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Smithsonian Institution

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 476

Revised List of the North American Freshwater Planarians (Platyhelminthes: Tricladida: Paludicola)

Roman Kenk

SMITHSONIAN INSTITUTION PRESS

City of Washington

1989

ABSTRACT

Kenk, Roman. Revised List of the North American Freshwater Planarians (Platyhelminthes: Tricladida: Paludicola). Smithsonian Contributions to Zoology, number 476, 10 pages, 1989.—A list of the species and subspecies of freshwater triclads of North America, including the West Indies, is presented, arranged in taxonomical order. For each taxon, the valid name, existing synonyms, known distribution, and pertinent morphological literature are given. An appendix lists the specific and subspecific names in alphabetical order.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Roman Kenk, 1898–

Revised List of the North American Freshwater Planarians (Platyhelminthes, Tricladida, Paludicola).

(Smithsonian contributions to zoology; no. 476)

Supt. of Docs. no.: SI 1.27:476

Bibliography p.

1. Planaria—North America—Classification. 2. Freshwater invertebrates—North America—Classification. 3. Platyhelminthes—North America—Classification. I. Title II. Series.

QL1.S54 no. 476 591 s [595.1′23097] 88-600390 [QL391.P7]

Contents

Introduction

List of Freshwater Triclads

Appendix

Literature Cited

Requirements for Smithsonian Series Publication

Revised List of the North American Freshwater Planarians (Platyhelminthes: Tricladida: Paludicola)

Roman Kenk

____________

Roman Kenk (deceased), Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Introduction

In 1972 I published a review of the freshwater planarians of North America north of Mexico, comprising 56 species and subspecies known at that time (Kenk, 1972). In the meantime the study of this animal group has proceeded at a rapid pace. Some of the taxa previously considered to be doubtful have been resolved, and additional species have been discovered and described. The present list does not repeat all the information given in the previous review, but intends to point out the necessary changes and to supply the recent additions to the North American fauna. It also extends the geographic area previously covered to include all the West Indies (Greater and Lesser Antilles) and all continental America north of the Isthmus of Panama.

Similar reviews of the freshwater triclads have been published recently for other areas of the globe: for Africa by Young (1976) and for Europe by Dahm and Gourbault (1978).

The following data are given for each species in the present list: the accepted name, synonyms when first used in the literature, known area of occurrence, and the principal descriptive literature that is helpful for the identification of the species. Purely distributional, ecological, physiological, or karyological records are omitted. Much of the information in this list is contained also in the first review (Kenk, 1972) and in the Index of Genera and Species of the World (Kenk, 1974). It is repeated here to make the list more useful to students of this group of flatworms.

List of Freshwater Triclads

Order TRICLADIDA

Suborder PALUDICOLA

Family DUGESIIDAE

Genus Cura Strand, 1942

1.

Cura foremanii (Girard, 1852)
Dugesia foremanii Girard, 1852

Planaria foremanii.—Stimpson, 1857

Planaria simplissima Curtis, 1900

Planaria simplicissima.—Morgan, 1904

Planaria lugubris.—Morgan, 1901 [not O. Schmidt, 1861]

Curtisia simplicissima.—Graff, 1916

Curtisia foremanii.—Kenk, 1930

Dugesia (Cura) foremanii.—Ball, 1974

?Planaria gonocephala.—Pearl, 1903 [not Dugès, 1830]

?Dugesia modesta Girard, 1893

Eastern and central North America.

Curtis, 1900:447–462; Stevens, 1904:208–218; Hyman, 1931b:322–323; Kenk, 1935:80–83, 1944:10–13, 1972:19; Anderson and Johann, 1958:375–383; Kawakatsu and Mitchell, 1982.

Genus Dugesia Girard, 1850

Subgenus Girardia Ball, 1974

2.

Dugesia (G.) tigrina (Girard, 1850)
Planaria maculata Leidy, 1847 [not Fabricius, 1826, not Darwin, 1844]

Planaria tigrina Girard, 1850

Dugesia maculata.—Girard, 1851

Phagocata tigrina.—Diesing, 1862

Planaria lata Sivickis, 1923

Euplanaria maculata.—Kenk, 1930

Euplanaria lata.—Kenk, 1930

Euplanaria novangliae Hyman, 1931

Euplanaria tigrina.—Kenk, 1935

?Euplanaria microbursalis Hyman, 1931

?Dugesia microbursalis.—Hyman, 1939

United States, southern Canada, Mexico.

Curtis, 1902; Sivickis, 1923:114–119; Hyman, 1939b:266–273; Kenk, 1935:83–87, 1944:13–18, 1972:20–21; Kawakatsu and Mitchell, 1981b.

3.

Dugesia (G.) dorotocephala (Woodworth, 1897)
Planaria dorotocephala Woodworth, 1897

Planaria agilis Stringer, 1909

Euplanaria dorotocephala.—Kenk, 1930

Euplanaria agilis.—Kenk, 1930

Euplanaria philadelphica Hyman, 1931

Dugesia agilis.—Hyman, 1939

Dugesia diabolis Hyman, 1956

United States, southern Canada, Mexico.

Woodworth, 1897:7–8; Hyman, 1925:51–80, 1929:406–413, 1931c:336–338; Kenk, 1944:18–21, 1972:22; Kawakatsu and Mitchell, 1981d.

4.

Dugesia (G.) arizonensis Kenk, 1975
Arizona.

Kenk, 1975a.

5.

Dugesia (G.) jenkinsae Benazzi and Gourbault, 1977
Texas.

Benazzi and Gourbault, 1977.

6.

Dugesia (G.) azteca Benazzi and Giannini, 1971
Cura azteca Benazzi and Giannini, 1971

Dugesia (Cura) azteca.—Ball, 1974

Mexico.

