THE 9"'^^
NAUTILUS
THE PILSBRY QUARTERLY
DEVOTED TO THE INTERESTS
OF CONCHOLOGISTS
VOL. 85
JULY, 1971 to APRIL, 1972
EDITORS AND PUBLISHERS
R. TUCKER ABBOTT "' '
Delaware Museum of Natural History, Greenville, Delaware ^ /
CHARLES B. WURTZ
MRS. HORACE B. BAKER
11 Chelten Road
Havertown, Pennsylvania 19083
\ \h /. C t >
h ■■
PONY PRINTING, UPPER DARBY, PA.
[Uj I LIBRARY
April, 1972 NAUTiLUsy*!;^ ^ J ^pdex) iii
CONTENTS N^ ^< ^^
Alasmidonta fabula in Kentucky 60
Amblema costata (Rafinesque), reproduction 146
Anodonta grandis Say, in Texas 144
Aplysiid opisthobranchs, feeding 37
Arionid slug from Washington state 100
Arizona, Pseudosuccinea columella in . 71
Armiger exigua Leonard, n. sp 81
Aroapyrgus colombiensis Malek and Little, n. sp 20
Baker, H. Burrington (1889-1971) 1
Biomphalaria glabrata, egg-laying 43
Cayman Islands, land mollusks 69
Charonia variegata (Lamarck), habits and food 84
Chromosomes of Spisula solidissima 93
Clappiella saludensis, type locality of 36
Conus patae Abbott, n. sp 49
Corbicula fluminea, in New Mexico 144
Corbicula manilensis (Philippi) in Oklahoma 145
Corbicula manilensis, in Texas 35
Cyphoma gibbosum, spawn Ill
Dates of The Nautilus no. 1, iii
Ecology of Charonia variegata 84
Egg laying of Biomphalaria 43
Egg laying of Cyphoma Ill
Egypt, Lake Birket Qarun 141
Endodontid snails, fluorescence in 17
Eulimastoma Bartsch, review of 51
Fasciolaria hullisi Lyons, n. sp 96
Florida, aplysiid opisthobranchs in 37
Fluorescence in endodontid snails 17
Fluorescence in Mesodon clausus (Say) 65
Food of Charonia variegata 84
Galapagos land snails 5
Gastrocopta abbreviata (Sterki) 74
Gastrocopta armifera (Say) 73
Gastrocopta clappi (Sterki) 76
Gastrocopta ruidosensis (Cockerell) 75
Gastrocopta similis (Sterki) ,..,.. 75
Gastrocopta venusta Leonard, n. sp - . 80
'-'>'',■■ 'o'-''
iv NAUTILUS Vol. 85 (Index)
Georgia, new species of Hydrobiidae 120
Goniodostomia Pilsbry and Johnson 51
Greggelix Miller, new genus 128
Gulf of Mexico, new Fasciolaria 96
Helminthoglyptidae, reproductive anatomy of 61
Hemidonax, familial affinities of 9
Hemphillia burringtoni Pilsbry 101
Hemphillia dromedarius Branson, n. sp 100
Hertlein, Leo G. (1898-1972) no. 4, iii
Illinois, Pleistocene gastropods of 78
Introduced Mollusks, to New Mexico and Texas 144
Kentucky, Alasmidonta fabula in 60
Lake Birket, Qarun, Egypt 141
Littorina irrorata, shell growth 136
Locomotion of Marginella olivaeformis 110
Malacology, research trends in 67
Marginella olivaeformis Kiener, locomotion 110
Mayer-Eymar, Karl, collection no. 4, iii
McCargo Lake, New York, gastropods 38
Melampus bidentatus (Say), distribution 106
Mesodon clausus (Say), fluorescence in 65
Minnesota, Amblema costata in 146
Mississippi, Pisidium compressum in 71
Missouri, Pleuroceridae in 26
Mudalia potosiensis (Lea), variation 26
Naesiotus achatellinus (Forbes) 5
Nassarius, reproductions 126
Neovolusia Emerson 30
New Mexico, introduced mollusks 144
Niso imbricata (Sowerby) 30
North Carolina, Strombus raninus in 72
Oklahoma, Corbicula manilensis in 145
Oklahoma, Sphaerium lacustre in 71
Otsego Lake, New York, mollusks of 70
Ovoviviparous Nassarius 126
Parasitology, Paragonimus 20
Parodostomia Laseron, review of 51
Pisidium compressum Prime, in Mississippi 71
Pleistocene gastropods of Illinois 78
April, 1972 nautilus (Index) v
Pleistocene land snails, California 32
Pleuroceridae from Missouri 26
Pseudosuccinea columella (Say), in Arizona 71
Punctum parvulum Leonard, n. sp 84
Pyramidellacea, review of 51
Radix auricularia (L.), in New Mexico 145
Reproduction in Amblema costata 146
Research trends in malacology 67
Shell growth in Littorina irrorata 136
Somatogyrus alcoviensis Krieger, n. sp 120
South Carolina, reef mollusks of 114
South Carolina, Strombus gigas in 72
Sphaerium lacustre (Miiller), in Oklahoma 71
Sphaerium simile (Say), in Tennessee no. 1, iii
Spisula solidissima (Dillwyn), chromosomes 93
Stagnicola montanensis (Baker), type locality 145
Stenotrema barbatum (Clapp) 16
Stenotrema burringtoni Grimm, n. sp 14
Stenotrema hirsutum (Say) 15
Stenotrema simile Grimm, n. sp 12
Strombus gigas L., in South Carolina 72
Strombus raninus Gmelin, in North Carolina 72
Succinea exile Leonard, n. sp 82
Telloda Hertlein and Strong, review of 51
Tennessee, Sphaerium simile in no. 1, iii
Texas, Corbicula manilensis in 35
Texas, introduced mollusks 144
Tryonigens remondi (Tryon), anatomy of 61
Type locality of Clappiella saludensis 36
Unionidae, in Kentucky 60
Variation in Mudalia shell 26
Veronicellids in the Gulf Coast 72
Vertigo briarensis Leonard, n. sp 79
Vertigo occulta Leonard, n. sp 78
Washington state, new Hew.phillia 100
Yen, John Teng-Chien (1903-1972) no. 4, iii
vi NAUTILUS Vol. 85 (Index)
INDEX TO AUTHORS
Abbott, R. Tucker 49
Abbott, R. T. and C. B. Wurtz 1
Athearn, Herbert D no. 1, iii
Bingham, Frasier 0 136
Blankenship, Shaw 60
Boss, Kenneth J 9
Branson, Branley A 26, 100
Gather, J. N. and M. E. Crovo Ill
Clench, W. J 69, 145
Corgan, James X 51
Dundee, Dee S 67, 72
Emerson, William K 30
Gale, William F 71
Grandy, John W. IV 106
Grimm, F. Wayne 12
Harmon, Willard N 70
Horst, T. J. and R. R. Gosta 38
Hubricht, Leslie 73
Imlay, Marc J 146
Johnson, Donald Lee 32
Joy, James E 43
Kaicher, Sally Diana 126
Krakauer, Janet M 37
Krieger, K. A 120
Leonard, A. Byron 78
Lyons, William G 96
Malek, E. A. and M. D. Little 20
Metcalf, Artie L. and Richard Smartt 145
Miller, Walter B 61, 128
Murray, Harold D 36
Perchard, Peter L 84
Rawls, H. G. and John M. Baum 65
Rawls, H. G. and R. L. Yates 17
Ropes, John W 93
Rose, Kenneth D 141
Roth, Barry 110
Russell, Richard H 71, 145
Shoemaker, Alan H 72, 114
Smith, Allyn G 5
Townes, George F no. 1, iii
April, 1972 nautilus (Index) vii
IMPORTANT NOTICE
to subscribers and contributors to The Nautilus
Beginning with volume 86, number 1, to be published in July,
1972, the size, format, editorial policies and subscription prices
will change.
The new format will consist of two columns of text and/or illus-
trations on a semi-gloss page, 8x10^2 inches. Tables will be set in
type. For a similar publication, see the recent Bulletin for 1971,
vol. 37, of the American Malacological Union or the Proceedings
of the National Shellfisheries Association.
An editorial committee of malacologists has been formed in order
to facilitate the review of research papers in various special fields.
The annual subscription price is now $7.00 for individuals and
$12.00 for institutions. Subscriptions may begin either in January,
or in July when a new volume begins. Send check or money order
to "The Nautilus" to Mrs, Horace B. Baker, Business Manager,
11 Chelten Road, Havertown, Pa. 19083.
Contributors should send a short abstract witli their papers.
Authors should follow the style prescribed by the Style Manual for
Biological Journals which may be purchased from the American
Institute of Biological Sciences, 2000 P Street, NW, Washington,
D. C. 20036.
Charges: All authors or their institutions will be charged 50 cents
per line of tabular material and taxonomic keys. The publishers
reserve the right, seldom exercised, to charge $32 per page.
[The editors would like to thank Pony Printing and Mr. Sten
Dalstrom for their excellent services rendered over past years.]
Vol. 85 JULY, 1971 No. 1
THE
NAUTILUS
THE PILSBRY QUARTERLY
DEVOTED TO THE INTERESTS OF CONCHOLOGISTS
EDITORS AND PUBLISHERS
R. Tucker Abbott, du Pont Chair of Malacology
Delaware Museum of Natural History, Greenville, Del. 19807
Charles B. Wurtz, Biology Departmi'n^
La Salle College, Philadelphia, Pa. 19U
psnne Biological Labora
LI BRA P
CONTENTS JUL 2 ?^ 1971
Horace Burrington Baker (1889-1971) — an obitdary. WOODS HOLE MASS
By R. T. Abbott and C. B. Wurtz '. j i . . — I 1
New record for a rare Galapagos land snail.
By Allyn G. Smith 5
Familial affinities of Hemidonax (Bivalvia).
By Kenneth J. Boss 9
Two new Stenotrema, with notes on S. hirsutum and
S. barbatujn. By F. Wayne Grimm 12
Fluorescence in endodontid snails.
By H. C. Rawls and R. L. Yates 17
Aroapyrgus colombiensis n. sp. (Gastropoda: Hydrobiidae), snail
intermediate host of Paragonimus caliensis in Colombia.
By E. A. Malek and M. D. Little 20
Variation in the shell Mudalia potosiensis (Lea) (Pleuroceridae)
from a single locality. By Branley A. Branson 26
Niso (Neovolusia) imbricata (Sowerby, 1834) rediscovered
(Gastropoda: Eulimidae). By William K. Emerson 30
Pleistocene land snails on the Channel Islands, California:
a call for research. By Donald Lee Johnson 32
Notes 35
$5.00 per year ($5.75 to Foreign Countries) $1.50 a copy.
Mrs. Horace B. Baker, Business Manager
11 Chelten Road, Havertown, Pennsylvania 19083
Second Class Postage paid at Spring House, Pa.
NAUTILUS:
A quarterly journal devoted to the study of moUusks, edited and published
by R. Tucker Abbott and Charles B. Wurtz. Business and subscription
manager: Mrs. Horace B. Baker, 11 Chelten Road, Havertown, Pennsylvania,
U.S.A. 19083.
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THE NAUTILUS
Vol. 85 July, 1971 No. 1
HORACE BURRINGTON BAKER
1889 - 1971
By R. Tucker Abbott and Charles B. Wurtz
Horace Burrington Baker, former editor-in-chief of The Nautilus
and Professor Emeritus of Zoology, University of Pennsylvania, was
born January 25, 1889, in Sioux City, Iowa. He was one of
America's outstanding land mollusk anatomists, and had a keen
knowledge of systematics. For 40 years he gave unstintingly of
his time as business manager and editor of The Nautilus.
His father was Robert Folin Baker, born in England, and his
mother, Sophia Jane Burrington, was of English, Scotch and
French Huguenot origin. Horace received his high school educa-
tion in Flint, Michigan, and after working in a buggy factory for
a year, he attended the University of Michigan in 1906. In his
sophomore year he published his first paper on a "Key to the
genera of Gastropoda of Michigan." The summer of his graduating
year, 1910, was spent with Dr. A. G. Ruthven collecting mollusks
in Mexico. He was an instructor of biology at Michigan's Douglas
Lake biological station and, in 1913, an instructor at Colorado
College, Colorado Springs.
World War I interrupted his education. He was commissioned
a 2nd Lt., Field Artillery, in 1917, and spent eighteen months in
France and Germany with the 18th Field Artillery, Third Division.
Returning to the University of Michigan after the war, he partici-
pated in field work on mollusks in Venezuela, and received his
doctorate in 1920.
An early disappointment to Dr. Baker was the total loss of his
first doctoral dissertation after all the work was done, but before
it was submitted to his graduate committee. That first dissertation
was devoted to a study of midwestern grasshoppers, and it was lost
in a fire at the University of Michigan. This entailed a new
beginning for his doctoral research.
Most of Dr. Baker's professional life was spent as a professor
of zoology at the University of Pennsylvania where he was ap-
pointed an instructor in 1920, assistant professor in 1926, associate
1
2 NAUTILUS Vol. 85 ( 1 )
professor in 1928, and full professor in 1939. He became emeritus
in 1959. He was an instructor in the invertebrate summer courses
at Woods Hole, Massachusetts, in 1924 and 1925.
At the University of Pennsylvania Dr. Baker was best known
as an anatomist. He taught vertebrate anatomy to a great many
generations of aspiring medical students. He was also one of the
early teachers of ecology. The junior author, C. B. Wurtz, studied
ecology under him in 1947 and was one of his latest graduate
students, taking his doctorate under Drs. Baker and Pilsbry as
co-chairmen of his committee.
Dr. Baker was an excellent anatomical illustrator, as is evidenced
by his beautiful drawings. He would sit in a large, leather-covered,
reclining easy chair in his office and execute his drawing with a fine
draughtman's pen on a large wooden board, not by moving his pen,
but rather by moving the drawing paper. He made his drawings
usually 5 or 6 times as large as the intended final published figure.
Dr. Baker was a quiet, pleasant, somewhat introverted, gentle-
man. He was particularly kindly and considerate of women. He
enjoyed humorous stories and had an infectious quiet laugh. If
anyone asked him a question, particularly a technical or historical
one, he would promptly give a satisfactory answer, but after several
minutes or even hours of contemplation, he would return to his
interrogator with fuller, more detailed information. He was never
jealous of his prerogatives or his position. When he asked the
senior editor to handle The Nautilus after his first heart attack in
1968, he volunteered explicit permission to modify the journal in
any way necessary.
When a course in systematics was first given at the Academy of
Natural Sciences (about 1950), Dr. Baker conducted that part of
the program devoted to the statistical analysis of data. He enjoyed
statistics the way many enjoy crossword puzzles. He was also
persistent in pointing out the errors that can occur through the
incorrect application of statistical methods.
Dr. Baker was an avid collector of land and freshwater mollusks,
and always kept meticulous locality data. His many field trips
included those to Mexico (1910 and 1926), Venezuela (1919),
Dutch Leeward Islands (1922), southern Appalachians (1928),
Idaho (1929 and 1931), Jamaica (1933), Hawaii (1935). Puerto
Rico (1939) and several occasions in Florida.
July, 1971
NAUTILUS
HorjKi' Burrington Baker. ISS'I-I'ITI. Formal portrait was taken about 1941.
Lower left shows him as a graduate student about 1920. Lower right shows
him in 1968 at age 79 (photo by R. Robertson).
4 NAUTILUS Vol. 85(1)
In 1935, at the invitation of C. Montague Cooke, Jr., Dr. Baker
was invited for an eight-month Yale-Bishop Museum Fellowship in
Hawaii, where he worked on his monumental "Zonitid Snails from
the Pacific Islands" (Bulletins 158, 165 and 166 of the Bernice P.
Bishop Museum). He was an Honorary Research Associate in
Malacology at the Bishop Museum from 1937 until his death in
1971. Altogether, he published 218 papers and notes, amounting
to about 2,237 pages. All of his papers after 1941 were either
published in The Nautilus or the Proceedings of the Academy of
Natural Sciences of Philadelphia. At the latter, he was a Research
Associate and Fellow from 1925 until his death. After his retire-
ment from the University of Pennsylvania he gave his full volun-
teer time to renovating the land and freshwater collections and
publishing a list of the holotype specimens at the Academy.
Upon the death of Charles W. Johnson in 1932, Dr. Baker took
over the business managership of The Nautilus and served as an
associate editor for 25 years. Upon Henry A. Pilsbry's death in
1957, he became editor-in-chief. He was editor of the mollusk
section of the Biological Abstracts for many years, beginning in
1925. He was active in the American Malacological Union, becom-
ing a charter member in 1932, the ninth President in 1940, and
Honorary Life Member in 1958. He was a charter member of the
Philadelphia Shell Club (1955), gave a series of formal course
lectures to the members from 1963 to 1965, and was made the
Honorary Life President in 1968. He was also for years a member
of Sigma Xi, the American Society of Naturalists, the American
Society of Zoologists, the American Ecological Society and the
Pennsylvania Academy of Science.
Dr. Baker was married December 21, 1941, to Bernadine C.
Barker of Massachusetts. She had been secretary for the Boston
Malacological Club for many years and was a member of the
American Malacological Union. "Bunny," as she is called, has
served as business manager for The Nautilus since 1958. He was
a devoted husband and theirs was a harmonious marriage for 30
years. They have two daughters, Elizabeth Coffin and Abigail
Burrington (married Richard Woodhull Smith, March 15, 1965).
Dr. Baker suflfered a mild heart attack in 1968, and again three
years later when he died quietly early in the morning of March
11, 1971.
