawyer, 1986). Externally, T. biannulatum is variable. The body is nearly translucent, color is olive-green and flecked with black chromatophores or with longitudinal rows of cream yellow spots on the dorsum (Klemm, 1982).


Theromyzon tessulatum(Müller, 1774)

Hirudo tessulata Müller, 1774

Hirudo tesselata Bosc, 1802

Hirudo tessulatum Braun, 1805

Erpobdella tessulata Fleming, 1822

Nephelis tessellata Savigny, 1822

Ichthyobdella tesselata de Blainville, 1828

Erpobdella vulgaris var. tesselata de Blainville, 1828

Clepsine sanguinea de Filippi, 1837

Clepsine tessulata Müller, 1844

Glossiphonia tesselata Thompson, 1846

Glossiphonia eacheana Thompson, 1846

Glossiphonia tessellata Moquin-Tandon, 1846

Glossiphonia sanguinea Moquin-Tandon, 1846

Clepsine tessulatum Diesing, 1850

Hirudo vitrina Dalyell, 1853

Haemocharis eacheana Thompson, 1856

Glossiphonia vitrina Johnston, 1865

Theromyzon pallens de Filippi, 1867

Hemiclepsis tessellata Vejdovsky, 1884

Clepsine tesselata Weltner, 1887

Glossiphonia tessulatum Blanchard, 1892

Hemiclepsis tesselata Scharff, 1898

Protoclepsis tesselata Livanow, 1902

Protoclepsis tessellata Harding, 1910

Theromyzon tessulatum Pawlowski, 1936

Theromyzon tessulata Autrum, 1936

Protoclepsis granata Endrigkeit, 1940

Theromyzon tessulatum is widely distributed in Eurasia with a center of distribution in mid-Europe. It has also been recorded in North America, South America, and Africa (Wilkialis and Davies, 1980a; 1980b; van der Lande, 1983). Like other Theromyzon species, T. tessulatum blood feeds on waterfowl, feeding within mucosa of the nasal chamber and conjunctiva of the eyes (Wilkialis and Davies, 1980; Klemm, 1982; van der Lande, 1983). Eyes are in four pair: in II, III, IV, V (Sawyer, 1986). Four annuli separate gonopores. The male gonopore is at X a2/a3 and the female gonopore is at XI/XII (Autrum, 1936). This species is nearly translucent with an overall amber or greenish coloration (Klemm, 1982). Six longitudinal rows of regularly positioned yellow spots or two thin paramedial black lines are present on the dorsal surface (Klemm, 1982; Sawyer, 1986).


Theromyzon rude (Baird, 1869)

Glossiphonia rudis Baird, 1869

Clepsine occidentalis Verrill, 1874

Glossiphonia occidentalis Vaillant, 1890

Hemiclepsis occidentalis Moore, 1912

Protoclepsis occidentalis Moore, 1922

Theromyzon occidentalis Bere, 1929

Theromyzon rude Moore and Meyer, 1951

Theromyzon occidentale Meyer and Moore, 1954

Theromyzon rude is a blood feeding leech infesting the nares of North American waterfowl (Davies, 1984). This species is recorded only from the nearctic and is characteristic of the Rocky Mountain duck migration routes of the Canadian Provinces and Territories, and American States west of the Great Lakes and the American Midwest (Davies and Wilkialis, 1980). Like other Theromyzon species, there are four pairs of eyes in two sub-parallel rows in inner-paramedian positions in II, III, IV, V (Oosthuizen and Davies, 1992). Gonopores are separated by two annuli (male: XI/XII, and female XII a2/a3), salivary glands are diffuse, and six pairs of testisacs are intersegmentally arranged from XII/XIV to XVIII/XIX (Oosthuizen and Davies, 1992). Theromyzon rude is olive green in color and may have two paramedial pairs and marginal pairs of yellow, orange, or brown spots on the dorsum (Klemm, 1982).


Torix baicalensis (Stschegolew, 1922)

This species is endemic to Lake Baikal. Two pairs of eyes are currently recognized and the anterior pair is larger than the posterior pair (Autrum, 1936; Sawyer, 1986). This is a non-blood-feeding leech, but hosts are unknown. Autrum (1936) recognized this species as "species inquirenda" because it was not possible to determine the appropriate genus from the original description of internal anatomy. The location of somites also could not be determined from the original description because Torix baicalensis has only one or two annuli per somite rather than the expected three annuli.