Benazzi and Giannini, 1971; Kawakatsu and Mitchell, 1984a.

7.

Dugesia (G.) typhlomexicana Mitchell and Kawakatsu, 1973
Here possibly also D. barbarae. Mexico.

Mitchell and Kawakatsu, 1972[1973]:642–646.

8.

Dugesia (G.) barbarae Mitchell and Kawakatsu, 1973
Possibly identical with D. typhlomexicana. Mexico.

Mitchell and Kawakatsu, 1972[1973]:646–649.

9.

Dugesia (G.) guatemalensis Mitchell and Kawakatsu, 1973
Mexico.

Mitchell and Kawakatsu, 1972[1973]:649–652; Kawakatsu and Mitchell, 1981c.

10.

Dugesia (G.) mckenziei Mitchell and Kawakatsu, 1973
Mexico.

Mitchell and Kawakatsu, 1973.

11.

Dugesia (G.)festai (Borelli, 1898)
Planaria festae Borelli, 1898

Dugesia festae.—Hyman, 1939

Dugesia festai.—Marcus, 1946

Widely distributed in South America, but occurs also in Curaçao.

Marcus, 1960:44–50; Kawakatsu and Mitchell, 1984c.

12.

Dugesia (G.) aurita (Kennel, 1888)
Planaria aurita Kennel, 1888

Species inquirenda. Trinidad.

Kennel, 1888:464–468; Ball, 1971:9–13.

13.

Dugesia (G.) arimana Hyman, 1957
Lesser Antilles (Trinidad, Saint Vincent, Santa Lucia). Hyman, 1957:2–4; Ball, 1971:3–9; Kawakatsu and Mitchell, 1984d.

14.

Dugesia (G.) miltgeni Gourbault, 1980
Lesser Antilles (Guadeloupe, Dominica, Martinique). Gourbault, 1980.

15.

Dugesia (G.) antillana Kenk, 1941
Puerto Rico.

Kenk, 1941; Ball, 1971:9; Codreanu and Balcesco, 1973:77–78.

16.

Dugesia (G.) cubana Codreanu and Balcesco, 1973
Cuba.

Codreanu and Balcesco, 1973:71–76; Gourbault, 1979.

Subgenus Schmidtea Ball, 1974

17.

Dugesia (S.) polychroa (Schmidt, 1861)
Dugesia lugubris.—Ball. 1969

This European species has been introduced to the St Lawrence River system (eastern Canada and northeastern United States).

Ball, 1969; Kenk, 1972:24.

Family PLANARIIDAE

Genus Phagocata Leidy, 1847

A detailed classification of this large and polymorphic genus, distributed over the entire northern hemisphere, has not been worked out (see Ball and Reynoldson, 1981:82).

18.

Phagocata velata (Stringer, 1909)
Planaria velata Stringer, 1909

Fonticola velata.—Kenk, 1930

Phagocata gracilis monopharyngea Hyman, 1948

Fonticola gracilis monopharyngea.—Ball, 1969

Phagocata monopharyngea.—Kenk, 1970

Phagocata vernalis Kenk, 1944, in part

Fonticola vernalis.—Ball, 1969, in part

Eastern and midcentral United States and Canada.

Stringer, 1909:258–259; Castle, 1928; Castle and Hyman, 1934; Kenk, 1944:28–35, 1972:35; Ball et al., 1981:9–15.

19.

Phagocata fawcetti Ball and Gourbault, 1975
California.

Ball and Gourbault 1975.

20.

Phagocata virilis Kenk, 1977
Maryland.

Kenk, 1977a:648–650.

21.

Phagocata bulbosa Kenk, 1970
North Carolina and Maryland.

Kenk, 1970a: 14–19, 1972:36.

22.

Phagocata crenophila Carpenter, 1969
Utah, California, New Mexico, and Nevada(?).

Carpenter, 1969; Kenk, 1970a: 19–23, 1972:39.

23.

Phagocata tahoena Kawakatsu, 1968
Phagocata nivea tahoena Kawakatsu, 1968

California and Nevada (Lake Tahoe).

Kawakatsu, 1968:6–11, 1973:67–69, 72–74; Kenk, 1970a:23–29, 1972:38.

24.

Phagocata hamptonae Kenk, 1982
Nevada.

Kenk, 1982.

25.

Phagocata oregonensis Hyman, 1963
Fonticola oregonensis.—Ball, 1969

Oregon and Washington.

Hyman, 1963:2–4; Kenk, 1970a:29–31, 1972:32.

26.

Phagocata nivea Kenk, 1953
Phagocata nivea nivea.—Kawakatsu, 1968

Fonticola nivea nivea.—Ball, 1969

Alaska.

Kenk, 1953:164–168, 1972:42.

27.

Phagocata morgani morgani (Stevens and Boring, 1906)
Planaria truncala Leidy [not Abildgaard, 1789]

Dendrocoelum truncatum.—Girard, 1893

Fonticola truncata.—Hyman, 1931

Planaria morgani Stevens and Boring, 1906

Fonticola morgani.—Castle and Hyman, 1934

?Dendrocoelum superbum Girard, 1850

?Galeocephala superba.—Stimpson, 1857

?Phagocata cavernicola Hyman, 1954

?Fonticola cavernicola.—Ball, 1969

Eastern half of the United States and Canada, also introduced to California.

Stevens and Boring, 1906:7–8; Stringer, 1918:338; Hyman, 1931b:328; Kenk, 1935:92–105, 1944:33–36, 1972:40.

28.

Phagocata morgani polycelis (Kenk, 1935)
Fonticola morgani polycelis Kenk, 1935

Virginia and Ontario.

Kenk, 1935:103–105, 1972:41.

29.

Phagocata spuria Kenk, 1987
North Carolina.

Kenk, 1987:669–671.