July, 1971 NAUTILUS 5
NEW RECORD FOR A RARE
GALAPAGOS LAND SNAIL
By Allyn G. Smith
California Academy of Sciences
San Francisco, Calif. 94118
Nacsiotus achatillinus (Forbes, 1850) has been collected only five
times according to jjublished records. The species is based on
material discovered on the (Jalapagos Island of Chatham (Isla San
Cristobal) in 1846 by Kellett and Wood of the British surveying
ship Pandora. However, there is an earlier collecting record.
According to Stearns (1893:405), the British collector Hugh
Cuming, whose mollusks from the Galajjagos were obtained in the
early 1830's, furnished Dr. Philip P. Carpenter with a list of his
land shells including A^. achatcllinus, which the latter published
(Carpenter, 1856:359). This record is confirmed by a single,
brilliantly-colored, nearly adult, live-taken specimen deposited in
the United States National Museum, Mollusk Division (no.
105150), with a label stating it was collected by Cuming and
placed subsequently in the Isaac Lea Collection. Unfortunately, the
island from which this shell came was not indicated.
The s])ecies was taken again in 1868, this time on Hood Ishnid
(Isla Espahola), by Dr. Simeon Habel, whose collection was re-
ported upon by August Wimmer in 1879. The next record is that
of Dr. Theodor Wolf, an Ecuadorian geologist, who collected on
the Galapagos Islands in 1875, although an account of the land
shells he obtained was not published until 1892 by Paul Reibisch.
While Wolf's shells were limited in numbers and generally were ol
poor quality, aj^parently he did find a single sj^ecimen of A'.
achatcllinus on "Chatham Island on mossy rocks at an elevation of
900 to 2000 feet . . ." 14ie fifth collecting record is by Dr. (ieorge
Baur, who made a collection in 1890 supported with important field
notes, which formed the basis for a classic discussion of insular land
shell faunas by William H. Dall in 1896. Evidently Baur found
only one specimen on Chatham Island at about 10(^0 feet elevation
on the undersides of the leaves of trees and bushes (USNM no.
107303), but this contained the soft parts from which Dall was
able to extract and describe the jaw and the radula.
A^. achatcllinus was not found by Charles Darwin during the
visit of the Beagle to the Galapagos in 1835; nor by Snodgrass and
Heller during the Stanford-Hopkins Galapagos Expedition of
NAUTILUS
Vol. 85(1)
Fig. 1 .A.-E. Naesiotus achat ellinus (Forbes). Freshwater Bay, Chatham
Island, Galapagos. L.G. Hertlein, Coll. 31 Dec. 1931 (Calif. Acad. Sci. Geol.
Dept. loc. no. 27208). Length of largest shell (IB). 16.6 mm.
1898-99; neither was it collected by W. H. Ochsner and the party of
the California Academy of Sciences Expedition to the Galapagos
in 1905-1906, which spent considerable time collecting land snails
on Chatham and Hood Islands, among others. No specimens of it
turned up dining the Galapagos International Scientific Project in
1 964. Thus it is with considerable satisfaction to be able to report
a si.xth collecting record accomplished by the well-known paleon-
tologist and conchologist to whom a previous issue of The Nautilus
was dedicated (vol. 84, no. 2) — Dr. Leo George Hertlein of the
California Academy of Sciences.
Dr. Hertlein was a participant in the Allan Hancock Expedition
to the Galapagos Islands in 1931-32 using the R/V Valero III.
Among numerous other collecting activities while the members of
the expedition were on Chatham Island, Dr. Hertlein tells me that
he filled a sizeable sack with leaf mold and forest litter at an
elevation of 300-400 meters from the sides of a small, permanent
fresh-water stream flowing into Freshwater Bay. Subsequent sorting
of this material produced five specimens of A^ achatcllmus as well
as a rich haul of other land-snail species, as follows:
July, 1971 NAUTILUS 7
Naesiotus canaliferus (Reibisch, 1892) 50-|-
curtus (Reibisch, 1892) 12+ juvs.
" chemnitzoidcs {Forhes, 1850) 1 subadult
Gastrocopta cf. G. munita (Reibisch, 1892) 100+
Pupisoma dioscoricola (C. B. Adams, 1845) 5
Habroconus? galapaganus (Dall, 1893) 25+
Retinella? chathamensis (Dall, 1893) 12
Succinea cf. S. bettii E. A. Smith, 1877 10-|- juvs.
Tornatellides chathamensis (Dall, 1892) 100+
HelicinanesioticaT)a.ll, 1892 100+
Unfortunately, as is often true in sortings of this kind, most of the
snails were dead although many of the shells were still in fairly
good condition.
The five shells of N. achatellinus consist of three adults and two
sub-adults (figures 1 A-E). Along with others collected at the same
time and place they have been accessioned into the California Acad-
emy of Sciences Geology Department Collection, locality no. 27208.
Measurements (in mm) of Dr. Hertlein's five shells are as follows:
Max. No. of
Remarks
Bleached; no color pattern shows.
Somewhat bleached; color band-
ing shows faintly.
15.1 8.0 7/2 Well preserved; shows color pat-
tern of 2 narrow brown bands on
a cream-colored background and
2 similar bands on the base.
12.8 7.3 7 Subadult; shows color pattern.
10.5 7.6 5% Bleached subadult; shorter and
stouter than the other shells; lacks
the well developed nodulous
sutures.
There is no doubt that A^. achatellinus is one of the rarer
Galapagos land snails. It is of special interest because of its super-
ficial resemblance to certain species of Hawaiian tree snails of the
genus Achatinella. Also, its straight-sided, conical shape, its non-
impressed, nodulose sutures, and its relatively bright color pattern
set it apart from any other known species of Naesiotus from the
Galapagos Islands or from the South American mainland. This led
Pfeiflfer to create a new subgeneric (or sectional) name Raphiellus
8 NAUTILUS Vol. 85(1)
for it in 1855, a taxonomic step concurred in by Dall (1896:429;
1928:152), although Pilsbry (1897-98:94) did not agree, placing
Raphiellus in the synonymy of the genus Naesiotus Albers, 1850.
The veiy fine, closely-spaced, transverse riblets on the nuclear whorls,
characteristic of Naesiotus, shows on one of Dr. Hertlein's speci-
mens although they have been lost through age and wear, or both,
on the other four shells.
The true systematic position of N. achatellinus with relation to
other Galapagos Island species of Naesiotus can be established
more firmly after collection of an adequate series of living snails for
which the exact habitat and other ecological factors have been
recorded accurately. It would be helpful to know, for example,
whether the species lives in an aboreal habitat as Dall thought it
might, or whether, in fact, it is more of a ground-loving species with
only occasional sorties up into tlie lower branches of bushes and
trees. The current interest in the Galapagos Islands and the easing
of travel problems to and from them may provide more and better
specimens than the few now available, together with more informa-
tion on the conditions under which they live on both Chatham and
Hood Islands.
I am indebted to Mr. Maurice Giles, scientific photographer
of the California Academy of Sciences, for the illustrations accom-
panying this report.
Selected References
Carpenter, Philip Pearsall, 1857. Report on the present state of our
knowledge with regard to the Mollusca of the West Coast of
North America. Rept. British Assoc. Adv. Sci. for 1856, pp.
358-362. Taylor & Francis, London.
Dall, William Healey, 1896. Insular landshell faunas as illustrated
by the data obtained by Dr. G. Baur in the Galapagos Islands.
Proc. Acad. Nat. Sci., for 1896, pp. 395-459, pis. 15-17.
1920. On the relations of the sectional groups of Bulimulus of
the subgenus Naesiotus Albers. Jour. Wash. Acad. Sci., 10 (5) :
117-122.
Dall, William Healey, and Washington Henry Ochsner, 1928.
Landshells of the Galapagos Islands. Proc. Calif. Acad. Sci., ser.
4, 17 (5): 141-185, pis. 8-9.
Forbes, Edward, 1850. On the species of Mollusca collected during
the surveying voyage of the Herald and Pandora, by Capt. Kellett,
R.N., C.B., and Lieut. Wood, R.N. Proc. Zool. Soc. for 1850, p.
56, pi. 19, figs. 5a-5b.
Pfeiffer, Louis, Versuch einer Anordnung der Heliceen nach
naturilchen Gruppen. Malacologische Blatter, vol. 2, p. 160.
July, 1971 NAUTILUS 9
FAMILIAL AFFINITIES OF HEMIDONAX (BIVALVIA)
By Kenneth J. Boss
Museum of Comparative Zoology
Harvard University, Cambridge, Mass. 02138
The placement of the genus Hemidonax Morch has long been
controversial but the anatomical and conchological features of
species in the genus show that it is allied to the family Cardiidae.
Recently Keen [in] Moore (1969: N629) considered Hemidonax
as a member of the tellinacean family Donacidae. In tlie past,
several authors have commented on the affinities of this genus
and it has alternately been referred to the Cardiidae (Vest, 1875),
the Tancrediidae (Fischer, 1887), the Crassatellidae (Hedley, 1906;
1909) as well as the Donacidae (Lamy, 1917). As subsequently
shown, several conchological and anatomical features preclude its
assignation to the tellinacean families Donacidae or Tancrediidae,
and its hinge structures are totally unlike those found in the
Crassatellidae or the related Astartidae and Carditidae. Iredale
and McMichael (1962) have gone as far as considering Hemidonax
the sole genus of a new family, the Hemidonacidae; however, they
did not characterize the family or establish its systematic position
in relation to other bivalves.
Early naturalists recognized the peculiar features of Hemidonax.
Thus, Deshayes (1835) remarked on Donax cardioides Lamarck:
II serait curieux de voir et d'etudier I'animal de cette espece, car
il est probable qu'elle n'appartient pas aux donaces; I'impression
palleale n'est point echancree posterieurement, et sa charniere se
rapproche plus de celle du cardium medium que de celle des
donaces.
Romer (1869) invited the description of a genus for Cardium
donaciforme when he said:
Eine der seltsamsten Formen, die wahrscheinlich wenn das Thier
bekannt sein wird, eine besondere Gattung bilden muss.
A response to this invitation was quickly received from Morch
(1870) who established Hemidonax and from Vest (1875) who
produced a junior subjective synonym, Donacicardium.
Little more was said about the genus until Lamy (1917), in a
paper on the Crassatellidae, discussed Hemidonax and concluded
that it was referable to the Donacidae since its hinge line was
similar to that of Donax denticulatus. Hedley (1923), who had
earlier (1906; 1909) advocated that die genus be placed in the
10 NAUTILUS Vol. 85(1)
Crassatellidae because it bore a great resemblance to Cyamiomactra,
responded with:
Dr. Lamy rejects my suggestion that Hemidonax might enter the
Crassatellitidae (sic) and restores it to the Donacidae. I feel
more confident that the entire pallial margin should exclude it
from the Donacidae than it should provide admission to the
Crassatellitidae. It is not clear why Dr. Lamy disallowed
Fischer's reference to Tancrediidae.
Thiele (1935) followed Lamy and made reference to Fischer
(1887), who had placed Hemidonax in the Tancrediidae, as had
Prashad (1932). Nearly all authors overlooked Pelseneer's (1911)
paper which described the gross anatomy of H. donaciforme. From
Pelseneer's figure (1911, pi. 13, fig. 4), the observations that
Hemidonax lacks a pallial sinus made sense: the genus has neither
retractable siphons nor the complex of retractor muscles that form
the scar of a pallial sinus. Further, it lacks a cruciform muscle and
the concomitant pallial scar of tellinaceans (Boss, 1966). The
Tancrediidae is a Mesozoic family of tellinacean affinities which
flourished from the Upper Triassic to the Upper Cretaceous (Cox
[in] Moore, 1969). The Donacidae is also tellinacean (i.e. having
a pallial sinus and cruciform muscle scars) and usually possesses
shells with comparatively smooth sculpture. Hemidonax has rela-
tively strong radial ribs and lacks both a pallial sinus and cruciform
muscle scars; thus, it must be excluded from both the Tancrediidae
and the Donacidae.
Pelsenecr (1911) indicated that the laterally compressed foot of
Hemidonax is unlike the elongate, geniculate foot of the cardiids
and placed the genus in close proximity to Crassatella. However,
the dentition of Hemidoriax (Lamy, 1917, p. 269) difiFers from that
found in either the Astartacea (Astartidae + Crassatellidae) or the
Carditacea and closely resembles that found in the Cardiidae
particularly Hemicardium. Thus, both conchological and anatomi-
cal features of Hemidonax show affinities with the family Cardiidae.
Characteristics usually typical of the Cardiidae and exhibited by
Hemidonax include: the lack of a pallial sinus, the concomitant
lack of an elaboration of incurrent and excurrent siphons, the
development of sti^ong radial sculpture, and the configuration of
the hinge and its dentition. Anatomically, Hemidonax compares
favorably with cardiids: 1) gills plicate, with both inner and outer
demibranchs; 2) siphons short, reduced, not markedly extensile or
retractile and not provided with elaborate siphonal retractor
July, 1971 NAUTILUS 11
muscles; 3) adductor muscles subequal and mantle gape extensive,
from the anterior adductor muscle to the incurrent aperture.
Thus Hemidonax can be considered a member of the Cardiidae
as first suggested by Vest (1875). However, the genus does not
have the elongate, geniculate foot, the pedal-byssal groove, the
pallial eyes, or the dorso-ventrally skewed axis of the ctenidium of
many cardiids and therefore forms a distinct taxon recognizable
by its bilaterally compressed foot, blunted cardinal dentition and
general donaciform shape. I think it merits nothing more than
subfamilial rank, for which we can use the emended name Hemi-
donacinae, Iredale and McMichael, 1962, and it may possibly be
related to the lineages which have radiated in the Pontian Basin
(Davitashvili and Merklin, 1966).
The species which constitute this genus and respective subfamily
are presently restricted in their distribution to the area of the
Philippine Islands, Indonesia, and Australia. Although Lamy
(1917) recognized a single species, several workers have augmented
that number so that there are now seven available nomina:
Cardium donaciforme Schroter 1786, Einleit. Conch., 2: pi. 7,
fig. 14; 3: 68 (type-locality, not known, subsequently given as the
Philippine Islands, see Romer, 1869, p. 100).
Cardium donaceum Spengler 1798, Skrivt. Naturh. Selsk, 6: 37
(type-locality, Drontheim in Norwegen, in error).
Donax cardioides Lamarck 1818, Anim. sans Vert., 5: 550-551
(type-locality, mers de la Nouvelle-Hollande, a I'ile Saint-Pierre-
Saint Francois [Nuyts Archipelago, Great Australia Bight]; see
Delessert, 1841, pi. 6, figs. 14a-c).
Cardium australiense Reeve 1844, Conch. Icon., vol. 2, Cardium.
pi. 5, fig. 24 (type-locality. Port Lincoln, South Australia); 1845.
Proc. Zool. Soc. London for 1844, p. 168.
Donax (Serrula) pictus Tryon 1871, Amer. J. Conch., 6: 23, pi.
1, fig. 1 (no locality given).
Hemidonax chapmani Gatliflf and Gabriel 1923, Vict. Nat., 40:
10, pi. 11 (type-locality, San Remo, Ocean beach, Victoria).
Hemidonax dactylus Hedley 1923, Proc. Linn. Soc. New South
Wales, 41: 303-304, pi. 31, fig. 13 (type-locality, Kiama, New
South Wales).
Literature Cited
Boss, K. J. 1966. The subfamily Tellininae in the Western Atlantic.
The genus Tellina (Part 1). Johnsonia, 4: 217-272, pis. 127-142.
Davitashvili, L. Sh. and R. L. Merklin. 1966. Spravochnik po
ekologii morshikh dvustvorok. (Monograph on the ecology of
marine bivalves) . Moscow, 348 pp.
Delessert, J. P. B. 1841. Recueil de coquilles decrites par Lamarck
dans son Histoire naturelle des animaux sans vertebres, et non
12 NAUTILUS Vol. 85(1)
encore figurees. Paris, 40 pis.
Deshayes, G. P. 1835. Histoire Naturelle des Animaux sans
Vertebres par J. B. P. A. de Lamarck, 2 Ed., vol. 6, Balliere, Paris,
600 pp.
Fischer, P. 1887. Manuel de Conchyliologie, Savy, Paris, 1369 pp.,
1158 figs., 23 pis.
Hedley, C. 1906. Studies on Australian Mollusca. Part IX. Proc.
Linn. Soc. N.S.W. (1905), pt. 4, pp. 520-546, pis. 31-33.
Hedley, C. 1909. Mollusca from the Hope Islands, North Queens-
land. Proc. Linn. Soc. N.S.W. (1909), 24 (3): 429-466.
Hedley, C. 1923. Studies on Australian Mollusca. Part XIV. Proc.
Linn. Soc. N.S.W., 48: 301-16, 21 figs.; pp. 303-304: H. dactylus,
pi. 31, fig. 13.
Iredale, T., and D. F. McMichael. 1962. A reference list of the
marine Mollusca of New South Wales, Aus. Mus. Mem., //,
109 pp.
Lamy, E. 1917. Revision des Crassatcllidae vivants du Museum
d'Histoire Naturelle de Paris. Jour, de Conchyl. 62: 197-270,
lOfigs., pi. 6.
Moore, R. C, ed. 1969. Treatise on Invertebrate Paleontology.
Part N. vol. 2 (of 3), Mollusca 6, Bivalvia, pp. N 491-951.
Morch, O. A. 1870. Ubersich der von Lorentz Spengler beschrie-
benen Conchylien. Malak, Blatt., 17: 99-124.