30.

Phagocata angusta Kenk, 1977
West Virginia.

Kenk, 1977a:645–648.

31.

Phagocata holleri Kenk, 1979
North Carolina.

Kenk, 1979a, 1979b:94.

32.

Phagocata carolinensis Kenk, 1979
North Carolina.

Kenk, 1979b, 1987:671–672.

33.

Phagocata pygmaea Kenk, 1987
North Carolina.

Kenk, 1987:667–669.

34.

Phagocata procera Kenk, 1984
North Carolina.

Kenk, 1984:209–213, 1987:667.

35.

Phagocata bursaperforata Darlington, 1959
Fonticola bursaperforata.—Ball, 1969

Georgia and Alabama.

Darlington 1959:276–284; Kenk, 1972:37.

36.

Phagocata notorchis Kenk, 1987
Tennessee.

Kenk, 1987:664–667.

37.

Phagocata gracilis (Haldeman, 1840)
Planaria gracilis Haldeman, 1840

Phagocata gracilis.—Leidy, 1847

Euplanaria gracilis.—Kenk, 1930

Fonticola gracilis.—Castle and Hyman, 1934

Phagocata subterranea Hyman, 1937

Fonticola subterranea.—Ball, 1969

Eastern United States north to Maryland, west to Missouri. Kenk, 1935:88–92, 1970c:2–10, 1972:43; Hyman, 1937a:302–305.

38.

Phagocata woodworthi Hyman, 1937
Phagocata gracilis woodworthi Hyman, 1951

Fonticola gracilis woodworthi.—Ball, 1969

Northeastern United States south to Maryland, eastern Canada as far west as Ontario.

Woodworth, 1891 (misidentified as P. gracilis); Hyman, 1937a:305–306; Kenk, 1970c: 10–14, 1972:44.

39.

Phagocata nordeni Kenk, 1977
Florida, Louisiana, Mississippi.

Kenk, 1977a:650–652.

Genus Hymanella Castle, 1941

40.

Hymanella retenuova Castle, 1941
Phagocata vernalis Kenk, 1944, in part

Eastern and midwestern United States and Ontario. Castle, 1941:85–95; Kenk, 1944:22–28 (as P. vernalis), 1972:31; Hyman, 1955:30–32; Ball et al., 1981:3–9. Genus Paraplanaria Ball and Gourbault, 1978

41.

Paraplanaria dactyligera (Kenk, 1935)
Planaria dactyligera Kenk, 1935

Planaria dactyligera dactyligera.—Kenk, 1969

Planaria dactyligera musculosa Kenk, 1969

The subspecies musculosa shows transitions to the nominate subspecies without geographic differentiation, and should therefore be withdrawn. Eastern United States (Virginia, Maryland, Tennessee, North Carolina, Louisiana).

Kenk, 1935:105–110, 1969:541–549, 1972:28–29; Ball and Gourbault, 1978:31–34.

42.

Paraplanaria occulta (Kenk, 1969)
Planaria occulta Kenk, 1969

Virginia.

Kenk, 1969:549–556, 1972:30.

Genus Polycelis Ehrenberg, 1831

Subgenus Polycelis Ehrenberg, 1831

43.

Polycelis (P.) coronata coronata (Girard, 1891)
Phagocata coronata Girard, 1891

Polycelis borealis Kenk, 1953

Polycelis coronata borealis Kenk, 1972

The species or subspecies borealis is not clearly separated from P. coronata; as many transitional forms exist. I therefore combine the two species, retaining the older name coronata. Western United States (Wyoming, Utah, Montana, New Mexico, Washington, Oregon, Alaska, Idaho) and Canada (Alberta).

Kenk, 1953:168–173, 1972:25, 27, 1973a:3–8.

44.

Polycelis (P.) coronata monticola Kenk and Hampton, 1982
Polycelis monticola Kenk and Hampton, 1982

California and Nevada.

Kenk and Hampton, 1982

45.

Polycelis (P.) coronata brevipenis Kenk, 1972
Polycelis coronata Hyman, 1931

South Dakota and Colorado.

Hyman, 1931a:124–131; Kenk, 1952:194–197, 1972:26, 1973a: 8–9.

Subgenus Seidlia Zabusov, 1911

46.

Polycelis (S.) sierrensis Kenk, 1973
California and Nevada.

Kenk, 1973a:9–12.

Family DENDROCOELIDAE

Subfamily KENKIINAE

Genus Sphalloplana Beauchamp, 1931

Subgenus Sphalloplana Beauchamp, 1931

47.

Sphalloplana (Sph.) percoeca (Packard, 1879)
Dendrocoelum percoecum Packard, 1879

Fonticola percoecum.—Hyman, 1931

Fonticola percaeca.—Hyman, 1935

Sphalloplana alabamensis Hyman, 1945

Kentucky, Alabama, Tennessee, and West Virginia. Beauchamp, 1931:317–330, in part; Hyman, 1937b:467–471, 1951:163; Buchanan, 1936; Kenk, 1972:45, 1977b:3–5.

48.

Sphalloplana (Sph.) georgiana Hyman, 1954
Somewhat problematic species. Georgia. Hyman, 1954:566–570; Kenk, 1972:47, 1977b:5–6.

49.

Sphalloplana (Sph.) evaginata Kenk, 1977
Missouri.

Kenk, 1977b:6–8.

50.

Sphalloplana (Sph.) kansensis Hyman, 1945
Species not well defined. Kansas.

Hyman, 1945:478–479, 1951:164; Kenk, 1972:49, 1977b:8.

51.

Sphalloplana (Sph.) californica Kenk, 1977
California.

Kenk, 1977b:8–10.

52.

Sphalloplana (Sph.) culveri Kenk, 1977
West Virginia.

Kenk, 1977b: 10–11.