Pelseneer, P. 1911. Les lamellibranches de I'expedition du Siboga.
Partie Anatomique. Leiden, 125 pp., 25 pis.
Prashad, B. 1932. The Lamellibranchia of the Siboga Expedition.
Systematic Part. Pelecypoda II (exclusive of the Pectinidae).
Sffcoga-Expeditie, Monographic 53c, pp. 1-353, 9 pis.
Romer, E. 1869-1870. Die Familie der Dreiecks-oder Stumpf-
muscheln, Donacidae. [in] Martini-Chemnitz, Syst. Conch. -
Cab. (2), 10(3) : 122 pp., pis. 1-21, 21a.
Thielc, J. 1935. Handbuch der systematischen Weichtierkunde.
Gustav Fischer, Jena, 2, pp. 779-1022, 110 figs.
Vest, W. von. 1875. Ober die Genera Adacna, Monodacna und
Didacna Eichwald und deren Stellung im System. Jahrbiicher der
Deutschen Malakozoologischen Gescll., 2: 322, 324.
TWO NEW STEISOTREMA, WITH NOTES ON
S. HIRSUTUM AND S. BARBATUM
By F. Wayne Grimm
Visiting investigator, Mollusc Section,
National Museum of Natural Sciences,
National Museums of Canada, Ottawa 4
Stenotrenia simile, new species Figure A
Shell imperforate, thin, subtranslucent to opaque, subglobose
with a low conoid spire, a well rounded periphery and convex base.
Color variable, ranging from deep cinnamon-brown to olive buff
(of Ridgway). Embryonic whorl coarsely granulate with radially
July, 1971 NAUTILUS 13
elongate granules. Later whorls densely pilose, covered with
moderately long, stiff, curved hairs with rounded bases. Parietal
tooth slender, sinuous, not quite reaching the level of the basal lip,
gently curved into the interdenticular sinus, then curving gently
toward the outer lip tooth. Basal lip moderately wide, adnate to
the body whorl on the outer margin for its entire length, the inner
margin being conspicuously thickened around the lip notch. Lip
notch deep, wide, U-shaped, with a thick, nob-like tooth on its
peripheral edge. Interdenticular sinus deep and rounded. Outer
lip tooth small. Fulcrum quite short, barely projecting beyond the
basal lip callus, clearly visible through the shell, pointing radially
from near the axis.
Height 6.7 mm. Diameter 9.3 mm. Whorls 5V4. Holotype.
Distribution. — Maryland: (type locality) Garrett Co.: leaf
mould at base of steep slope near sharp bend of Bear Creek, 2.9
miles west of jet. U.S. -219 and Md.-42 (F. W. and G. F. Grimm,
G. B. Morris, coll.), holotype 61543, paratypes 61544 National
Museum of Canada; gorge of Bear Creek 2.8 miles east of Friends-
ville; leaf mould and logs near summit of Backbone Mountain,
Roth Rock Fire Tower. West Virginia : Monongalia Co. : Cooper's
Rock State Park. Nicholas Co.: wet sandstone talus 6.0 miles nortli
of Richwood; cool wet woods along North Fork of Cherry River,
2-3 miles north of Richwood. Pocahontas Co.: damp maple hillside
5.3 miles west of Mill Point.
Stenotrema simile differs from S. hirsutum (Say) by being a bit
larger, having coarser granulations on the embryonic whorl, a
denser pile of longer hairs, a shorter fulcrum, a more distinct but
thinner callus lining the lower lip, a deeper, more heavily callused
lip notch which has a nob-like tooth on its peripheral margin, a
deeper interdenticular sinus, and a more sinuous, thinner parietal
tooth. Stenotrema simile is a rare, solitary species which inhabits
cool, wet woods and ravines on the Appalachian Plateau. In the
same region, S. hirsutum is a gregarious species which inhabits dry
upland oak woods, cut over woods, and clearings.
Stenotrema simile differs from S. barbatum (Clapp) by having
coarse granulations on the embryonic whorl (the embryonic whorl
of S. barbatum is smooth to finely striate), smaller hairs on the
surface of the shell, a much shorter, stouter fulcrum, a rounder
oudine with a proportionally higher spire and rounder base, a
thicker callus on the lower lip, a deeper, more heavily callused lip
14
NAUTILUS
Vol. 85(1)
Figure 1. Shells of A. Stenotrema simile Grimm, holotype; B. Stenotrema
barbatum (Clapp), Washington Co., Md.; C. Stenotrema hirsutum (Say),
Howard Co., Md.; D. Stenotrema burringtoni Grimm, holotype.
notch with a nob-like tooth on its peripheral margin, a deeper
interdenticular sinus, a higher parietal tooth and a narrower
aperture. The inner half of the basal lip is adnate to the body
whorl in S. barbatum; the entire basal lip is adnate in S. simile and
S. hirsutum.
Stenotrema burringtoni, new species Figure D
Shell small, imperforate, subtranslucent, depressed globose with
a low conoid spire, a well rounded periphery and convex base.
Color pale cinnamon-brown (of Ridgway). Embryonic whorl
coarsely granulate with rounded granules. Later whorls covered
with moderately long, stiff, straight hairs with rounded bases.
Parietal tooth comparatively stout, quite sinuous, reaching the
level of the basal lip, strongly curved into the interdenticular sinus,
its outer edge curved toward the outer lip tooth. Basal lip wide,
sinuous, thickened on the inner edge. Lip notch deep, U-shaped,
thickened at both edges, with a thick tooth at its peripheral edge.
July, 1971 NAUTILUS 15
Interdenticular sinus deep, forming a trough, rounded. Outer lip
tooth quite distinct. Fulcrum short, very stout, projecting consider-
ably beyond the basal lip callus, clearly visible through the shell,
pointing tangentially from the axis and slanted toward the aperture.
Height 5.5 mm. Diameter 8.4 mm. Whorls 5. Holotype.
Type-locality. — Virginia: Highland Co.: limestone rubble 3.1
miles south-southwest of Mustoe on U.S. -220, holotype 61545,
para types 61546 National Museum of Canada and Del. Mus. Nat.
Hist. No. 41544.
Stenotrema burringtoni differs from S. hirsutum (Say) by
averaging slightly smaller, having coarser, more rounded granula-
tions on the embryonic whorl, a denser pile of longer hairs, a
shorter, stouter fulcrum which is slanted toward the aperture (that
of S. hirsutum is slanted away from the aperture), a thicker, more
sinuous callus on the basal lip, a deeper, more heavily callused lip
notch, a deeper, more rounded interdenticular sinus, and a higher,
more sinuous parietal tooth which is turned into the interdenticular
sinus, then swings upward toward the outer lip tooth. The parietal
tooth of S. hirsutum points directly at the outer lip tooth.
Stenotrema burringtoni is named in honor of Horace Burrington
Baker.
Stenotrema hii*sutuni (Say) Figure C
Because this species is likely to be confused with both of the
preceding species, and frequently has been confused with Steno-
trema barbatum (Clapp), a short discussion of it is presented here.
Little can be added to the accurate description given by Pilsbry
(1940, p. 662) except to state that despite its wide range, S. hirsu-
tum is quite uniform in its appearance. After examining many lots
of this species in the collections of the United States National
Museum, the Academy of Natural Sciences of Philadelphia, the
University of Michigan Museum of Zoology, and the National
Musuem of Canada, I noted no significant variation in sculpture or
in the appearance of the aperture. The diameter varies from about
6.4 mm. to 8.8 mm., and the height from 4.5 mm. to nearly 6.0 mm.
Extremes in size are rather rare. The fulcrum of S. hirsutum is
moderately large, about ^4 to ^3 the width of the body whorl, and
is slanted posteriad.
Stenotrema hirsutum ranges from Connecticut to northern
Georgia in the Piedmont and Appalachians, west on unglaciated
land to southern Indiana, south through central Kentucky and
16 NAUTILUS Vol. 85(1)
Tennessee to northern Alabama and northeastern Mississippi.
Throughout its range it occupies upland oak woods, mixed oak-pine
woods, cut over woods, and dry clearings. Often it is most abun-
dant in ecotonal situations such as roadsides and the edges of
partially cleared upland woods.
Stenotrenia barbatum (Clapp) Figure B.
Polygyra (Stenotrema) barbata Clapp, 1904, Nautilus 18 (8):
85.
Polygyra hirsuta yarmouthensis F. C. Baker, 1927, Nautilus 40
(4): 115.
Stenotrema hirsutum form yarmouthense (F. C. Baker), Pilsbry,
1940, Land Moll. N. Am. / (2) : 665.
Stenotrema hirsutum (Say) in part, Pilsbry, op. cit.: 662-665.
Stenotrema hirsutum barbatum (Clapp), Pilsbry, op. cit.: 665-
666.
Stenotrema hirsutum (Say), Oughton, 1948, Univ. of Toronto
Biol. Ser. 57: 10.
Stenotrema harbatwn (Clapp), Hubricht, 1950, Nautilus 64 (1):
7. Listed, no discussion.
Stenotrema barbatum (Clapp), Hubricht, 1962, Sterkiana 8: 1.
Discussion.
Stenotrema hirsutum (Say) in part, LaRocque, 1970, Bull. Geol.
Svy. Ohio 62 (4) : 566.
Stenotrema barbatum may be separated from S. hirsutum by its
smooth to obsoletely ribbed embryonic whorl, by its larger, more
widely spaced, stiffer hairs, by the weaker armature of its aperture,
and by its longer, more slender fulcrum. The basal lip of S.
barbatum is reflected, and does not become adnate to the body
whorl until the lip notch is reached. The basal lip of S. hirsutum
and the two species herein described is adnate to the body whorl
for its entire length. The lip notch of S. barbatum is quite weak
and shallow, nearly disappearing in some examples, and the aper-
ture is wide because of the weakness of its teeth and thin basal lip.
Stenotrema barbatwn is quite variable in size, ranging from 6.5
mm. to slightly over 11 mm. in diameter, and 4.1 mm. to 7 mm.
in height. Usually it averages between 8.5 mm. and 10.5 mm. in
diameter. Proportionally, its spire is lower than that of the other
species discussed here. The largest examples are from floodplains
near the Gulf Coast, the smallest are from loess and silt in the
July, 1971 NAUTILUS 17
northern Mississippi and Ohio valleys and from low ground in
Michigan and southern Ontario.
This species ranges from Massachusetts to southern Minnesota
(including the southern tip of Ontario), south in the west through
Iowa to Kansas, in the east to Mississippi and Alabama, north on
the Coastal Plain and Piedmont through the Carolinas to southern
New England. Reports of Stenotrema hirsutum from northern
Ohio, central and western New York, Michigan, Wisconsin, Iowa
and Kansas are based upon S. barbatum.
Throughout most of its range Stenotrema barbatum is hygro-
phile, preferring shaded floodplains and marshes. It has been
called the "lowland form" of S. hirsutum. It is most abundant
where the calcium content of the soil is high, but it is found on
acid soils as well. In scattered upland areas, usually near flood-
plains, it occupies ravines, clearings, and fields. I have found it
with S. hirsutum several times in ravines at the bases of hills near
floodplains on the Piedmont of Maryland, and have seen no
intergradation.
Literature Cited
Baker, F. C, 1927. Descriptions of new forms of Pleistocene land
mollusks from Illinois with remarks on other species. Nautilus
40 (4): 114-120.
Clapp, George H., 1904. New forms of Polygyra from Alabama.
Nautilus 18 (8) : 85-86.
Hubricht, Leslie, 1950. The Polygyridae of Pittsylvania County,
Virginia. Nautilus 64 (1) : 6-9.
, 1962. Pleistocene land snails of southern Mississippi and
adjacent Louisiana. SterkianaS: 1-11.
LaRocque, Aurele, 1970. Pleistocene Mollusca of Ohio. Bull. Geol.
Survey of Ohio 62 (4) : 566.
Oughton, John, 1948. A zoogeographical study of the land snails
of Ontario. University of Toronto Biological Series *57: 10.
Pilsbry, H. A., 1940. Land Mollusca of North America (north of
Mexico). Acad. Nat. Sci. Phila. Monogr. 3, / (2): 662-666.
FLUORESCENCE IN ENDODONTID SNAILS
By Hugh C. Rawls and Roger L. Yates
Department of Zoology, Eastern Illinois University
Charleston, Illinois 61920
A review of the literature related to fluorescence reveals little
concerning the occurrence of this phenomenon in animals. Furreg
and Querner (1929, 1930) described fluorescence of the shells of
gastropods of the families Trochidae and Turbinidae. Latham
(1953) reported the occurrence of fluorescence in the fur of the
18 NAUTILUS Vol. 85(1)
least weasel, Mustata frenata noveboracensis. More recently,
Zahl, (1963, 1968) presented popular accounts of this phenomenon
in various kinds of animals, including certain coelenterates, arthro-
pods and marine molluscs. Heretofore, it appears, fluorescence has
not been observed in any of the land molluscs. We wish to report
the discovery of this phenomenon in representatives of three of the
genera of the family Endodontidae, and to suggest its probable
occurrence in the remaining genera of the family.
Our first observation of this phenomenon occurred as we were
attempting solution of a problem involving chromatographic study
of mucus from snails of several families, including endodontids.
Resorting to the use of ultraviolet light in an effort to visualize
better a chromatogram of the mucus from Anguispira alternata, we
found that a particularly brilliant, bluish fluorescence appeared
under this illumination. With this observation as a clue, we
turned to chromatograms of mucus from specimens of Anguispira
kochi and discovered a similar fluorescence, but one which appeared
to be distinctly different from that which we had previously ob-
served. Our interest heightened, we prepared strip chromatograms
of mucus from Discus patulus and were rewarded by the appear-
ance of yet another seemingly distinctive fluorescence. The
chromatograms were prepared on F' x 8" strips of No. 4 Whatman
paper. We used the solvent reported by Kirk, Main and Beyer
(1954): 1-butanol, acetic acid, and water (100:22:50 v/v). The
mucus was collected from snails by the use of standard blood capil-
lary tubes which were inserted into the apertures of the shells; this
technique provided a means of stimulating the snails to produce
large quantities of mucus, as well as a means by which suitable
volumes of mucus could be collected and delivered to the paper
strips. Because of the viscosity, we found it necessary to expel the
collected mucus samples onto the paper by gently blowing through
the capillary tube. Three such applications were made on each
strip and allowed to dry before hanging the strip from the solvent
tray of the chromatographic jar. Solvent was allowed to descend
for three hours, after which the strip was removed and air-dried.
This technique gave a good distribution of the fluorescent material
present in the mucus, and it provided a convenient source for that
material for other aspects of a continuing study. We used a Min-
eralite-SL unit for our initial observations, but we found that the
fluorescence which appeared was primarily in response to long-
July, 1971 NAUTILUS 19
wave ultraviolet illumination; weaker fluorescence was observed
under short-wave light.
A somewhat natural progression of events led us to the observa-
tion that we could locate specimens of Anguispira and Discus in
the dark, by simply playing the U-V light over the tray on which
numerous snails had been spread; the mucus on the foot and body
of each endodontid glowed brightly, as did the mucus trails left as
these snails moved over the tray. Other specimens, all polygyrids,
failed to exhibit any sign of the fluorescence, and we began to view
the presence of fluorescence in the endodontids as something of a
diagnostic characteristic.
Further investigation revealed that alcohol -preserved specimens
of Anguispira and Discus could be identified readily by the fluor-
escence of the fluid in which the specimens had been kept. Later,
we observed that snails of these genera could be recognized by
simply holding the vials, containing the snails, under the U-V
light and observing the characteristic fluorescence. We did find,
however, that any endodontid contamination of alcohol used to
preserve specimens of other families produced a degree of fluor-
escence which could be misleading. One vial of Stenotrema, for
example, was observed to fluoresce even though all other specimens
which we had of this genus did not; the data on this vial corres-
ponded with data on several vials of Anguispira, and we concluded
that specimens of both genera had been collected and preserved in
common before being sorted.
Our experience with the preserved specimens of Anguispira and
Discus led us to the further discovery that Helicodiscus parallelus,
which we had available only in preserved form, exhibited still
another apparently distinctive fluorescence. On the basis of our
observations so far, we believe that Punctum and Radiodiscus,
neither of which is represented in our preserved or live collections,
will display a fluorescence similar to that exhibited by the genera
which we have been able to investigate.
The fluorescence which we have observed in specimens of the
three genera noted is extremely long-lived, being as bright and as
distinctive in specimens preserved for twenty years and more as it
is in living snails. Moreover, it is not simply a local phenomenon
peculiar to snails of the central Illinois region. Mucus taken from
specimens of Anguispira and Discus collected in several widely
separated localities in Illinois responded to U-V illumination; and
20 NAUTILUS Vol. 85(1)
alcohol -preserved specimens of all three genera mentioned, from
numerous locations throughout their ranges, likewise exhibited the
characteristic fluorescence.
Some animals fluoresce under U-V light because of the presence
of specific minerals; others, because of the presence of certain
organic compounds; still others fluoresce because of the presence of
specific microorganisms. Certain evidence now available to us
suggests that this phenomenon occurs in endodontids because of
the presence of two or more as yet unidentified organic compounds
in the mucus of these snails. Preliminary spectrophotometric work
indicates a significant difference in the chemistry of the compounds;
fluorescence peaks appear between 275 and 300 nm, and again
between 350 and 400 nm, in mucus samples from all three genera
investigated. We have also found, hov.^ever, that the mucus of all
living endodontids in our collection invariably contains bacteria of
the genus Pseudomonas. We are testing the role, if any, that these
bacteria play in the production of fluorescence in the snails. At the
same time, several other avenues are being explored and we expect
to report on these investigations in a subsequent paper.
Literature Cited
Furreg, E. and F. R. Querner. 1929. Uber Eigenartige Fluorezzenz-
erscheinungen an Gastropoden Schalen (Fam. Trochidae; Tur-
binidae). Anz. Akad. Wiss. Wien, 66: 96-98. 1929.
, 1930. Uber Fluorezzenzerscheinungen an Gastropoden
Schalen. Z. Wiss. Zool. Leipzig, 136: 355-375. 1930.
Kirk, R. L., A. R. Main and F. G. Beyer. 1954. Paper Chroma-
tography in Taxonomic Work. Biochem. Journal, 57 (3): 440.
1954.
Latham, Roger M. 1953. Simple Method for Identification of the
Least Weasel. J. Mammal, 34:385. 1953.
Zahl, Paul A. 1963. Fluorescent Gems from Davy Jones's Locker.
Nat. Geog. Mag., 164 (2): 260-271. 1963.
, 1968. Scorpions: Living Fossils of the Sands. Nat. Geog.
Mag., /33 (3): 436-442. 1968.
AROAPYRGJJS COLOMBIENSISN. SP. (GASTROPODA:
HYDROBIIDAE), SNAIL INTERMEDIATE HOST OF
PARAGOJSmUS CALIENSIS FN COLOMBIA^
By Emile a. Malek and M. D. LmLE
Department of Parasitology, School of Public Health and
Tropical Medicine, Tulane University, New Orleans, La. 70112
During investigations of paragonimiasis in Colombia (Little and
Epler, unpublished report) a hydrobiid snail was found to be
naturally infected with a new species of Paragonimus, P. caliensis
July, 1971
NAUTILUS
21
bm buccal mass ey eye ovi oviduct
bu bursa copulatrix gi gill sh shell
dg digestive gland oes oesophagus te tentacle
eg egg capsule ov ovary ve verge
em embryo
Fig. 1. Aroapyrgus colomhiensis, showing shell and operculum. Fig. 2.
Female specimen with the reproductive organs. Fig. 3. Male specimen showing
the verge. The gill shows through the shell.
Little, 1968. The snails were collected from swift, shallow streams
2 to 6 feet wide, and with fairly steep banks. The snails were most
readily collected in quiet pools on dead decaying leaves. Laboratory
colonies were successfully established, with maple leaves being used
as food. The availability of field and laboratory-reared snails per-
mitted a study of the systematics of this new hydrobiid belonging
to the genus Aroapyrgus, to which A. colomhiensis is assigned.
Holotype: Delaware Museum No. 41542. Paratypes: Delaware
Museum No. 41543; Field Mus. Nat. Hist. 168647; Acad. Nat. Sci.
Phila. 321818. Type locality: Tributaries of Rio Pichinde, near
Pichinde, Municipio de Cali, Departamento de Valle de Cauca,
Colombia.
Methods
Specimens were relaxed with menthol before they were fixed in
Bouin's or FAA. Morphological studies were carried out with some
1 This study was supported by a U. S. Public Health Service Re.search Career
Award K6-AI- 18,424 to the senior author, and by the Tulane University Inter-
national Center for Medical Research and Training Grant TW-00143, and
grants AI-04919 and AI-00002 from the National Institutes of Health, U.S.
Public Health Service. Acknowledgments are made to the Seamens Research
Laboratory, USPHS Hospital for many facilities.
22 NAUTILUS Vol.85 (1)
shells of these snails, while others were used for in toto mounts,
after being stained with Semichon's carmine. Because of the small
size of the snail it was found necessary to study its morphological
details from serial paraffin sections, stained with Ehlrich's hema-
toxylin and eosin. The radula and operculum were studied in
glycerin preparations.
Description of the snail
The Shell (Fig. 1) — Field collected specimens are covered with
a dark brownish deposit. In laboratory-reared specimens, and a few
field ones, however, the shell is smooth, light yellowish in color
and translucent. Usually there is no size difference between the
males and females. Very few females, however, are slightly broader
than the males of the same age. The shell in the fully developed
specimens has 5 to 6 evenly rounded whorls. The body whorl is
rapidly increasing; some of the growth lines are raised in the form
of very fine ridgelets. Fine microscopic ridgelets are also present on
the body whorl, and on the penultimate whorl. The protoconch
consists of one whorl only. The sutures are deeply impressed; the
umbilicus is open, and relatively large. The aperture is ovoid and
almost vertical; the peristome is simple and sharp, and is slightly
flaring in the palatal region. The columellar margin is simple. The
operculum is thin, corneus and paucispiral. Average shell measure-
ments from 15 specimens in millimeters are as follows: height 3.1,
width 1.6, aperture height 1, and aperture width 0.8.
The Animal (Figs 2 and 3) — ^The foot is spatulate, truncate
anteriorly, and bluntly pointed posteriorly. The tentacles are
slender on both sides of the moderately pigmented snout. The gill
of Aroapyrgus colomhiensis consists of about 30 lamellae and starts
near the mantle collar, where it is wide, and narrows down pos-
teriorly under the roof of the mantle, through which it shows very
clearly.
The radula is similar to that of other hydrobiids (Baker 1930,
Berry 1943, Thompson 1968). The central tooth has from 12 to 15
cusps on its reflected margin, and 3 basal cusps. The lateral has
9 to 13 cusps, the 6th from the inside (the mesocone) is twice as
large as any of the others, and the tooth is attached by an elongate
base which extends obliquely laterad and posteriad from its squar-
ish body. The handle is elongate and narrow. The inner marginal
tooth is spoon-shaped, its concave blade carries about 25 to 35
small cusps on the reflection. The outer marginal is thinner and
July, 1971 NAUTILUS 23
more slender than the inner, and has much smaller cusps which are
numerous, and are not easily seen except in fresh preparations.
Description of the reproductive organs is based on reconstructions
from serial paraffin sections. In the male the testis is located at the
level of the third whorl, and is embedded among the tubules of the
digestive gland. Seminal vesicles are located at the beginning of
the vas deferens. The latter continues as a narrow duct until it
opens into the verge. A large prostate gland extends transversely
about the level of the penultimate whorl. The verge arises inside a
circular fold to the right of the mid-dorsal line. As is characteristic
of species belonging to this genus the verge (Fig. 3) is large, simple
(devoid of any papillae and is unbranched); its terminal half is
smaller in diameter than the base. The seminal duct runs along
the outer margin of the verge.
In the female (Fig. 2) the ovary is located at the same level as
is the testis in the male. The oviduct arises from the ovary as a
narrow channel before it enlarges to form a "brood pouch," which
is the equivalent of the pallial oviduct in other hydrobiids. A
bursa copulatrix and a seminal receptacle open at about the junc-
tion of the two parts of the oviduct. Numerous embryos, up to 30,
are found in the brood pouch. The active embryos, each with a
•shell of about one whorl, are located within the egg capsules from
which they are apparently released at birth. The birth pore is
found in the proximal end of the brood pouch at the mantle collar.
Discussion
Baker (1930) proposed Aroa as a subgenus of Potamopyrgus,
with P. (Aroa) ernesti vivens as type. (The species ernesti was
described as Hydrobia ernesti by von Martens 1873, as a subfossil
from Lago de Valencia). The reasons Baker gave are that the
verge is simpler than in Potamopyrgus and Littoridina, and still
is not bifid, as in Amnicola or Hydrobia ( Paludestrina) . Later,
Baker (1931) renamed it Aroapyrgus, and Morrison (1946) used
Aroapyrgus as genus with the genotype Aroapyrgus ernesti vivens
H. B. Baker. According to Morrison (1946) the genus is known to
occur from Panama to Cayenne in French Guiana. Some species
described under Amnicola in the region from Mexico to Costa Rica
might also belong to this genus but will remain uncertain until
animal characteristics are better known.
Morrison (1946) named and described the species Aroapyrgus
alleei, A. chagresensis and A. joseana from the Panama region.
24 NAUTILUS Vol. 85(1)
A. alleei has faintly shouldered whorls, whereas A. chagrcsensis has
evenly rounded whorls. A. joseana has axially shorter whorls, than
the other two species, and can also be distinguished from them by
its smaller aperture and narrower conical outline. A. colombiensis
can be distinguished from the above 3 species by its somewhat
turreted spire, by having more whorls (up to 6 rounded ones),
and by a narrower aperture. Except for the genus characteristics
of the copulatory organ Morrison (1946) did not mention any
other anatomical features of his new species. Baker (1930)
described Aroapyrgus ernesti vivens ( —Potamopyrgus (Aroa)
ernesti vivens) as a new subspecies from Venezuela. He noted that
it was the commonest form in the small streams of central and
western Venezuela. A. colombiensis differs from A. ernesti vivens
in the fact that generally there is no sexual dimorphism in the
Colombian species contrary to the case with the Venezuelan form.
Pilsbry (1935) described Potamopyrgus laciranus, as a new species
from La Cira formation near Zopffs, La Cira District, Colombia.
The shells were found to be abundant in the La Cira hematitic
sandstones. The description of the shell of A. colombiensis as given
in the present paper agrees in the main with that described by
Pilsbry.
A note on the nomenclatorial history of this hydrobiid group
seems in order. Pilsbry (1891) regarded the two genera Pyr-
gophorus (Ancey) and Potamopyrgus (Stimpson) as synonymous.
Morrison (1939), however, was of the opinion that Potamopyrgus
is strictly a New Zealand genus, and that Pyrgophorus is found
only in the Americas. He also thought that Lyrodes (Doering) is
a synonym of Pyrgophorus. Species belonging to Aroapyrgus can
be differentiated from species belonging to the latter genera by the
characteristics of the verge. The verge is very simple in Aroapyrgus
spp., whereas it is bifid, and is provided with very distinct papillae
in Pyrgophorus.
Several hydrobiids have been described from South America
under various genera. It is obvious from a review of the literature
that further studies are still needed to clarify their taxonomic
status. Weyrauch (1963) for example described, on the basis of
the shell only, a new species of Potamopyrgus from Peru, P.
mirandoi, and some new species of Littoridina from Argentina and
Peru. Parodiz (1960) reported from Argentina, a new species of
Lyrodes, L. doellojuradoi whose shells resemble those of Aroapyrgus
July, 1971 NAUTILUS 25
spp. However, their anatomy remains to be studied.
The morphology of various British and European prosobranchs,
in particular Potaraopyrgus jenkinsi, were dealt with in detail by
Fretter and Graham (1962). The comparative morphology of the
various organ systems of these mollusks were elucidated. The
morphology of several hydrobiids was treated by various authors,
viz. Pomatiopsis spp. by van der Schalie and Dundee (1956),
Dundee (1957), van der Schalie and Getz (1962); Oncomelania
nosophora by Roth and Wagner (1957); Cochliopa texana and
Lyrodes cheatumi by Dundee and Dundee (1969); various hydro-
biids from Florida by Thompson (1968). The above reports as well
as the present report show the usual hydrobiid morphology and,
indicate that generic and specific differences shown are based on
details of the structure of certain parts of the genitalia, particularly
the verge, and the oviduct. The pallial portion of the oviduct is
considerably expanded in the viviparous forms to accommodate the
fully developed embryos. Other features of diagnostic value are:
the radula, shape of the gill and number of gill lamellae, size of
the osphradium, branching of the ovary, and shape of the bursa
copulatrix and prostrate gland.
Literature Cited
Baker, H. B. 1930. The mollusca collected by the University of
Michigan-Williamson expedition in Venezuela. Occ. Papers
Mus. Zool. Univ. Mich. No. 210: 1-94.
, 1931. A new name in Potamopyrgus. Nautilus, 44: 143.
Berry, E. G. 1943. The Amnicolidae of Michigan: Distribution,
ecology and taxonomy. Misc. Publ. Mus. Zool. Univ. Mich., No.
57: 1-68.
Dundee, D. S. 1957. Aspects of the biology of Pomatiopsis lapidaria
(Say). Misc. Publ. Mus. Zool. Univ. Mich., No. 100; 1-37.
and Dundee, H. A. 1969. Notes concerning two Texas
molluscs, Cochliopa texana Pilsbry, and Lyrodes cheatumi, Pils-
bry (Mollusca: Hydrobiidae). Trans. Amer. Micros. Soc, 88:
205-210.
Fretter, V. and Graham, A. 1962. British prosobranch molluscs.
Their functional anatomy and ecology. The Ray Society, Lon-
don, 755 pp.
Little, M. D. 1968. Paragonimus caliensis sp. n. and paragonimiasis
in Colombia. J. Parasit., 54:738-746.
Morrison, J. P. E. 1939. Notes on the genera Potamopyrgus and
Lyrodes. Nautilus, 52: 87-88.
, 1946. The nonmarine mollusks of San Jose Island, with
notes on those of Pedro Gonzalez Island, Pearl Islands, Panama.
Smithsonian Misc. Collect. 106, No. 6: 1-49.
26 NAUTILUS Vol. 85(1)
Parodiz, J. 1960. Neotype for Lyrodes guaranitica Doering, and
description of a new species. Nautilus, 74: 24-27.
Pilsbry, H. A. 1891. Land and fresh- water mollusks collected in
Yucatan and Mexico. Proc. Acad. Nat. Sci. Phila., 43: 310-334.
and Olsson, A. A. 1935. Tertiary fresh-water mollusks of
the Magdalena embayment, Colombia. Proc. Acad. Nat. Sci.
Phila., 87:7-39.
Roth, A. A. and Wagner, E. D. 1957. The anatomy of the male
and female reproductive systems of Oncomelania nosophora.
Trans. Amer. Micros. Soc, 76: 52-69.
Thompson, F. G. 1968. The aquatic snails of the family Hydro-
biidae of Peninsular Florida. Univ. Florida Press. Gainsville,
268 pp.
van der Schalie, H. and Getz, L. L. 1962. Morphology and develop-
ment of the sex organs in the snail Pomatiopsis cincinnatiensis
(Lea). Trans. Amer. Micros. Soc, 8/:332-340.
and Dundee, D. S. 1956. The morphology of Pomatiopsis
cincinnatiensis (Lea), an amphibious prosobranch snail. Occ.
Papers, Mus. Zool. Univ. Mich., No. 579: 1-17.
Weyrauch, W. K. 1963. Cuatro nuevas especies de Hydrobiidae de
Argentina y Peru (Gastropoda, Prosobranchia). Acta Zoo-
logica Lilloana, 19: 243-259.
VARIATION IN THE SHELL OF
MUD ALIA POTOSIEISSIS (LEA)
(PLEUROCERIDAE) FROM A SINGLE LOCALITY
By Branley A. Branson
Eastern Kentucky University
Richmond, Kentucky 40475
A number of workers have attempted to correlate aquatic gas-
tropod shell variation with environmental factors. Wiebe (1926),
for example, reported a direct correlation between shell size and
shape and exposure to wave action, reporting, however, that differ-
ences in obesity index arose via two main avenues: (1) because of
apical erosion and (2) because of actual differences in shell size
and shape. Several mechanisms underlying apical erosion have
been suggested in the literature, from a combination of physical
and chemical factors (Jones and Branson, 1964; Goodrich, 1936)
to mechanical abrasion by water-borne silt (Bailey, Pearl, and
Winsor, 1932, 1933 a). The latter investigators (1933 b) were
not able to correlate shell characters with any features of the
environment. However, Adams (1915) demonstrated headwaters-
to-mouth variation in the shells of lo, and Cheatum and Mouzon
(1934) found that shells of Goniobasis comalensis Pilsbry averaged
longer and wider when secured from ponded areas as contrasted
July, 1971
200
NAUTILUS
27
9.0 10.0
OlAMETEfl
12.0
Figure 1. Regression of length on diameter in Mudalia potosiensis. F,
female; m, male; t, total; n=120 (61 female; 59 male). Open stars, female;
.solid stars, male. Measurements in mm.
with ones from stream situations, and that the mean ratio diameter-
to-length was greater in river shells than in ones from ponds,
although these differences were not statistically significant.
Since none of the above authors investigated possible correlations
between sex and shell variability, that question seems to be a
natural avenue for investigation: are there measurable differences
28 NAUTILUS Vol. 85(1)
between the shells of the two sexes? Since shell variation has been
noted in specimens secured from different habitats and localities,
all the specimens utilized in this study were secured from a single
locality: the downstream side of a rock — approximately 2 feet by
1.5 feet in size — Shoal Creek, Redding's Mill, 2 miles south of
Joplin, Jasper County, Missouri, 24 March 1962. From the 3,150
specimens collected at this site (deposited in the U.S. National
Museum: USNM 262804), a random sample of 100 specimens
was secured by means of blind-selection, one specimen at a time.
The snails were killed by immersion in boiling water, the bodies
removed from the shell, and the sex of each specimen determined
following the technique of Jones and Branson (1964) and Stimpson
(1864). Each shell was marked for later identification.
Although apical erosion was present in most of the shells, I do
not believe this presents a serious problem as regards the analyses
discussed below. I have assumed a normal distribution of the
erosion in both sexes, and hence the quantitative results are
probably valid and true. The measurements of greatest length
and width (distance between the outermost point on the peristome
and the opposite side of the body whorl) were made by means of
a vernier caliper and are believed to be accurate within ±0.05 mm.
All computations were effected by means of Fortran programs.