53.

Sphalloplana (Sph.) consimilis Kenk, 1977
Virginia and Tennessee.

Kenk, 1977b: 11–12.

54.

Sphalloplana (Sph.) subtilis Kenk, 1977
Virginia (now possibly extinct).

Kenk, 1977b: 12–14.

Subgenus Speophila Hyman, 1937

55.

Sphalloplana (Sp.) virginiana Hyman, 1945
Species inquirenda. Virginia.

Hyman, 1945:477–478; Kenk, 1972:48, 1977b: 14.

56.

Sphalloplana (Sp.) holsingeri Kenk, 1977
Virginia (now possibly extinct).

Kenk, 1977b: 15–17.

57.

Sphalloplana (Sp.) weingartneri Kenk, 1970 Indiana.
Kenk, 1970b:314–319, 1972:54, 1977b:17.

58.

Sphalloplana (Sp.) buchanani (Hyman, 1937)
Speophila buchanani Hyman, 1937

Kentucky.

Hyman, 1937b:468–469; Kenk, 1972:51, 1977b:17–18.

59.

Sphalloplana (Sp.) pricei (Hyman, 1937)
Speophila pricei Hyman, 1937

Pennsylvania.

Hyman, 1937b:462–466; Kenk, 1972:50, 1977b:18–19.

60.

Sphalloplana (Sp.) hubrichti (Hyman, 1945)
Speophila hubrichti Hyman, 1945

Missouri and Illinois.

Hyman, 1945:479–480; Kenk, 1972:52, 1977b:20–21.

61.

Sphalloplana (Sp.) chandleri Kenk, 1977
Tennessee, Indiana, Virginia.

Kenk, 1977b:21–23.

62.

Sphalloplana (Sp.) hypogea Kenk, 1984 Virginia.
Kenk, 1984:213–215.

63.

Sphalloplana (Sp.) hoffmasteri (Hyman, 1954)
Speophila hoffmasteri Hyman, 1954

Macrocotyla hoffmasteri.—Kenk, 1975

Kenkia hoffmasteri.—Kawakatsu and Mitchell, 1981

The pharyngeal musculature does not quite conform with that of the genus Kenkia, which places the species back into the genus Sphalloplana. West Virginia and Maryland. Hyman, 1954:570–573; Kenk, 1972:53, 1975b:333–336.

Subgenus Polypharyngea Carpenter, 1971

64.

Sphalloplana (P.) mohri Hyman, 1938
Sphalloplana kutscheri Mitchell, 1968

Sphalloplana sloani Mitchell, 1968

Sphalloplana zeschi Mitchell, 1968

Sphalloplana reddelli Mitchell, 1968

Texas.

Hyman, 1939c:276–280 ; Mitchell, 1968:598–610, 1974:402–429; Kenk, 1972:55, 1977b:23–24.

Genus Kenkia Hyman, 1937

65.

Kenkia rhynchida Hyman, 1937
Oregon.

Hyman, 1937b:457–462; Kenk, 1972:56; Kawakatsu and Mitchell, 1981a:126–135.

66.

Kenkia glandulosa (Hyman, 1956)
Macrocotyla glandulosa Hyman, 1956

Missouri.

Hyman, 1956:9–14; Kenk, 1972:57, 1975b:324–329.

67.

Kenkia lewisi (Kenk, 1975)
Macrocotyla lewisi Kenk, 1975

Missouri.

Kenk, 1975b:329–333.

Subfamily DENDROCOELINAE

Genus Dendrocoelopsis Kenk, 1930

68.

Dendrocoelopsis vaginata Hyman, 1935
Western United States (Montana, Oregon, Washington). Hyman, 1935:339–345; Kenk, 1972:58, 1973b:2–6.

69.

Dendrocoelopsis piriformis Kenk, 1953
Alaska and western Canada.

Kenk, 1953:174–178, 1972:59, 1973b:6–8; Holmquist, 1967:453–465.

70.

Dendrocoelopsis alaskensis Kenk, 1953
Alaska.

Kenk, 1953:178–181, 1972:60, 1973b:8–9.

71.

Dendrocoelopsis hymanae Kawakatsu, 1968
California and Nevada (Lake Tahoe).

Kawakatsu, 1968:11–15; Kenk, 1972:61, 1973b:9–10.

72.

Dendrocoelopsis americana (Hyman, 1939)
Sorocelis americana Hyman, 1939

Oklahoma, Arkansas, Texas, Missouri.

Hyman, 1939a:422–424, 1939c:280–283; Kenk, 1972:62, 1973b:10–14; Kawakatsu and Mitchell, 1984b.

Genus Procotyla Leidy, 1857

73.

Procotyla fluviatilis Leidy, 1857
Dendrocoelum lacteum.—misidentified by early American authors

Dendrocoelum graffi Wilhelmi, 1909

?Dendrocoelum pulcherrimum Girard, 1850

?Oligocelis pulcherrima.—Stimpson, 1857

?Procotyla leidyi Girard, 1893

Eastern and midwestern United States and Canada.

Woodworth, 1897:2–6 (as Dendrocoelum lacteum); Hyman, 1928:222–245; Kenk, 1944:36–38, 1972:63.

74.

Procotyla typhlops Kenk, 1935
Virginia, Maryland, Florida.

Kenk, 1935:111–117, 1972:64; Hyman, 1951:165–166.

Genus Rectocephala Hyman, 1953

75.

Rectocephala exotica Hyman, 1953
Bdellocephala exotica.—Kenk, 1972

Species inquirenda. Introduced to aquatic gardens in Washington, District of Columbia.

Hyman, 1953; Kenk, 1972:65.

Species Incertae Sedis

76.

Planaria fissipara Kennel, 1888
Not observed in a sexually mature state. Apparently widely distributed in South America, but recorded also from Trinidad and probably Mexico.