Discussion — Utilizing Hubbs and Hubbs' (1953) method of
graphical presentation, in which two standard errors are plotted on
each side of the sample mean (no overlap indicates samples from
different populations), demonstrates no significant differences be-
tween males and females as regards length and width (Figures 2
and 3). However, plotting these same measurements on a scatter-
graph demonstrates some points of interest. The regression line for
male specimens scarcely diverges from that for the whole popula-
tion (Figure 1). The slope of the line for females does diverge
somewhat from the other two. The reason for this divergence is
obvious by inspection of the graph. As regards diameter, at levels
below 8.5 mm there is a disparity of 12:7 in favor of females, and
above 9.5 mm, a 19:15 ratio, again in favor of the females. In
length, there is a similar degree of disparity in favor of females:
9:5 above 17.0 mm, and 3:1 below 13.0 mm.
These kind of data, of course, tend to smooth out some types of
graphs (such as the Hubbs' type), and hence to obscure certain
aspects of populations, in this example, a moderately developed
July, 1971 NAUTILUS 29
60 7.0 8.0 9.0 100 110 12.0 13.0
Figure 2. Shell diameter in Mudalia potosiensis. Horizontal line, range;
vertical line, mean; open box, one standard deviation on each side of mean;
closed box, two standard errors on each side of mean. Other symbols, abbrevia-
tions, numbers and measurements as in Figure 1. Confidence intervals set at
99% level.
10.0 10.5 11.0 II.S 12.0 I2.S 13.0 I3.S 14.0 14.5 IS.D IS.5 IB.O 16.5 170 I7.S IB.O IB.5 19.0 IB.S2Q.0
IS.065I • 0.409S
IJ.JIK * 0.ltSl
I4.t320 - 0.1401
Figure 3. Shell length in Mudalia potosiensis. Data as in Figure 1 and 2.
trend toward sexual dimorphism in shell characteristics. The
significance of these findings, although interesting, may be of
minimal value to the researcher interested in separating specimens
according to sex, although the odds would be 2:1 in his favor when
lengths above 17.0 mm were used as the distinguishing criterion.
Literature Cited
Adams, C. C. 1915. The variations and ecological distribution of
the snails of the genus lo. Mem. Nat. Acad. Sci. 12: 1-184.
Bailey, J. L., R. Pearl and C. P. Winsor. 1932. Variation in
Goniobasis virginica and Anculosa carinata under natural con-
ditions. Biol. Gen. 8: 607-630.
Bailey, J. L., R. Pearl and C. P. Winsor. 1933 a. Variation in
Goniobasis virginica and Anculosa carinata under natural con-
ditions. II. The relations between size of the shells and environ-
mental factors. Biol. Gen. 9: 301-336.
Bailey, J. L., R. Pearl and C. P. Winsor. 1933 b. Variation in
Goniobasis virginica and Anculosa carinata under natural con-
ditions. III. Correlations shape of shells, and conclusions. Biol.
Gen. 9:48-69.
Cheatum, E. P. and E. D. Mouzon. 1934. Biometrical study of
Goniobasis comalensis Pilsbry from two diverse habitats. Field
30 NAUTILUS Vol. 85(1)
and Lab. 3:18-23.
Goodrich, C. 1936. Goniobasis of the Coosa River, Alabama. Misc.
Pub. Mus. Zool. Univ. Mich. 31: 1-60.
Hubbs, C. L. and Clark Hubbs. 1953. An improved graphical
analysis and comparison of series of samples. Syst. Zool. 2:
49-56; 92.
Jones, W. C. and B. A. Branson. 1964. The radula, genital system,
and external morphology in Mudalia potosiensis (Lea) 1841
(Gastropoda: Prosobranchiata: Pleuroceridae) with life history
notes. Trans. Amer. Micros. Soc. 83: 41-62.
Stimpson, W. 1864. On the structural characters of the so-called
melanians of North America. Amer. J. Sci. Arts 38: 41-53.
Wiebe, A. H. 1926. Variations in the freshv^ater snail, Goniobasis
livescens. Ohio J. Sci. 26: 49-68.
mSO (JSEOVOLUSIA) IMBRICATA
(SOWERBY, 1834) REDISCOVERED
( GASTROPODA : EULIMID AE )
By William K. Emerson
American Museum of Natural History
New York, New York 10024
At the time I reviewed the eastern Pacific representatives of the
genus Niso (Emerson, 1965), the only Recent specimen known for
Niso imbricata (Sowerby, 1834a) was the holotype, which was
stated to have been dredged by Hugh Cuming off Santa Elena,
Ecuador. Fossil specimens of this taxon, however, had been re-
ported from a Pliocene deposit in Ecuador (Pilsbry and Olsson,
1941). As a result of collections made by Mesdames Carmen
Angermeyer and Jacqueline DeRoy in the waters of the Galapagos
Islands, additional specimens of this rare Niso have been recovered.
This species is now known to occur also at Clarion Island, Revil-
lagigedo Islands, Mexico, on the basis of specimens taken more than
35 years ago by the Allan Hancock Pacific Expeditions (Dr. James
H. McLean, personal communication). These new records, to-
gether with notes on the holotype of Sowerby's taxon, form the
foundation for this report.
The Angermeyers dredged one, well-preserved, specimen in 10 to
15 fathoms off Isla Rabida (here illustrated, left figure), and they
found a fragmental specimen in the digestive tract of a fish that
had been caught off Isla Santa Fe (AMNH No. 132795), in 1965.
The DeRoys also obtained specimens by dredging in 5 to 6.5
fathoms in Academy Bay, Isla Santa Cruz, in November, 1965 and
January, 1966. Two of these specimens, which measure 19.5 and
22 mm. in length, are now in the respective collections of Mr.
July, 1971 NAUTILUS 31
Niso (Neovolusia) iinhricata (Sowerby, 1834)
Left fig. Off Isla Rabida [Jervis Island], Galapagos Islands, in 10-15
fathoms, Angermeyer coll.; 18.9 mm. in length.
Right fig. Holotype of Eulima imbricata Sowerby, 1834, off Santa Elena.
Ecuador, in 6-8 fathoms, British Museum (Natural History) No. 1965226;
18.2 mm. in length.
Anthony D'Attilio of San Diego, California, and Dr. Donald R.
Shasky of Redlands, California.
Three additional specimens came to light when the collection of
gastropods of the Allan Hancock Foundation of the University of
Southern California was transferred on loan to the Los Angeles
County Museum of Natural History, in 1967. According to Dr.
McLean, these are hermit crab specimens. They were collected
intertidally at Sulphur Bay, Clarion Island, of the Revillagigedo
Islands, on January 5, 1934 (Fraser, 1943, p. 280; AHF Sta.
141-34).
All of these specimens are without doubt referable to Sowerby's
long-lost species, the holotype of which was examined by me during
a recent visit to the British Museum (Natural History). The type
specimen (here illustrated, right figure) is somewhat faded, but
the characteristic color pattern of axially arranged, reddish brown
hneations on a whitish surface is retained. The whorls are not
as angulated as depicted in the illustrations of Sowerby (1834b, fig.
4; Adams, 1854, pi. 70, fig. 10; Reeve, 1866, fig. 3). The axial lines
also are not weakly varicose, as the drawings suggest, but they
have this appearance to the naked eye. The recently obtained
Galapagan specimens, however, have slightly more inflated whorls
and the suture is not as strongly developed as that of the holotype
(see figures).
Neovolusia Emerson (1965, p. 8) was proposed to replace the
homonym Volusia A. Adams, 1861, and it was afforded subgeneric
32 NAUTILUS Vol. 85(1)
status to include Niso imbricata, which, on the basis of the illus-
trations of the type specimen, I wrongly concluded was weakly
varicose. Dr. McLean has kindly pointed out to me that Niso
imbricata, together with the Panamic Niso excolpa Bartsch, 19 17,
form a distinctive group in which the periphery projects beyond
the whorl below. Therefore, I have retained Neovolusia as a
subgeneric unit to include the species of Niso with this morpho-
logical character.
Acknowledgements: In addition to Mesdames Angermeyer and
DeRoy, and Dr. McLean, I am also indebted to Mr. S. Peter Dance,
formerly of the British Museum (Natural History), for providing
a photograph of the holotype of Niso imbricata.
Literature Cited
Adams, Arthur, 1854. Monographs of the genera Eulima, Niso,
Leiostraca, Obeliscus, Pyramidella, and Monoptygma. In Sower-
by, G.B. II, Thesaurus conchyliorum. London, 2: 793-825, pis.
169-172.
Adams, Arthur, 1861. On some new species of Mollusca from the
north of China and Japan. Ann. Mag. Nat. Hist., ser. 3, 8:
299-309.
Bartsch, Paul, 1917. A monograph of west American melancllid
mollusks. Proc. U.S. Nat. Mus., 53: 295-356, pis. 34-49.
Emerson, W. K., 1965. The eastern Pacific species of Niso.
(Mollusca: Gastropoda). Amer. Mus. Novitates, no. 2218, pp.
1-12, 11 figs.
Fraser, C. M., 1943. General account of the scientific work of the
Velero III in the eastern Pacific, 1931-41, pt. 3. Allan Hancock
Pacific Expeds., / (3): 255-431, 115 charts.
Pilsbry, H. A., and A. A. Olsson, 1941. A Pliocene fauna from
western Ecuador. Proc. Acad. Nat. Sci. Philadelphia, 93: 1-79,
pis. 1-19, 2 text figs.
Reeve, L. A., 1866. Monograph of the genus Niso. Conchologia
iconica. London, vol. 15, Niso pi. and text.
Sowerby, G. B., I, 1834a. Characters of new species of Mollusca
and conchifera, collected by Hugh Cuming. Proc. Zool. Soc.
London, for 1834, pp. 6-8.
Sowerby, G. B., I, 1834b. A catalogue of the Recent species of the
genus Eulima. In Sowerby, G. B. II, The Conchological illustra-
tions. London, 2 pp, 2 pis.
PLEISTOCENE LAND SNAILS ON THE CHANNEL
ISLANDS, CALIFORNIA: A CALL FOR RESEARCH
By Donald Lee Johnson
Department of CJeography
University of Illinois, Urbana, 111. 61801
In the course of soil and geomorphological studies carried out
July, 1971
NAUTILUS
33
2S MILES
Figure 1. Location map of the California Channel Islands.
over a number of years on the Channel Islands of California the
writer has been increasingly impressed with the malacological re-
search potentials offered by the islands. The location of the islands
is shown in Figure 1.
While each of the four outermost islands^ of San Clemente, San
Nicolas, Santa Rosa, and San Miguel offer more than casual
interest for the malacologist concerned with possibly tracing evolu-
tionary changes of characters of a species, the latter island, San
Miguel, is by far most interesting and potentially most scientifically
rewarding. On this island. Pleistocene mollusks number literally in
the millions and appear to the writer, who is untrained in mala-
cology, to be primarily of the single species Helminthoghjpta
ayresiana (Newcomb), although a specialist's eye might discrimi-
nate others. The mollusks occur at many places over the island in
some of the oldest as well as very recent Quaternary deposits
(Figure 2), and are still extant. -
The modern shells are noticeably smaller than their Pleistocene
counterparts, an observation earlier made by Cockerell (1937) but
not emphasized by him. Although Cockerell was of the opinion
' Santa Rosa Island is privately owned and, as experience has shown, is difilcuh
(.{ access. San Clements and San Nicolas are both owned and administered by
the U.S. Navy; permission to do scientific research is usually granted by writing
directly to the basa commanders, respectively. San Miguel Island is also owned
by the U. S. Navy, but is administered by the National Park Service; parmission
to work on the island must come from the Channel Islands National Monu-
ment, Oxnard, California.
34
NAUTILUS
Vol.85 (1>
Figure 2. Typical scene on San Miguel Island. California. Subfossil
Helminthog,lypta aijresiana shells weathering out of a paleosol and over-lying
sediments.
"I do not see that it is possible to distinguish the fossil shells from
the recent . . ." his view was based on two brief visits to the island
in July, 1937 (Cockerell 1938) and in May 1938 (Cockerell 1939)
before any detailed stratigraphic studies had been carried out, and
before the profusion and age of the shells established. Whether
linear relationship between time and shell size exists remains to
be determined.
- Cockerell (1938) described a fossil subspecies H. aijresiana lesteri from San
Miguel Island, but sines shells occur ubiquitously from oldest through youngest
nonmarine sediments we have no way of knowing where his specimens fit
temporally, or morphologically, in the chronologic column. Additionally,
probably it is premature to assign a subspecitic rank based on non-quantitative
parameters, especially when the .shells occur in quantities sufficient for statistical
analysis.
July, 1971 NAUTILUS 35
The late Quaternary subaerial (nonmarine) sedimentary record
on San Miguel Island is probably more complete than anywhere
else on the west coast of North America. For example, in one
particular exposure which consists of a complex of paleosols (fossil
soils) and eolian sand deposits, the entire span of radiocarbon time
is represented, and the age of the lower zones of the sedimentary
unit is well beyond radiocarbon range. Land snails occur through-
out the deposit. Similar exposures abound on the island, and are
contributing to a Quaternary chronology.
The large number of snail shells present in the Quaternary
deposits should easily lend themselves to quantitative analyses of
shell morphological characters, such as size, banding, and so on.
Additionally, the proven association of snail shells with vertebrate
remains, fossil plants, and archaeological sites on the island attests
to their cross-disciplinary importance, and as Taylor points out
"Pleistocene history will be most advanced by combined researches
on stratigraphy, soils, fauna, and flora; through such investigations
research on fossil mollusks will not only yield more but receive
more from neighboring sciences" (Taylor 1965).
In summary, the Channel Islands of southern California,
especially San Miguel, offer a fund of untapped malacological
research possibilities. The nonmarine Quaternary deposits on San
Miguel Island contain a virtually continuous record of subfossil
land snails from late Pleistocene times to the present, presumably
of a single species which occurs in great numbers. To the writer's
knowledge no detailed malacological studies of these subfossil
snails have been made. This vacuum of research activity should be
filled.
References Cited
Cockerell, T. D. A., 1937. Naut. 51 (2) : 71-72.
, 1938. Naut. 52 ( 1 ) : 24-25.
,1939. Bios /O (11): 99-106.
Taylor, D. W., 1965. Quaternary of the U.S., (Princeton U.
Press), Princeton, pp. 597-61 1.
NOTES
New^ Records of Corbicula manilensis (Philippi) in Texas
— The spread of the "Asiatic clam," Corbicula manilensis, in the
United States is well documented in the literature by various
authors. Since the reports by Dundee and Harman (1963, Nautilus
77:30) of C. manilensis from Louisana and by Metcalf (1966,
Nautilus 80:16-20) witli tlie first records of the species from west-
36 NAUTILUS Vol. 85(1)
ern Texas near El Paso, C. manilcnsis has been expected in eastern
and southeastern Texas. Mrs. Constance Boone, Houston, Texas,
forwarded to me specimens of C. manilensis obtained in February,
1969, from the Mexican side of Falcon Lake which boarders
United States and Mexico. This suggested that the species occurred
on the U.S. side of the lake.
On April 18, 1969, Kery Hummel, a Trinity student, collected
C. manilensis from Monte Alto Resei^voir (locally called Delta
Lake) in Hidalgo County, Texas. This reservoir has a sand and
mud bottom, and the water is used for irrigation of vegetable and
fruit farming in the extreme southeastern part of Texas. Monte
Alto Reservoir is approximately 150 miles east of Falcon Dam and
over 750 miles east of the first Texas recorded by Metcalf. Kery
Hummel recovered additional specimens of C. manilensis on May
3, 1969 from an irrigation canal beside U. S. Highway 281 near
Relampago, Hidalgo County. Specimens from Monte Alto Reservoir
are judged to be 3-4 years old, and specimens from near Relampago
are 1-2 years old which indicates a more recent introduction to the
Relampago area. Since Metcalf suggested his specimens were
approximately four years old in 1964, establishment of C. manilensis
probably occurred first in western Texas and secondly in south-
eastern Texas.
In August, 1969, Mrs. Boone forwarded to me a single specimen
of C. manilensis collected from Lake Mathis, 25 miles northwest of
Corpus Christi, Texas. Persons attending the 1968 American
Malacological Union in Corpus Christi will recall the visit to this
lake which at that time yielded no C. manilensis. Lake Mathis is
on the Nueces River approximately 170 miles north of the popula-
tions in the Lower Rio Grande drainage in Hidalgo County.
C. manilensis is now established in southeast Texas and appears
to be moving in a northern direction. The next few years will
probably see additional Texas records as this species spreads into
eastern Texas areas where irrigation for cotton and rice farming
is common. — Harold D. Murray, Trinity University, San Antonio,
Texas 78212.
Protection of the Type Locality of Clappiella saludensis
(Morrison) — Clappiella saludensis (Morrison) is listed as a rare
and endangered eastern land snail by Dundee (Malacologia, Vol.
10, No. 1, May, 1970). The type locality in Greenville County,
S. C, as described by Morrison (Proc. Biol. Soc. Wash., Vol. 50,
July, 1971 NAUTILUS iii
p. 59, 1937), is now included vvithin the watershed area of a
reservoir maintained by the Greenville (S. C.) Waterworks. The
area is maintained in a natural state and all public access is
denied. This type locality is probably effectively and permanently
protected against most adverse factors. — George F. Townes, P. O.
Box 10128 F. S., Greenville, S. C. 29603.
Sphaerium simile (Say) in Tennessee. — In the summer of
1962, I collected S. simile Say 1816 (S. sulcatum Lamarck 1818 is
a synonym) from Grassy Cove, Cumberland County, a large sink-
hole measuring nearly four miles across and with a bottom eleva-
tion of about 1560 feet above sea level. The surrounding mountains
of the Cumberland Plateau range up to 2900 feet in elevation. This
is the first record from the Tennessee River drainage. The species
does not extend through the underground streams of Grassy Cove
drainage into the Tennessee Valley proper. Davenport Spring,
from which these specimens were taken, is located about 35 yards
west of Highway 68 and one mile north of Grassy Cove Church.
Other mollusks found in this spring and its sluggish outlet were:
Goniobasis teres Lea, G. laqueata castanea Lea, Helisoma trivolvis
Say, Physa microstoina Haldeman, Pisidium compressum Prime and
P. casertanum Poli. — Herbert D. Athearn, R-5, Box 376, Cleveland,
Tennessee 37311.
Dates of The Nautilus. — Vol. 84, no. 1 mailed July 16, 1970.
No. 2, Oct. 5, 1970. No. 3, Jan. 25, 1971. No. 4, April 26, 1971.
WORLD WIDE SPECIMEN SHELLS for sale. New 1971
Price List on request. "Illustrated Catalog of Popular Marginella
Species" now available also, 117 species shown with full data
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WILLIAM H. WEEKS SHELL COLLECTION: New price lists
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Sea Shells of
Tropical West America
MARINE MOLLUSKS FROM
BAJA CALIFORNIA TO PERU
SECOND EDITION
A. Myra Keen
With the Assistance of James H. McLean. This
revision and expansion of a classic work treats
all of the molluscan species recorded from the
Panamic province — the tropical and subtropical
shores of the Eastern Pacific and the offshore
islands as far as the Galapagos. Some 3,325
species are treated in this book, and for the
majority the authors provide nomenclatural data,
synonyms, detailed descriptions, typical dimen-
sions, range data, notes on relationships, and one
or more illustrations. Also included are synopses
for all taxonomic groups above species, anatomical
drawings for most major groups, identifying keys
to the subgenus level, introduction and system-
atics, a list of excluded species, glossary compre-
hensive bibliography, four coastal maps, a list of
place names, source data on all illustrations, and an
index to names and synonyms. Illustrated with
some 4,000 halftones and line drawings; 22 pages
in full color illustrate 85 species (including 29
nudibranchs and other opisthobranchs ) .
1040 pages. $25.00
Stanford University Press
Vol. 85 OCTOBER, 1971 No. 2
THE
NAUTILUS
THE PILSBRY QUARTERLY
DEVOTED TO THE INTERESTS OF CONCHOLOGISTS ,
EDITORS AND PUBLISHERS
R. Tucker Abbott, du Pont Chair of Malacology
Delaware Museum of Natural History, Greenville, Del. 19007 —.
Chakles B. Wurtz, Biology Department \}^^^ LauOratOi'
La Salle College, Philadelphia, Pa. 19141 ^' ^^
r- '■ 1971
CONTENTS lyinno ».n. r>
WOOUS HOLE, MASS
The feeding habits of aplysiid npi-t]inhr;inrhi in Finn'na
By Janet M. Krakauer 37
Distribution patterns of five selected gastropod species from
McCargo Lake. By T. J. Horst and R. R. Costa 38
The influence of light conditions upon the egg-laying of the
planorbid snail, Biomphalaria glabrata. By James E. Joy . . 43
Conus patae, a new Caribbean gastropod. By R. Tucker Abbott. 49
Review of Parodostomia, Telloda, Goniodostomia and
Eulimastoma (Gastropoda: Pyramidellacea).
By James X. Corgan 51
Notes on Alasmidonta fabula (Lea) in Kentucky (Unionidae).
By Shaw Blankenship 60
The reproductive anatomy of Tryonigens remondi (Try on, 1863) :
Helminthoglyptidae. By Walter B. Miller 61
Fluorescence in Mesodon clausus (Say). By H. C. Rawls and
John M. Baum 65
United States research trends in malacology. By Dee S. Dundee . 67
Notes 69 Publications received iii
$5.00 per year ($5.75 to Foreign Countries) $1.50 a copy.
Mrs. Horace B. Baker, Business Manager
11 Chelten Road, Havertown, Pennsylvania 19083
Second Class Postage paid at Spring House, Pa.
NAUTILUS:
A quarterly journal devoted to the study of moUusks, edited and published
by R. Tucker Abbott and Charles B. Wurtz. Business and subscription
manager: Mrs. Horace B. Baker, 11 Chelten Road, Havertown, Pennsylvania,
U.S.A. 19083.
AUTHORS PLEASE NOTE
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THE NAUTILUS
Vol. 85 October, 1971 No. 2
THE FEEDING HABITS OF
APLYSIID OPISTHOBRANCHS IN FLORIDA
By Janet M. Krakauek
Department of Biology, Hollins College, Virginia 24020
While collecting data on the ecology of Aphjsia willcoxi for a
Master's thesis at the University of Florida, some incidental obser-
vations were made on the other two aplysiids, Bursatella leachi plei
and Phyllaplysia engeli, which are known to occur in the Cedar Key
area. The question arose as to whether or not the food preferences
of these animals, all purported to be herbivores (Hyman, 1967),
overlapped.
A survey was made of the macroscopic algae found in the Cedar
Keys and the crop contents of 47 A. willcoxi were analyzed. The
crop contents and fecal material from two B. leachi were also exam-
ined. Observations were made with a dissecting microscope on two
living P. engeli while they were feeding.
Analysis of the crop contents indicate that A. willcoxi eats chiefly
macroscopic red algae. Less than 5% of the crops contained frag-
ments of green and brown algae. Fragments of the smaller sea
grasses, Divlanthera and Syringodium, were also found in some of
the sea hares' crops. The only animal matter present was attached
or slow moving organisms such as bryozoans and some unidentified
gastropods. In aquaria, A. willcoxi fed on both red and green algae,
but refused brown algae. Captive sea hares were never observed to
feed on animal matter.
The crop contents of B. leachi include a variety of materials of
both plant and animal origin. There was an assortment of diatoms,
sand grains, ostracod exoskeletons, some very small fragments of the
red alga Spyridia, and some other unrecognizable detritus. The
fecal material contained chiefly sand grains and some living diatoms.
B. leachi, in aquaria, was observed to feed on bits of shrimp, and
appeared to be more adept at feeding from the bottom of the
aquarium than was A. willcoxi.
Although no crop contents were examined in the case of P. engeli,
observations were made on some living animals while they fed.
37
38 NAUTILUS Vol. 85 (2)
P. engeli is much smaller than the other two species, and is well
camouflaged to blend in with the turtle grass, Thalassia, to which it
clings while feeding. P. engeli feeds on the microscopic epiphytes of
the turtle grass.
The three aplysiids differ appreciably in their food preferences
and do not compete with each other. A. willcoxi is restricted to the
red algae in the Cedar Keys, probably because the green algae,
although palatable to the sea hares, are chiefly intertidal and are
not generally available to subtidal organisms. The animal matter
in the sea hares' crops appears to have been ingested by accident.
B. leachi shows the food preferences of a bottom feeding organism.
It apparently ingests a variety of food. Whether or not diatoms are
a part of B. leachi's diet is open to question since living diatoms
were found in the fecal material.
P. engeli appears to restrict its feeding activities to the epiphytes
of the turtle grass. In this it resembles P. taylori which Beeman
(1969) reports feeds on the diatoms on eel grass.
Literature Cited
Beeman, R. D. 1969. An autoradiographic demonstration of stomach
tooth renewal in Phyllaplysia taylori Dall, 1900 (Gastropoda:
Opisthobranchia). Biol. Bufl. 136: 141-146.
Hyman, L. H. 1967. The invertebrates: Mollusca I. McGraw-Hill,
New York.
DISTRIBUTION PATTERNS OF FIVE SELECTED
GASTROPOD SPECIES FROM McCARGO L\KE
By Thomas J. Horst and R. R. Costa
Division of Biology
Kansas State University
Manhattan, Kansas 66502
Introduction — Selected gastropod species were studied with refer-
ence to their distribution in McCargo Lake, Orleans County, New
York. The lake is located on the Fancher Campus of SUNY College
at Brockport. It has a surface area of 31,948 m.^ and a maximum
depth of 5.75 m.
The results of investigations conducted during July and August
1970 are contained in this report. The species studied were Amni-
cola limnosa, Valvata tricarinata, Physa sayii, Gyraulus parvus and
Promenetus exacuous. Other gastropods present in the lake but not
McCargo Lake
Fig. 1. Location of stations along the liorth-^-outh and East-West transects.
included in this study are Helisoma trivolvis, Viviparus georgianus
and Ferrissia sp.
The habitats and ranges of water temperature associated with
these gastropods are discussed by Baker (1928) and Harman
(1968). Boycott (1936) considers the most iniportant features of
molluscan habitats to be clean water, "absence of disturbance" and
the presence of lime.
Methods and Materials — Two transects were sampled during the
summer of 1970. Three Ekmann (6 in. by 6 in.) bottom samples
and a Kemmerer water sample were taken from the hydrosol layer
at each of the twenty stations (Fig. 1).
Water samples were analyzed for temperature, dissolved oxygen,
alkalinity, carbon dioxide, hydrogen sulfide, turbidity and color.
Water analyses were performed according to Standard Methods
(APHA, 1965). Hydrogen sulfide, turbidity and color were moni-
tored using the HACH colorimeter.
Analysis of bottom samples was performed according to the
techniques of Duncan (1955). Gastropod counts obtained from
replicate samples were analyzed and for all stations were found to
be acceptable at a 95% confidence limit.
Results — ^Twenty stations which varied in depth from 0.75 m. to
5.75 m. were sampled along two transects.
Temperature profiles indicated that the epilimnion extended to
a depth of 2.0 m. during the sampling period. Photometeric read-
40 NAUTILUS Vol. 85 (2)
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October, 1971
NAUTILUS
41
Table 2, Two way analysis of variance, North-South and East-West transects
NORTH-SOOTH TRANSECTS, STATIONS 14-20
-ignificant at O.05 level
ings revealed only 5% light transmittance at the 2.0 m. zone. Secchi
disk determinations ranged from 1.25 m. to 1.75 m.
Dissolved oxygen concentration at stations v^^here live gastropods
were collected ranged from 2.6 ppm to 8.1 ppm. A 0.70 correlation
coefficient between gastropod density and dissolved oxygen was
found. A negative correlation ( — 0.64) was found to exist between
hydrogen sulfide and dissolved oxygen.
Phenolphthalein alkalinity was present at stations on the north
and south shores while carbon dioxide was detected at all other sta-
tions. Total alkalinity values ranged from 228.0 ppm to 286.0 ppm.
A summary of chemical and physical data is presented in Table 1.
Live gastropods were collected at eight of the twenty stations
(Figs. 2-3). These stations ranged in depth from 0.75 m. to 2.50 m.
Discussion — Water samples taken in the profundal zone, below
2.5 m. were found to have chemical and physical properties different
from those of the littoral zone. Water color and turbidity were most
pronounced in the profundal zone. Water samples taken below
3.0 m. were found to have low dissolved oxygen and high hydrogen
sulfide concentrations. Alkalinity and carbon dioxide concentrations
reflected lower pH values for the deeper stations.
42
NAUTILUS
Vol. 85 (2)
Aanieol* lljgios*
ValY«t« trlcarinat*
nyraulua parvus
Proffenetus ex
hig. 2. Kean number of gastropods per iktnann sample collected at stations
along the iast-x'est transect.
Limited light penetration and its apparent effect upon the com-
position of the algal community were factors associated with the
absence of gastropods. Samples collected below 1.5 m. contained
Oscillatoria sp. and were associated with lower gastropod density.
October, 1971 nautilus 43
Sediments of the littoral zone were lighter in color and contained
more detrital matter than those of the deeper layers. The only
sediments containing gravel were collected on the south shore. This
is the only shore of the lake with high relief.
Results of analysis of variance indicated that the south shore
gastropod community was significantly different than that of the
north shore (Table 2). There was no significant difference, how-
ever, at the east and west shores. A. limnosa was the most abundant
species at all shores except the south. V. tricarinata was most abun-
dant there.
Summary and Conclusions — A study of the distribution of the
gastropods of McCargo Lake was conducted during the summer of
1970.
Water of the littoral and profundal regions was found to have
different chemical and physical properties. Low oxygen tension,
high hydrogen sulfide concentration and low pH probably accounted
for the absence of gastropods in the profundal zone. Limited light
penetration in the lake and its effect upon the algal community
appeared to be related to gastropod distribution.
Additional work is needed to relate sediment composition and
algal communities to gastropod distribution.
Literature Cited
A.P.H.A. 1965. Standard Methods for the Examination of Water and Waste-
water. 12th ed. New York. 769 p.
Baker, F. C. 1928. The Freshwater Mollusca of Wisconsin, Part 1: Gastropoda.
Wisconsin Nat. Hist. Sur. Bull. 70(1): 1-507.
Boycott, A. E. 1936. The Habitats of Freshwater Molluscs in Britain. J. of
Animal Ecology. 5: 116-186.
Duncan, C. J. 1959. The Life Cycle and Ecology of the Freshwater Snail Physa
fontinalis. }. of Animal Ecology. 28: 97-117.
Harman, W. N. 1968. The Distribution and Ecology of the Aquatic Gastropoda
of Central New York with Illustrated Keys to the Genera and Species. Un-
published Ph.D. Thesis. Cornell University, Ithaca, New York. 398 p.
THE INFLUENCE OF LIGHT CONDITIONS
UPON THE EGG-LAYING OF THE
PLANORBID SNAIL, BIOMPHALARIA GLABRATA
By James E. Joy
College of Veterinary Medicine
Department of Veterinary Parasitology
Texas A&M University
College Station, Texas 77843
The influence of light on the behavior of snails has been given
little attention. Pesigan et al (1958) showed that the reaction of
44 NAUTILUS \'ol. 85 (2)
Oncomelania quadras: to light was one of avoidance as tlie snails
invariably crawled awa}- n-oni a high intensity' light source, the
speed of their movement ina"easmg as the intensity of the light
increased. That Oncoryieluv.ia avoids direct sunlight or strong direct
light rays has also been documented by Abbott (1948), Kawamoto
(1952). Komiya et al (1959). and Moose and Williams (1961-62).
In an attempt to correlate the distribution of B. glahrata with
en\T.ronmental factors in Puerto Rico. Harn' and Aldricli (1958)
stated that, "'.A. glahratus seems little affected by the amount of
light whicli reaches its habitat. It is often found exposed to direct
sunlight and in habits partially or well shaded." On the other
hand, Deschiens (1957) maintained that glabrata withdrew into its
shell immediately when subjected to a strong direct light. Deschiens'
belief that B. glabrcta favored darkness and light of a low intensit}',
led he and Bijan to study the behavior of this snail mider conditions
of total darkness for a period of 90 days. Deschiens and Bijan
(1956) noted that, "que ce mollusque conser\'e. dans ces conditions,
son comportement et son acti\ite nonnaux; le rytlime des pontes,
leur nombre. la croissance des embrv'ons, des lar^■es et des foniies
juveniles ne sont pas entravees par I'absence de lumiere." Although
reproductive activity- appeared nonnal tlie autliors gave no specific
numbers of clutches, eggs, or percent hatchabilit}'.
The data presented here show differences in egg-laying of B.
glabrata wiili relation to light energ}' and day length.
Materials and ^lethods — All snails used in this study were main-
tained individually in 250 ml of dechlorinated tap water at a tem-
perature of 23 to 25 C. Plaster of Paris was added at tlie rate of
2.5 mg per liter of water. The water, not being aerated, was
changed ever}- third da}'. Romaine lettuce was always available
for the snails. A sodium alginate food supplement was given the
snails one da}' per week.
Five gi'oups of snails were maintained under the followir.g con-
ditions of light period and energ}'.
1) continuous darkness (the changing of water, feeding and
counting of eggs required a certain amount of light. These
light periods were of short duration, usually less than 10
minutes each day, and at a very low — but unmeasured — light
energy level).
2) continuous light at 355 uav cm-
October, 1971
NAUTILUS
45
3; continuous light at 890 /iw/cm^
4) 12-hour dark period/ 12-hour light period at 355 ^tw/cm^
5) 12-hour dark period '12-hour light period at 890 fiw/cm-
Light energy levels were attained through the combination of
four 8-foot, 40-watt cool white fluorescent lamps and twelve 25-watt
incandescent bulbs. The light energ}^ of this source three feet above
the snails was calculated at 890 /iw/cm- with 617 aw/cm- in the
visible spectrum and 273 ftw/crar in the infra red spectrum.
5 J59
2 JM-
2 03-
2 jn-i
1 2 3
WEEKS
Fig. 1
m OJO-
2 2
WE Etc 3
F'9. 3
Figure 1. Mean clutches per snail per hour fa) and mean eggs per snail per
hour (b) laid hy B. glahraza maintained under cGnunuous darkness. Closed
circles represent clutches and eggs laid during the nighttime (8 P.^L to 8 AJ»L) ;
open circles represent clutches and eggs laid during the dajtime (8 .\M. to 8
P3^). Numbers abcrve closed circles in (a) represent ovipositing snails.
Figure 2. Mean clutches per snail per hour (a) and mean eggs per snail per
hour (b) laid by B. glabrata maintained under continuous light at an energy
le-vel of 355 ^w/cm^. Closed circles represent clutches and eggs laid during the
nighttime (8 P.M. to 8 A.M); open circles represent clutches and eggs laid
during the daytime (8 A_M. to 8 P.\L). Numbers above closed circles in (a)
represent ovipositing snails.