Kennel, 1888:468–473; Hyman, 1957:4–5; Marcus, 1948:176–177.

Not Recognizable Species

77.

Hydrolimax bruneus Girard, 1891
This may be a triclad. Pennsylvania (Fairmount Park, Philadelphia).

Girard, 1893:163–164; Kenk, 1972:67.

78.

Planaria fuliginosa Leidy, 1851
Planaria (Typhlolepta?) fuliginosus Leidy, 1851

Anocelis fuliginosa.—Stimpson, 1857

Typhloplana(?) fuliginosa.—Leuckart, 1854

New Jersey (Rancocas Creek, Burlington County).

Leidy, 1851:225; Kenk, 1972:66.

79.

Planaria simplex Woodworth, 1896
Lake Michigan.

Woodworth, 1896a:94, 1896b:239–240; Kenk, 1972:66.

80.

Planaria unionicola Woodworth, 1897
Illinois (Illinois River near Havana).

Woodworth, 1897; Kenk, 1972:66.

Appendix

Names of Species and Subspecies

(includes valid names and synonyms; numbers refer to numbered paragraphs in text)

agilis, 3

alabamensis, 47

alaskensis, 70

americana, 72

angusta, 30

antillana, 15

arimana, 13

arizonensis, 4

aurita, 12

azteca, 6

barbarae, 7, 8

borealis, 43

brevipenis, 45

bruneus, 77

buchanani, 58

bulbosa, 21

bursaperforata, 35

californica, 51

carolinensis, 32

cavernicola, 27

chandleri, 61

consimilis, 53

coronata, 43, 44, 45

crenophila, 22

cubana, 16

culveri, 52

dactyligera, 41

diabolis, 3

dorotocephala, 3

evaginata, 49

exotica, 75

fawcetti, 19

festae or festai, 11

fissipara, 76

fluviatilis, 73

foremanii, 1

fuliginosa or -us, 78

georgiana, 48

glandulosa, 66

gonocephala, 1

gracilis, 18, 37, 38

graffi, 73

guatemalensis, 9

hamptonae, 24

hoffmasteri, 63

holleri, 31

holsingeri, 56

hubrichti, 60

hymanae, 71

hypogea, 62

jenkinsae, 5

kansensis, 50

kutscheri, 64

lacteum, 73

lata, 2

leidyi, 73

lewisi, 67

lugubris, 1, 17

maculata, 2

mckenziei, 10

microbursalis, 2

miltgeni, 14

modesta, 1

mohri, 64

monopharyngea, 18

monticola, 44

morgani, 27, 28

musculosa, 41

nivea, 23, 26

nordeni, 39

notorchis, 36

novangliae, 2

occulta, 42

oregonensis, 25

percoeca or -um, 47

philadelphica, 3

piriformis, 69

polycelis, 28

polychroa, 17

pricei, 59

procera, 34

pulcherrima or -um, 73

pygmaea, 33

reddelli, 64

retenuova, 40

rhynchida, 65

sierrensis, 46

simplex, 79

simplicissima, 1

simplissima, 1

sloani, 64

spuria, 29

subterranea, 37

subtilis, 54

superba or -um, 27

tahoena, 23

tigrina, 2

truncata or -um, 27

typhlomexicana, 7, 8

typhlops, 74

unionicola, 80

vaginata, 68

velata, 18

vernalis, 18, 40

virginiana, 55

virilis, 20

weingartneri, 57

woodworthi, 38

zeschi, 64

Literature Cited

(Papers mentioned only in the list of synonyms of individual species are not entered here; most of these may be found in the bibliography of Kenk, 1974.)

Anderson, J.M., and J.C. Johann

1958.   Some Aspects of Reproductive Biology in the Fresh-Water Triclad Turbellarian Cura foremanii. Biological Bulletin, 115:375–383.

Ball, I.R.

1969.   Dugesia lugubris (Tricladida: Paludicola), a European Immigrant into North American Fresh Waters. Journal of the Fisheries Research Board of Canada, 26:221–228.

1971.   Systematic and Biogeographical Relationships of Some Dugesia Species (Tricladida, Paludicola) from Central and South America. American Museum Novitates, 2472: 25 pages.

Ball. I.R., and N. Gourbault

1975.   The Morphology, Karyology and Taxonomy of a New Freshwater Planarian of the Genus Phagocata from California (Platyhelminthes: Turbellaria). Life Sciences Contributions, Royal Ontario Museum, 105: iii + 19 pages.

1978.   The Phyletic Status of the Genus Planaria (Platyhelminthes, Turbellaria, Tricladida). Bijdragen tot de Dierkunde, 48:29–34.

Ball, I.R., N. Gourbault, and R. Kenk

1981.   The Planarians (Turbellaria) of Temporary Waters in Eastern North America. Life Sciences Contributions, Royal Ontario Museum, 127: iii + 27 pages.

Ball, I.R., and T.B. Reynoldson

1981.   British Planarians (Platyhelminthes: Tricladida); Keys and Notes for the Identification of the Species. 141 pages. Cambridge: Cambridge University Press.

Beauchamp, P. de

1931.   Biospeologica, LVI: Campagne spéologique de C. Bolivar et R. Jeannel dans l'Amérique du Nord (1928), 2: Turbellariés Triclades. Archives de Zoologie Expérimentale et Générale, 71:317–331.

Benazzi, M., and E. Giannini

1971.   Cura azteca, nuova specie di planaria del Messico. Atti della Accademia Nazionale dei Lincei, Rendiconti della Classe di Scienze Fisiche Matematiche e Naturali, series 8, 50:477–481.

Benazzi, M., and N. Gourbault

1977.   Dugesia jenkinsae n. sp., a Fresh-Water Triclad (Turbellaria) from Texas. Transactions of the American Microscopical Society, 96:540–543.

Buchanan, J.W.

1936.   Notes on an American Cave Flatworm, Sphalloplana percaeca (Packard). Ecology, 17:194–211.

Carpenter, J. H.

1969.   A New Planarian from Utah, Phagocata crenophila n. sp. (Turbellaria). Transactions of the American Microscopical Society, 88:274–281.

Castle, W.A.

1928.   An Experimental and Histological Study of the Life-Cycle of Planaria velata. Journal of Experimental Zoology, 51:417–483.

1941.   The Morphology and Life History of Hymanella retenuova, a New Species of Triclad from New England. American Midland Naturalist, 26:85–97.

Castle, W.A., and LH. Hyman

1934.   Observations on Fonticola velata (Stringer), Including a Description of the Anatomy of the Reproductive System. Transactions of the American Microscopical Society, 53:154–171.

Codreanu, R., and D. Balcesco

1973.   Dugesia cubana n. sp., Planaire nouvelle de l'île de Cuba et ses affinités sur-américaines. In Résultats des expéditions biospéologiques cubano-roumaines à Cuba, 1:71–80, 4 plates. Bucuresti: Academia Republicii Socialiste România.

Curtis, W.C.

1900.   On the Reproductive System of Planaria simplissima, a New Species. Zoologische Jahrbücher, Abtheilung für Anatomie und Ontogenie der Thiere, 13:447–466, plates 31, 32.

1902.   The Life History, the Normal Fission, and the Reproductive Organs of Planaria maculata. Proceedings of the Boston Society of Natural History, 30:515–559, plates 9–19.

Dahm, A.G., and N. Gourbault

1978.   Tricladida et Temnocephalida (Turbellaria). In J. Illies, editor, Limnofauna Europaea, 2nd edition, pages 16–20.

Darlington, J.T.

1959.   The Turbellaria of Two Granite Outcrops in Georgia. American Midland Naturalist, 61:257–294.

Girard, C.

1852.   Descriptions of Two New Genera and Two New Species of Planaria. Proceedings of the Boston Society of Natural History, 4:210–212.

1891.   Deux espèces nouvelles de Planaires américaines. Le Naturaliste, 13:80.

1893.   Recherches sur les Planariés et les Némertiens de l'Amérique du Nord. Annales des Sciences Naturelles, Zoologie, series 7, 15:145–310, plates 3–6.

Gourbault, N.

1977.   Karyology of Dugesia arizonensis Kenk (Turbellaria, Tricladida). Caryologia, 30:63–68.

1979.   Données biologiques et cytotaxonomiques sur un Triclade de l'îsle de Cuba. Archives de Zoologie Expérimentale et Générale, 120:131–142.

1980.   Morphologie et cytotaxonomie d'une Planaire nouvelle de Guadeloupe (Turbellarié, Triclade). Bulletin du Muséum National d'Histoire Naturelle (Paris), series 2, section A, 3:751–757.

Holmquist, C.

1967.   Dendrocoelopsis piriformis (Turbellaria Tricladida) and Its Parasites from Northern Alaska. Archiv für Hydrobiologie, 62:453–466, 6 plates.

Hyman, L.H.

1925.   The Reproductive System and Other Characters of Planaria dorotocephala Woodworth. Transactions of the American Microscopical Society, 44:51–89.

1928.   Studies on the Morphology, Taxonomy, and Distribution of North American Triclad Turbellaria, I: Procotyla fluviatilis, Commonly but Erroneously Known as Dendrocoelum lacteum. Transactions of the American Microscopical Society, 47:222–255.

1929.   Studies on the Morphology, Taxonomy, and Distribution of North American Triclad Turbellaria, II: On the Distinctions between Planaria agilis and Planaria dorotocephala, with Notes on the Distribution of agilis in the Western United States. Transactions of the American Microscopical Society, 48:406–415.

1931a. Studies on the Morphology, Taxonomy, and Distribution of North American Triclad Turbellaria, III: On Polycelis coronata (Girard). Transactions of the American Microscopical Society, 50:124–135.

1931b. Studies on the Morphology, Taxonomy, and Distribution of North American Triclad Turbellaria, IV: Recent European Revisions of the Triclads, and Their Application to the American Forms, with a Key to the Latter and New Notes on Distribution. Transactions of the American Microscopical Society, 50:316–335.

1931c. Studies on the Morphology, Taxonomy, and Distribution of North American Triclad Turbellaria, V: Description of Two New Species. Transactions of the American Microscopical Society, 50:336–343.

1935.   Studies on the Morphology, Taxonomy, and Distribution of North American Triclad Turbellaria, VI: A New Dendrocoelid from Montana, Dendrocoelopsis vaginatus n. sp. Transactions of the American Microscopical Society, 54:338–345.

1937a. Studies on the Morphology, Taxonomy, and Distribution of North American Triclad Turbellaria, VII: The Two Species Confused under the Name Phagocata gracilis, the Validity of the Generic Name Phagocata Leidy, 1847, and Its Priority over Fonticola Komareck, 1926. Transactions of the American Microscopical Society, 56:298–310.

1937b. Studies on the Morphology, Taxonomy, and Distribution of North American Triclad Turbellaria, VIII: Some Cave Planarians of the United States. Transactions of the American Microscopical Society, 56:457–477.

1939a. New Species of Flatworms from North, Central, and South America. Proceedings of the United States National Museum, 86:419–439.

1939b. North American Triclad Turbellaria, IX: The Priority of Dugesia Girard 1850 over Euplanaria Hesse 1897, with Notes on American Species of Dugesia. Transactions of the American Microscopical Society, 58:264–275.

1939c. North American Triclad Turbellaria, X: Additional Species of Cave Planarians. Transactions of the American Microscopical Society, 58:276–284.