Figure 3. Mean clutches per snail per hour (a) and mean eggs per snail per
hour (b) laid by B. giabrata maintained und^ continuous light at an energy
level of 890 /iw/cm^. Closed circles represent clutches and eggs laid during the
nighttime (8 P.M. to 8 AJvI.) ; open circles represent clutches and eggs laid
during the daytime (8 A.M. to 8 P-M). Numbers above closed circles in (a)
represent ovipositing snails.
46
NAUTILUS
Vol. 85 (2)
To obtain a lower light energy level with the same spectral dis-
tribution, three layers of aluminum screening were placed ten inches
above the snails. This procedure reduced the original energy of
890 ^w/cnr to a total energy of 355 /.iw/cm^ with 250 juw/cm^ in
the visible spectrum and 105 ^iiw/cm-^ in the infra red spectrum.
Measurement of the light source was done with an ISCO Model SR
Spectroradiometer having a wave length range of 380 to 1050 mja.
Approximately three weeks after the onset of oviposition, clutches
and eggs of all snails were counted at 8 A.M. (to ascertain night-
.09-1
.06-
X
y
3 .03-
(J
y^
-o o
1.50 1
< 1.00-
z
o
{2 0.50-1
I I
1 2
WEEKS
Fig. 4
.09 1
.06-
T\ 03
2j00n
1.50-
uT 1.00*
< 0.50-1
1 1 I-
1 2 3
W E EKS
Fig. 5
Figure 4. Mean clutches per snail per hour (a) and mean eggs per snail per
hour (b) laid by B. glabrata maintained under a 12-hour/l 2-hour day. The
light period was at an energy level of 355 fiw/crrfi. Closed circles represent
clutches and eggs laid during the nighttime (8 P.M. to 8 A.M.); open circles
represent clutches and eggs laid during the daytime (8 A.M. to 8 P.M.). Num-
bers above closed circles in (a) represent ovipositing snails.
Figure 5. Mean clutches per snail per hour (a) and mean eggs per snail per
hour (b) laid by jB. glabrata maintained under a 12-hour/12-hour day. The
light period was at an energy level of 890 Atw/cm2. Closed circles represent
clutches and eggs laid during the nighttime (8 P.M. to 8 A.M.); open circles
represent clutches and eggs laid during the daytime (8 A.M to 8 P.M.). Num-
bers above closed circles in (a) represent ovipositing snails.
October, 1971 nautilus 47
time egg-laying activity regardless of whether the snails were in
darkness or light) and at 8 P.M. to determine the number of
clutches and eggs laid during the daytime (again disregarding light
conditions).
Results — ^There is no evidence to suggest a strict correlation of
egg-laying with light conditions. Although a predisposition to
nocturnal egg-laying existed in all groups, the disparity between
nighttime and daytime oviposition was lessened in those groups
maintained under continuous conditions of darkness and light
(Figs. 1 thru 5; Table 1). Mean egg-laying rates for the entire
experimental period, along with the rate of nighttime egg-laying
over that of the daytime, are shown in Table 1 .
Discussion — That B. glabrata "prefers" the nighttime for oviposi-
tion has been demonstrated. They retain this ability, although to
a lesser degree, even when maintained in continuous darkness or
light for extended periods of time. The same phenomenon has been
observed in other snails. Pesigan et al (1958), in working with
200 female snails on 12-hour/ 12-hour days for a period of 5 days,
noted that Oncomelania quadrasi favors the nighttime over daytime
for oviposition by a ratio of nearly 3 to 1. Clapp (1921) in working
with the river limpet, Ancylus fuscus, Cole (1925) in studying
Planorbarius corneus and Helisoma trivolvis, and Krull (1931) in
working with Gyraulus parvus, all observed that egg-laying occurred
at night. Cole even had groups of snails in continuous darkness
and continuous light which deposited their eggs at night. These
experiments were, however, of short duration (96 hours). In addi-
tional experiments Cole reversed the day and found that "Under
12-hour/12:5hour
(890 ;aw/cm ) 1.992 ■. 0.935 . 2.1
48 NAUTILUS Vol. 85 (2)
these conditions 14 masses were deposited, all of them during the
night, even though the normal environmental conditions of day and
night had been reversed."
Another point of interest concerns the snails maintained under
continuous light at 355 /xw/cm^. The snails in this group had the
highest rate of egg-laying in the daytime and second highest during
the nighttime (Table 1 ) . This is not too surprising, however, since
increased oviposition in continuous light has been noted before.
Wagner (1954-55) reported a definite trend toward increased
production of young by snails in continuous light. Van der Schalie
and Davis (1968) reported the same was true of Oncomelanin.
hupensis iormosana, as more young were produced when 5 males
and 5 females were kept in continuous light of 70-100 footcandles.
On the other hand, while Deschiens and Bijan (1956) believed the
reproductive activity of B. glabrata under continuous darkness to
be normal, this study reveals that of the five groups tested the
continuous darkness group had the poorest oviposition rates as
determined by eggs/snail/hour.
Clutch patterns (Figs, la thru 5a) have been included to empha-
size a minor point. They paralleled the egg patterns (Figs, lb thru
5b) quite well, indicating that by the fourth week of oviposition the
number of eggs/clutch had stabilized.
Acknowledgments — I would like to thank Dr. R. D. Powell and
Mr. Bobbie McMichael of the Department of Plant Sciences, Texas
A&M University, for use of laboratory equipment and providing
assistance in calculating light energy levels.
Literature Cited
Abbott, R. T. 1948. Handbook of medically important mollusks of the Orient
and the Western Pacific. Bull. Mus. Comp. Zool. Harvard. 100: 245-328.
Clapp, W. F. 1921. Eggs and young of the river limpet, Ancylus fuscus C. B.
Adams. Boston Soc. Nat. Hist. Occ. Papers. 5: 5-10.
Cole, W. H. 1925. Egg- laying in two species of Planorbis. Am. Nat. 59:
284-286.
Deschiens, R. 1957. Sur la perpetuation des elevages des mollusques vecteurs
des bilharzioses a I'obscurit^. Bull. Soc. Path. Exot. 50: 229-233.
and H. Bijan. 1956. Comf)ortement d'^levages do mollusques
vecteurs des bilharzioses a I'obscurit^. Bull. Soc. Path. Exot. 49: 658-661.
Harry, H.W. and D. V. Aldrich. 1958. The ecology of Australorbis glabratus
in Puerto Rico. Bull. WHO. 18: 819-832.
Kawanoto, S. 1952. On the photophobotaxis of Oncomclania nosophora. Med.
Biol. 23: 76-79.
Komiya, Y., K. Kuniko, and C. Koyama. 1959. A simple breeding method for
Oncomelania using a Petri dish. Jap. J. Parasitol. 8: 721-724.
October, 1971 nautilus 49
Krull, W. H. 1931. Importance of laboratory-raised snails in helminthology
with life history notes on Gyrciulus parvus. Mich. Univ. Mus. Zool. Occ.
Papers. 70(226): 1-10.
Moose, J. W. and J. E. WilHams. 1961-62. Medical General Laboratory (406),
U.S. Army Medical Command, Japan, Professional Reports. (Not seen; vide
van der Schalie).
Pesigan, T. P., N. G. Hairston, J. J. Jauregui, E. G. Garcia, A. T. Santos, B. C.
Santos, and A. A. Gesa. 1958. Studies on Schistosoma japonicum infections
in the Philippines. 2. The molluscan host. Bull. WHO. 18: 481-578.
van der Schalie, H. and G. M. Davis. 1968. Culturing Oncomelania snails
(Prosobranchia: Hydrobiidae) for studies of Oriental schistosomiasis. Mala-
cologia 6; 321-367.
Wagner, E. D. 1954-55. Annual progress report to the commission on parasitic
diseases of the U. S. Armed Forces Epidemiological Board, Loma Linda Uni-
versity, Loma Linda, Calif., U.S.A. (Not seen; vide van der Schalie and
Davis).
COISVS PATAE, A NEW CARIBBEAN GASTROPOD
By R. Tucker Abbott
Delaware Museum of Natural History
Through the kindness of Mrs. Patricia Nelson Ware, Miss Eliza-
beth K. Stapleton, J. M. Humfrey and Thomas L. McGinty, a new
species of Conus from Florida and Jamaica has been submitted for
description. I take pleasure in naming it for the original discoverer,
Mrs. Patricia N. Ware of Florida. This new species somewhat
resembles Conus mus Hwass, 1792, and C. cardinalis Hwass, 1792.
Conus patae, new species FigS- 1-6
Description — Shell 21 to 25 mm. in length, moderately heavy,
spirally sculptured and weakly and axially ridged. Whorls 71/2 or
8'/2- almost straight-sided. Shoulder carinate and smooth, although
in some specimens it may be weakly undulating. Color of shell
ivory-white with a blush of lavender at the base and a band of
tincture of rose at the shoulder. Over this are a few irregular, small
blotches of light brown. In Florida specimens which have been
buried in offshore sands, the lavender color has faded and the
brown blotches are yellow. Spire fairly low, straight to slightly
concave and with an angle of about 100°. Top of whorls concave
in the spire and have about 6 raised spiral threads crossed by
microscopic axial scratches. The thin, brown periostracum in live
specimens is similarly sculptured. Top of whorls with 6 to 8 color
blotches per whorl. Sides of whorls with long, axial, conspicuous,
rounded plaits, crossed by about two dozen spiral threads. Interior
of aperture pinkish in fresh specimens, white in dead ones. Soft
parts and operculum unknown.
50
NAUTILUS
Vol. 85 (2)
Figs. 1-6. Conus patae Abbott, new species. 1, holotype, 24.4 mm. 2, para-
type from Jamaica, 24.0 mm. 3 and 6, paratype from Jamaica, 20.8 mm. 4 and
5, paratype from off Lauderdale-by-the-Sea, 24.5 mm. (Del. Mus. Nat. Hist.
no. 40595).
Types — The type locality is 10 fathoms, from pipe dredgings, off
Pompano Beach, Broward County, Florida. Patricia N. Ware, coll.
1970. The holotype is in the Delaware Museum of Natural His-
October, 1971 nautilus 51
tory, No. 44097. Paratype from the same locality, DMNH 44096;
3 paratypes (DMNH 40595) from dredgings, 50 to 100 feet depth,
1/2 mile oflF Lauderdale-by-the-Sea, Elizabeth K. Stapleton, coll.
1970. Two paratypes (DMNH No. 44095) from Ocho Rios, north
central coast of Jamaica. J. M. Humfrey, coll. 1970, in 45 feet of
water near coral reefs; 3 paratypes from 80 feet, off Pompano Beach
from the John Finlay collection.
Range — From southeast Florida to Jamaica.
Remarks — Conus patae has the same general shape as Conus
daucus Hwass, 1792, and C. juliae Clench, 1942, but differs in
having strong axial and spiral sculpturing and lacking any color in
the nuclear whorls. C. patae has a non-coronate shoulder, unlike
C. mus Hwass, 1792, C. cardinalis Hwass, 1792, and the various
varicose forms of jaspideus Gmelin, 1791, such as verrucosus Hwass,
1792, and havanensis Aguayo and Farfante, 1947. The most out-
standing feature of this new species is the series of fine axial folds
that extend the length of the last whorl. Fresh specimens from
Jamaica have a pink to lavender blush that is more pronounced at
the base.
REVIEW OF PARODOSTOMIA, TELLODA,
GOmODOSTOMlA AND EVLIMASTOMA
(GASTROPODA: PYRAMIDELLACEA)
By James X. Corgan
Austin Peay State University
Clarkesville, Tennessee 37040
In 1959, Laseron (p. 200) concluded that the generic name
Scalenostoma Deshayes, 1863, had been incorrectly applied in
studies of the Australian fauna. All well-known Australian "Scal-
enostoma" are pyramidellacean gastropods of the odostomiid stock.
They have heterostrophic protoconchs, prominent parietal plica-
tions, and convex or flat-sided whorl profiles. Scalenostoma car-
inatum Deshayes, 1863, the type species of Scalenostoma, is poorly
known but it lacks a parietal plication and has a concave whorl
profile. Laseron doubted that Scalenostoma Deshayes belonged in
the Pyramidellacea.
Parodostomia Laseron, 1959, was proposed for Pacific pyram-
idellacean gastropods that had been incorrectly assigned to the
genus Scalenostoma Deshayes. The type species, Odostomia compta
Brazier, 1877, and two other Australian species were unequivocally
52 NAUTILUS Vol. 85 (2)
referred to Parodostomia. A North American species, Odostomia
(Scalenostoma) dotella Dall and Bartsch, 1909, was referred to
Parodostomia with query and Laseron suggested that a New Zea-
landic species might also belong in his new genus. Laseron's concept
of Scalenostoma Deshayes has won wide acceptance and Parodos-
tomia Laseron is a commonly used name (e.g.: Iredale and McMi-
chael, 1962).
Laseron was not the first person to note pronounced differences
between Scalenostoma Deshayes and Pacific species referred to
Scalenostoma. Telloda Hertlein and Strong (1951, p. 104) was
proposed for an Eastern Pacific pyramidellacean species that had
been placed in the eulimacean genus Scalenostoma Deshayes.
Though no general review of Scalenostoma was attempted, Hertlein
and Strong clearly meant Telloda to include species like those
Laseron placed in Parodostomia.
By original designation, the type species of Telloda Hertlein and
Strong is Odostomia (Scalenostoma) dotella Dall and Bartsch, 1909.
Yet, according to Laseron (1959, p. 200), ". . . S. dotella Dall and
Bartsch is apparently a Parodostomia." The type species of Telloda
and the type species of Parodostomia are very similar. They differ
in a single character. Species assigned to Parodostomia have spiral
sculpture on the interior of the outer lip while such sculpture is
absent in species assigned to Telloda. Traditionally, great taxonomic
significance is assigned to the presence of spiral sculpture on the
interior of the outer lip of pyramidellids (e.g.: Dall and Bartsch,
1909; Laws, 1940). In Parodostomia and Telloda all morphologic
features except internal lirations are virtually identical. Species
assigned to Parodostomia are, thus, regarded as congeneric with
those assigned to Telloda. Since species assigned to Parodostomia
and those placed in Telloda do differ in one important shell char-
acter and have distinctly different geographic distributions, they
are here regarded as subgenerically distinct.
Both Telloda and Parodostomia were established for pyramidel-
lacean gastropods previously placed in the eulimacean genus Scal-
enostoma. Prior to the establishment of either Parodostomia or
Telloda, Bartsch (1916) had concluded that Sca/enostoma Deshayes
should be placed in the eulimacean family Eulimidae. He also
reviewed American and African species of Scalenostoma. For
Odostomia (Scalenostoma) dotella Dall and Bartsch, 1909, a new
October, 1971 nautilus 53
taxon was proposed: Eulimastoma Bartsch, 1916. Since type species
are the same, Eulimastoma Bartsch, 1916, is a senior objective syn-
onym of Telloda Hertlein and Strong, 1951. It is also a senior sub-
jective synonym of Parodostomia Laseron, 1959, and Parodostomia
should be ranked as a subgenus of Eulimastoma. Eulimastoma
appears to be the oldest valid name for gastropods congeneric with
Odostomia (ScalenostomaJ dotella Dall and Bartsch, 1909.
Some indication of a possible synonymy between Parodostomia,
Telloda, and Eulimastoma is implicit in the original descriptions.
Each nominal genus involves both Scalenostoma Deshayes and
Odostomia dotella Dall and Bartsch. Goniodostomia Pilsbry and
Johnson (1917, p. 181) is here interpreted as another synonym of
Eulimastoma Bartsch, 1916, but the basis for this synonymy is
considerably more complex. The name Goniodostomia should
never have been proposed. It was introduced as the result of a
series of errors.
Eulimastoma Bartsch, 1916, was proposed to rectify an erroneous
interpretation of Scalenostoma Deshayes made by Dall and Bartsch
(e.g.: 1909). The one species referred to Eulimastoma by Bartsch
(1916) had been described as a Scalenostoma hy Dall and Bartsch
(1909). The species was not redescribed and Eulimastoma was not
formally characterized by Bartsch (1916). In Dall and Bartsch
(1909, p. 229), Scalenostoma Deshayes is diagnosed as "Smooth
Odostomias having a peripheral keel." Unfortunately, the first
species described under Scalenostoma in Dall and Bartsch (1909)
has an abundance of microscopic spiral striations and lacks a keel,
though the periphery is angular. Since it is neither smooth nor
keeled, this species bears no relationship to the genus-level diag-
nosis under which it was described. This is the species, Odostomia
dotella Dall and Bartsch, that became the type species of Eulimas-
toma Bartsch. Thus, Eulimastoma must include species with spiral
microsculpture. Pilsbry and Johnson (1917) apparently took the
diagnosis of Scalenostoma by Dall and Bartsch (1909) at face value.
They placed Eulimastoma-like species from the Oligocene of His-
panola in two groups. Smooth species were placed in Eulimastoma,
in accordance with the Dall and Bartsch (1909) diagnosis of
Scalenostoma. For species with spiral microsculpture, like the type
species of Eulimastoma, they created a new taxon, Goniodostomia
Pilsbry and Johnson (1917, p. 181).