1945.   North American Triclad Turbellaria, XI: New, Chiefly Cavernicolous, Planarians. American Midland Naturalist, 34:475–484.

1951.   North American Triclad Turbellaria, XII: Synopsis of the Known Species of Fresh-Water Planarians of North America. Transactions of the American Microscopical Society, 70:154–167.

1953.   North American Triclad Turbellaria, 14: A New, Probably Exotic, Dendrocoelid. American Museum Novitates, 1629: 6 pages.

1954.   North American Triclad Turbellaria, XIII: Three New Cave Planarians. Proceedings of the United States National Museum, 103:563–573.

1955.   Descriptions and Records of Fresh-Water Turbellaria from the United States. American Museum Novitates, 1714: 36 pages.

1956.   North American Triclad Turbellaria, XV: Three New Species. American Museum Novitates, 1808: 14 pages.

1957.   A Few Turbellarians from Trinidad and the Canal Zone, with Corrective Remarks. American Museum Novitates, 1862: 8 pages.

1963.   North American Triclad Turbellaria, XVI: Fresh-Water Planarians from the Vicinity of Portland, Oregon. American Museum Novitates, 2123: 5 pages.

Kawakatsu M.

1968.   North American Triclad Turbellaria, 17: Freshwater Planarians from Lake Tahoe. Proceedings of the United States National Museum, 124(3638):1–21, 2 plates.

1973.   Additional Notes on Freshwater Planarians from Lake Tahoe in the Sierra Nevada Mountains of North America. Bulletin of Fuji Women's College, series 2, 11:67–77.

Kawakatsu, M., and R.W. Mitchell

1981a. Redescription of Kenkia rhynchida, a Troglobitic Planarian from Oregon, and a Reconsideration of the Family Kenkiidae and Its Genera (Turbellaria, Tricladida, Paludicola). Annotationes Zoologicae Japonenses, 54:125–141.

1981b. A Note on the Morphology of Dugesia tigrina (Girard, 1850) from Louisiana, U.S.A. Proceedings of the Japanese Society of Systematic Zoology, 20:1–6, 1 plate.

1981c. An Additional Note on Dugesia guatemalensis Mitchell et Kawakatsu (Turbellaria, Tricladida, Paludicola), a Troglophilic Planarian from Mexico. Journal of the Speleological Society of Japan, 6:37–41.

1981d. Freshwater Planarians from the Southern United States and Mexico: Dugesia dorotocephala and Dugesia sp. (Turbellaria, Tricladida, Paludicola). Annotationes Zoologicae Japonenses, 54:191–206.

1982.   A Note on the Morphology of Cura foremanii (Girard, 1852) from Four Localities, U.S.A. (Turbellaria, Tricladida, Paludicola). Bulletin of Fuji Women's College, series 2, 20:91–100.

1984a. Redescription of Dugesia azteca (Benazzi et Giannini, 1971) Based upon the Material Collected from the Type Locality in Mexico, with Corrective Remarks (Turbellaria, Tricladida, Paludicola). Bulletin of the National Science Museum (Tokyo), series A (Zoology), 10:37–50.

1984b. Occurrence of Dendrocoelopsis americana (Hyman, 1939) in Soda Water Well, Texas, U.S.A. (Turbellaria, Tricladida, Paludicola). Proceedings of the Japanese Society of Systematic Zoology, 28:1–11, plates 1–3.

1984c. Redescription of Dugesia festai (Borelli, 1898) Based upon Material from Venezuela and Peru (Turbellaria; Tricladida; Paludicola). Bulletin of the Biogeographical Society of Japan, 39(1):1–12.

1984d. Redescription of Dugesia arimana Hyman, 1957, Based upon Material from Trinidad, St. Vincent and Venezuela (Turbellaria, Tricladida, Paludicola). Bulletin of Fuji Women's College, series 2, 22:63–77.

Kenk, R.

1935.   Studies on Virginian Triclads. Journal the Elisha Mitchell Scientific Society, 51:79–133, 8 plates.

1941.   A Fresh-Water Triclad from Puerto Rico, Dugesia antillana, New Species. Occasional Papers of the Museum of Zoology, University of Michigan, 436: 8 pages, 1 plate.

1944.   The Fresh-Water Triclads of Michigan. Miscellaneous Publications of the Museum of Zoology, University of Michigan, 60: 44 pages, 7 plates.

1952.   Fresh-Water Triclads (Turbellaria) of the Rocky Mountain National Park Region, Colorado. Journal of the Washington Academy of Sciences, 42:193–198.

1953.   The Fresh-Water Triclads (Turbellaria) of Alaska. Proceedings of the United States National Museum, 103:163–186, plates 6–8.

1969.   Freshwater Triclads (Turbellaria) of North America, I: The Genus Planaria. Proceedings of the Biological Society of Washington, 82:539–558.

1970a. Freshwater Triclads (Turbellaria) of North America, II: New or Little Known Species of Phagocata. Proceedings of the Biological Society of Washington, 83:13–34.

1970b. Freshwater Triclads (Turbellaria) of North America, III: Sphalloplana weingartneri, New Species, from a Cave in Indiana. Proceedings of the Biological Society of Washington, 83:313–320.

1970c. Freshwater Triclads (Turbellaria) of North America, IV: The Polypharyngeal Species of Phagocata. Smithsonian Contributions to Zoology, 80:1–17.

1972.   Freshwater Planarians (Turbellaria) of North America. Biota of Fresh-water Ecosystems, Identification Manual, 1: ix + 81 pages. Washington: Environmental Protection Agency. (Also second printing, 1976, Cincinnati, Ohio.)

1973a. Freshwater Triclads (Turbellaria) of North America, V: The Genus Polycelis. Smithsonian Contributions to Zoology, 135:1–15.

1973b. Freshwater Triclads (Turbellaria) of North America, VI: The Genus Dendrocoelopsis. Smithsonian Contributions to Zoology, 138:1–16.

1974.   Index of the Genera and Species of the Freshwater Triclads (Turbellaria) of the World. Smithsonian Contributions to Zoology, 183:1–90.

1975a. Freshwater Triclads (Turbellaria) of North America, VIII: Dugesia arizonensis, New Species. Proceedings oAf the Biological Society of Washington, 88:113–120.

1975b. Freshwater Triclads (Turbellaria) of North America, VII: The Genus Macrocotyla. Transactions of the American Microscopical Society, 94:324–339.

1977a. Freshwater Triclads (Turbellaria) of North America, X: Three New Species of Phagocata from the Eastern United States. Proceedings of the Biological Society of Washington, 89:645–652.

1977b. Freshwater Triclads (Turbellaria) of North America, IX: The Genus Sphalloplana. Smithsonian Contributions to Zoology, 246:1–38.

1979a. Freshwater Triclads (Turbellaria) of North America, XI: Phagocata holleri, New Species, from a Cave in North Carolina. Proceedings of the Biological Society of Washington, 92:389–393.

1979b. Freshwater Triclads (Turbellaria) of North America, XII: Another New Cave Planarian from North Carolina. Brimleyana, 2:91–96.

1982.   Freshwater Triclads (Turbellaria) of North America, XIII: Phagocata hamptonae, New Species, from Nevada. Proceedings of the Biological Society of Washington, 95:161–166.

1984.   Freshwater Triclads (Turbellaria) of North America, XV: Two New Subterranean Species from the Appalachian Region. Proceedings of the Biological Society of Washington, 97:209–216.

1987.   Freshwater Triclads (Turbellaria) of North America, XVI: More on Subterranean Species of Phagocata of the Eastern United States. Proceedings of the Biological Society of Washington, 100:664–673.

Kenk, R., and A.M. Hampton

1982.   Freshwater Triclads (Turbellaria) of North America, XIV: Polycelis monticola, New Species, from the Sierra Nevada Range in California. Proceedings of the Biological Society of Washington, 95:567–570.

Kennel, J.

1888.   Untersuchungen an neuen Turbellarien. Zoologische Jahrbücher, Abtheilung für Anatomie und Ontogenie der Thiere, 3:447–486, plates 18, 19.

Leidy, J.

1851.   Helminthological Contributions, II. Proceedings of the Academy of Natural Sciences of Philadelphia, 5:224–227.

Marcus, E.

1948.   Turbellaria do Brasil. Universidade de Säo Paulo, Boletins da Faculdade de Filosofia Ciéncias e Letras, Zoologia, 13:111–243.

1960.   Turbellaria from Curaçao. Natuurwetenschappelijke Studiekring voor Suriname en de Nederlandse Antillen, 21:41–51.

Mitchell, R.W.

1968.   New Species of Sphalloplana (Turbellaria: Paludicola) from the Caves of Texas and a Reexamination of the Genus Speophila and the Family Kenkiidae. Annales de Spéléologie, 23:597–620.

1974.   The Cave-Adapted Flatworms of Texas: Systematics, Natural History, and Responses to Light and Temperature. In N.W. Riser and M.P. Morse, editors, Biology of the Turbellaria, pages 402–430. New York: McGraw-Hill Book Company.

Mitchell, R.W., and M. Kawakatsu

1972 [1973].   Freshwater Cavernicole Planarians from Mexico; New Troglobitic and Troglophilic Dugesia from Caves of the Sierra de Guatemala. Annales de Spéléologie, 27:639–681. [Date on title page is 1972; actually published in 1973.]

1973.   A New Cave-Adapted Planarian (Tricladida, Paludicola, Planariidae) from Chiapas, Mexico. Association for Mexican Cave Studies, Bulletin, 5:165–170.

Sivickis, P.B.

1923.   Studies on the Physiology of Reconstitution in Planaria lata, with a Description of the Species. Biological Bulletin, 44:113–152.

Stevens, N.M.

1904.   On the Germ Cells and the Embryology of Planaria simplissima. Proceedings of the Academy of Natural Sciences of Philadelphia, 56:208–220, plates 13–16.

Stevens, N.M., and A.M. Boring

1906.   Planaria morgani n. sp. Proceedings of the Academy of Natural Sciences of Philadelphia, 58:7–9, plate 1.

Stringer, C.E.

1909.   Notes on Nebraska Turbellaria with Descriptions of Two New Species. Zoologischer Anzeiger, 34:257–262.

1918.   The Free-living Flatworms (Turbellaria). In H.B. Ward and G.C. Whipple, editors, Fresh-Water Biology, pages 323–364. New York:John Wiley and Sons.

Woodworth, W.M.

1891.   Contributions to the Morphology of the Turbellaria, I: On the Structure of Phagocata gracilis, Leidy. Bulletin of the Museum of Comparative Zoology at Harvard College, 21:1–44, plates 1–4.

1896a. Preliminary Report on Collections of Turbellaria from Lake St. Clair and Charlevoix, Michigan. Michigan Fish Commission, Bulletin 6:94–95.

1896b. Report on the Turbellaria Collected by the Michigan State Fish Commission during the Summers of 1893 and 1894. Bulletin of the Museum of Comparative Zoology at Harvard College, 29:239–244, 1 plate.

1897.   Contributions to the Morphology of the Turbellaria, II: On Some Turbellaria from Illinois. Bulletin of the Museum of Comparative Zoology at Harward College, 31:1–16, 1 plate.

Young, J.O.

1976.   The Freshwater Turbellaria of the African Continent. Zoologischer Anzeiger, 197:419–432.

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