54 NAUTILUS Vol. 85 (2)
Clearly, Goniodostomia Pilsbry and Johnson, 1917, must be
ranked as a junior subjective synonym of Eulimastoma Bartsch,
1916. Species that Pilsbry and Johnson placed in Eulimastoma pose
a more significant taxonomic problem. They do lack spiral micro-
sculpture and are, thus, morphologically distinct from the type
species of Eulimastoma. In odostomiids, and in other pyramidel-
lacean families, great taxonomic significance has been assigned to
the presence, or absence, of faint spiral striations (e.g.: Dall and
Bartsch, 1909). Recent research suggests that the mere presence or
absence of spiral microsculpture may not be taxonomically signifi-
cant. One recent study (Corgan, 1969) shows that similar micro-
scopic ornamentation can be produced by two different morphologic
features, superficial sculpture and internal structural differences
within the shell wall. Spiral micro-ornamentation is, thus, a diffi-
cult character to use as a taxobasis. Personal experience shows that
odostomiids may have spiral microsculpture at one ontogenetic stage
and lack it at another. In studies of monospecific populations of
adult odostomiids, both Willett (1937) and Abbott (1958, p. 102)
concluded that presence, or absence, of spiral microsculpture alone
was not a taxonomically significant character at the species level.
All data suggest that faint spiral striations can not be reasonably
cited as evidence of genus-level distinction. Species Pilsbry and
Johnson placed in Eulimastoma are properly referred to that genus.
The equivalence, or near equivalence, of the names Eulimastoma,
Goniodostomia, Telloda, Parodostomia, and "Scalenostoma" has
passed unnoted and, in the literature of the last few decades, new
congeneric species have been proposed under different generic
names. In addition, Telloda, Eulimastomji, Goniodostomia, and
"Scalenostomd" are, at least occasionally, ranked as subgenera of
Odostomia Fleming, 1813. Thus, six genus-level names are cur-
rently used for congeneric species. To clarify the contest, character-
istics, and distribution of Eulimastoma Bartsch, 1916, the genus is
briefly reviewed below.
Superfamily Pyramidellacea
Family Odostomudae
Genus EULIMASTOMA Bartsch, 1916
Eulimastoma BARTSCH (1916, p. 73).
Goniodostomia PILSBRY AND JOHNSON (1917, p. 181). Type
species by original designation Odostomia (Goniodostomia)
superans Pilsbry and Johnson.
October, 1971 nautilus 55
Telloda HERTLEIN AND STRONG (1951. p. 104). Type species
by original designation Odostomia (ScalenostomaJ dotella Dall
and Bartsch.
Parodostomia LASERON (1959, p. 200). Type species by original
designation Odostomia compta Brazier.
Type Species — By original designation, Odostomia (Scalenos-
tomaJ dotella Dall and Bartsch.
Diagnosis — Small to minute, high spired marine gastropods with
a single columellar plication that is generally expressed as a promi-
nent parietal tooth; with a pronounced angulation at the periphery;
with or without faint spiral striations; late teleoconch whorls gen-
erally attached low on the base of earlier whorls, exposing part of
the earlier whorl base; protoconch heterostrophic, low-spired, gen-
erally about tw'o whorls, unornamented, usually deeply and some-
times completely immersed in the first teleoconch whorl; teleoconch
whorls slightly convex to flat-sided; aperture generally oval, with
or without faint spiral sculpture inside the outer lip; with or without
umbilicus; known adult size range from about 1.2 to 5.5 mm,
average adult size about 2.5 mm.
Status — Many authors rank Eulimastoma, or a synonym, as a
subgenus of Odostomia Fleming, 1813 (e.g.: Hertlein and Strong,
1951). Others rank Eulimastoma, or a synonym, as a genus and
place it near the genus Eulimella, which is usually attributed to
Forbes, 1846 (e.g.: Cossmann, 1921). Most modern workers treat
Eulimastoma, or a synonym, as a genus and place is near Odostomia
(e.g.: Bartsch, 1955).
An immersed, to partly immersed, heterostrophic protoconch,
presence of a columellar plication, and a dirth of sculpture suggest
that Eulimastoma does belong to the odostomid stock. Slightly
convex to flat-sided whorls and a high spired growth form make
Eulimastoma quite distinct from Odostomia. Morphologic distinc-
tions, wide geographic distribution, and a significant duration in
time combine to suggest that Eulimastoma Bartsch, 1916, should be
ranked as a genus of the Odostomiidae.
Ecology — Odostomiids are external parasites of mollusks and/or
worms (e.g.: Fretter and Graham, 1962). Dietary preferences of
Eulimastoma have not been described. Dredged specimens have
been recovered between 24 to 162 feet. The exact depth range of
living specimens is unknown but data suggest that the genus is
restricted to the inner Continental Shelf. Laseron (1959, p. 201)
56 NAUTILUS Vol. 85 (2)
associates E. compta (Brazier) with a grass substrate. It may well
occur on a variety of substrates and it may, or may not, be typical
of the genus.
Subgenera — Two subgenera are recognized: Eulimastoma (Euli-
mastoma) and Eulimastoma (Parodostomia):
Eulimastoma (Eulimastoma)
Diagnosis — Eulimastoma without spiral sti'iations on the inner
surface of the outer lip.
Geologic Range — Oligocene to modern.
Content — One species is known from the Oligocene of New
Zealand: Scalenostoma southlandica Laws (1940, p. 158, pi. 13,
fig. 5). Six nominal species, first illustrated by Pilsbry (1922), have
been collected from the Oligocene of Hispanola. Two of these
species, with faint columellar plications, have not previously been
placed in Eulimastoma. They are Odostomia (Evalea?) vexator
Pilsbry and Johnson (1917, p. 180) and O. santodomingensis Pilsbry
and Johnson (1917, p. 179). These seem to be meaningful species.
The name O. santodomingensis originated as a replacement for
Aclis polita Gabb (1881, p. 226). Though the grounds offered for
replacement are no longer valid, there is an Aclis polita Verrill
(1872, p. 282) and the replacement, thus, is necessary.
Odostomia ( Goniodostomia) superans Pilsbry and Johnson (1917,
p. 180>) and O. (G.) circumvincta Pilsbry and Johnson (1917, p.
181) are rather similar species but, apparently, distinct. It seems
best to rank Odostomia (Eulimastoma) pyrgulopsis Pilsbry and
Johnson (1917, p. 179) as a senior synon}'m of O. (E.) bathyraphe
Pilsbry and Johnson (1917, p. 180). It has page priority and O.
bathyraphe seems to be an unusually broad individual of the same
species.
With seven nominal species of Eulimastoma known from the
Oligocene, it is surprising that Miocene species seem to be unre-
corded. One Pliocene species from Florida has been described
under two names: Eulimastoma harbisonae Bartsch (1955, p. 82,
pi. 16, fig. 1) and E. olssoni Bartsch (1955, p. 83, pi. 16, fig. 5).
These nominal species occur together and differ only in the pres-
ence, or absence, of an umbilicus. Personal experience with Euli-
mastoma and the observations of Laseron (1959, p. 200-201) show
that both umbilicate and non-umbilicate individuals occur in mono-
specific populations of Eulimastoma. Since it has page priority,
October, 1971 nautilus 57
Eulimastoma harbisonae Bartsch, 1955, is here selected as the senior
synonym.
An unpublished dissertation (Corgan, 1967) describes and illus-
trates two new species of Eulimastoma from Quarternary sediments
of the Mississippi River Delta. New species are part of a cool water
fauna that inhabited the area some 15,500 years ago. It seems
probable that these species are still part of the Carolinian fauna of
the Gulf of Mexico. Since the dissertation was prepared, I have
seen two additional new species of Eulimastoma in modern faunal
collections made by Miss Jean Andrews on the Texas Coast.
Described modern species of Eulimastoma (Eulimastoma) are
known only from the tropical coasts of western North America. The
two known species are Odostomia (Scalenostoma) dotella Dall and
Bartsch (1909, p. 230, pi. 30, fig. 5) and Odostomia (Telloda) sub-
dotella Hertlein and Strong (1951, p. 104, pi. 8, fig. 5). Both seem
to be meaningful taxa.
Eulimastoma (Par odostomia)
Diagnosis — Eulimastoma with spiral striations on the inner sur-
face of the outer lip.
Geologic Range — Known only from modern faunas.
Content — Eulimastoma (Par odostomia) is known only from the
Pacific. Two species occur in the fauna of New South Wales:
Scalenostoma subcarina Laseron (1951, p. 308, fig. 21) and S. pyra-
midata Laseron (1951, p. 308, fig. 24). One species occurs in
northern Australia: Odostomia compta Brazier (1877, p. 259).
Laseron (1959, p. 200, figs. 46-48) provides illustrations and a
synonymy. He correctly treats O. affinis Brazier (1877, p. 259),
O. parvula Brazier (1877, p. 260), and O. polita Brazier (1877,
p. 260) as junior synonyms of O. compta Brazier. The only other
known species of Eulimastoma (Parodostomia) is Odostomia
eutropia Melvill (1899, p. 94, pi. 1, fig. 14) from Pakistan.
Eulimastoma ?
A fairly detailed review of the world odostomiid fauna suggests
that no other described species can be unequivocally referred to
Eulimastoma. A few briefly described and/or unillustrated species
can not be adequately evaluated. Odostomia ijaquica Maury (1917,
p. 151, pi. 25, fig. 22) may be a Eulimastoma. It is, apparently,
known only from the holotype which came from Hispanola and is
probably of Oligocene age.
58 NAUTILUS Vol. 85 (2)
Odostomia engonia Bush (1885, p. 466) and O. engonia var.
teres Bush (1885, p. 467, pi. 45, fig. 9) resemble Eulimastoma in
growth form and sutural characteristics. Protoconch-teleoconch
relationships are not adequately described for either nominal taxon
and from published records they can not be identified with cer-
tainty. Garcia-Cubas (1963, p. 45, pi. 4, fig. 9) has recorded
Pyramidella engonia var. teres (Bush) from southeastern Mexico.
Like Bush, Garcia-Cubas does not provide a detailed description
but his illustration is good and the species is apparently a Eulimas-
toma. If the identification is correct. Bush's taxon is very widespread.
Summary
Eulimastoma Bartsch, 1916, must be regarded as a common and
widespread genus. It ranges from roughly 35° north, in the Atlan-
tic, to about 35° south, in the Pacific, and extends back in time
to the Oligocene. Eulimastoma and its subgenus Parodostomia
Laseron, 1959, include, or may include, the 22 nominal species
listed below. Species that, to the best of my knowledge, have not
previously been placed in Eulimastoma s.l. are indicated by an
asterisk. Each species listed is cited and discussed in the text.
Eulimastoma (Eulimastoma)
circumvincta (Pilsbry and Johnson); dotella (Dall and Bartsch);
harbisonae Bartsch (syn.: olssoni Bartsch); pyrgulopsis (Pilsbry
and Johnson) (syn.: bathyraphe (P. and J.)); *santodomingensis
(Pilsbry and Johnson) (syn.: polita (Gabb)); southlandica
(Laws); subdotella (Hertlein and Strong); superans (Pilsbry and
Johnson); *vexator (Pilsbry and Johnson).
Eulimastoma (Parodostomia)
compta (Brazier) (syn.: affinis (Brazier), parvula (Brazier) and
polita (Brazier)); *eutropria (Melvill); pyramidata (Laseron);
subcarina (Laseron).
Eulimastoma ?
engonia (Bush); teres (Bush); yaquica (Maury).
Reflrences Cited
Abbott, R. T., 1958, Marine mollusks of Grand Cayman Island,
British West Indies: Mon. Acad. Nat. Sci. Philadelphia, no. 11.
138 pp. 5 pis.
Bartsch, P., 1916, Eulimastoma a new subgenus of pyramidellids
and remarks on the genus Scalenostoma: Nautilus 30, p. 73-74.
, 1955, The pyramidellid mollusks of the Pliocene deposits of
North St. Petersburg, Florida: Smithsonian Inst. Misc. Coll. 125,
no. 2. 102 pp. 18 pis.
October, 1971 nautilus 59
Brazier, J. W., 1877, Shells collected during the Chevert Expedition,
with descriptions of new species: Proc. Linn. Soc. New South
Wales /, p. 249-261.
Bush, K. J., 1885, Additions to the shallow- water Mollusca of Cape
Hatteras . . .: Trans. Connecticut Acad. Arts Sci. 6, p. 452-480.
Corgan, J. X., 1967, Quarternary micromolluscan fauna of the
Mudlump Province, Mississippi River Delta: Unpub. Ph.D. dis-
sert., Louisiana State Univ. 300 pp. 8 pis.
, 1969, Odostomia cassandra Bartsch: Nautilus 83, p. 71-72.
Cossmann, M., 1921, Essai de paleoconchologle comparee. Pt. 12:
Privately printed. 348 pp. 6 pis.
Dall, W. H., and Bartsch, P., 1909, A monograph of the West
American pyramidellid mollusks: U.S. Nat. Mus. Bull. 68. 258
pp. 30 pis.
Fretter, V., and Graham, A., 1962, British prosobranch mollusks:
The Ray Society. 755 pp.
Gabb, W. M., 1881, On the topography and geology of Santo
Domingo: Trans. Amer. Phil. Soc. 15, pt. 2, p. 49-260.
Garcia-Cubas, A., 1963, Sistematica y distribution de los micro-
moluscos recientes de la laguna de terminos, Campeche, Mexico:
Univ. Nac. Aut. Mexico, Inst. Geol. Bol. no 67, pt. 4. 55 pp. 4 pis.
Herdein, L. G., and Strong, A. M., 1951, Eastern Pacific expeditions
of the New York Zoological Society. XLIIL Mollusks from the
west coast of Mexico and Central America: Zoologica 36, p.
67-120.
Iredale, T., and McMichael, D. P., 1962, A reference list of the
marine Mollusca of New South Wales: Mem. 11, Australian Mus.
109 pp.
Laseron, C. P., 1951, The New South Wales Pyramidellidae and the
genus Mathilda: Rec. Australian Mus. 22, p. 298-334.
, 1959, The Family Pyramidellidae (Mollusca) from North-
ern Australia: Australian Jour. Marine Freshwater Res. 10, p.
177-267.
Laws, C. R., 1940, Review of the Tertiary and Recent Neozelanic
pyramidellid molluscs. No. 7. Further odostomid (sic) genera:
Trans. Royal Soc. New Zealand 70, p. 150-160.
Maury, C. J., 1917, Santo Domingo type sections and fossils. Pt. 1.
Mollusca: Bull. Amer. Paleont, no. 29. 251 pp. 39 pis.
Melvill, J. C, 1899, Notes on the Mollusca of the Arabian Sea,
Persian Gulf, and Gulf of Oman . . .: Ann. Mag. Nat. Hist.,
ser. 7, 4, p. 81-101.
Pilsbry, H. A., 1922, Revision of W. M. Gabb's Tertiary Mollusca
of Santo Domingo: Proc. Acad. Sci. Philadelphia 73, p. 305-435.
, and Johnson, C. W., 1917, New Mollusca from the Santo
Domingo Oligocene: Proc. Acad. Nat. Sci. Philadelphia 6,9, p.
150-205.
Verrill, A. E., 1872, Recent additions to the Molluscan fauna of
60
NAUTILUS
Vol. 85 (2)
New England . . .: Amer. Jour. Sci., ser. 3, 3, p. 281-290.
Willett, G., 1937, An Upper Pliocene fauna from the Baldwin Hills,
Los Angeles County, California: Trans. San Diego Soc. Nat. Hist.
8, p. 379-406.
NOTES ON ALASMIDOISTA FABULA (LEA)
IN KENTUCKY (UNIONIDAE)
By Shaw Blankenship
Department of Biological Sciences
Eastern Kentucky University, Richmond, Ky. 40475
The mollusk fauna of Kentucky is fairly well-known by collected
material but more publications are needed (Bickel, 1967). This
paper will discuss the status of Alasmidonta (Pegias) fabula (Lea),
a small bivalve characteristic of the Cumberlandian fauna (Wilson
and Clark, 1914; Ortmann, 1924, 1925, 1926). Simpson (1914)
and Clench (1959) gave the distribution of A. fabula as the Cum-
berland and Tennessee rivers. Williamson (1905) reported A. fabula
in the Rockcastle River, a major tributary of the Cumberland.
However, no data were given as to ecology or abundance. Wilson
and Clark (1914) listed the species as rare after collecting only two
living specimens from the Cumberland Drainage. Again, the col-
Figure 1. Alasmidonta (Pegias) fabula (Lea) from Horse Lick Creek, Rock-
castle-Laurel County line, Kentucky: (A) left valve, male; (B) right valve,
male; (C) right valve, female (note highly corroded condition of specimen,
making age determination impossible; and (D) left valve, feinale.
October, 1971 nautilus 61
lection was secured from the Rockcastle River at Livingston, Ken-
tucky. An intensive search was made by Neel and Allen (1964),
but no specimens of A. fabula were obtained and thought to be
extinct.
On October 2, 1970, an empty shell was found near the mouth
of Horse Lick Creek, Rockcastle-Laurel County line, a tributary of
the Rockcastle River. One week later living specimens were located
about two miles upstream from the first collecting station. One of
each sex was taken for the initial study. Selected measurements
are: 5-year-old male (dead), length: 25.5 mm., height; 16.0 mm.
4-year-old male, length: 21.5 mm., height: 13.0 mm. Female (age
unknown), length: 25.0 mm., height: 13.5 mm.
As illustrated by the photograph, all specimens were badly cor-
roded. Not one specimen was found to be "dug in," but all were
free to be moved by the churning water causing additional wear.
It is hoped that this report will stimulate additional study as to
ecology and distribution of this unusual mussel.
Literature Cited
Bickel, D. 1967. Preliminary Checklist of Recent and Pleistocene Mollusca of
Kentucky. Sterkiana 28: 7-20.
Clench, W. J. 1959. In: Ward, H. B. and G. C. Whipple, Freshwater Biology.
John Wiley and Sons, Inc. New York. 1248 pp.
Neel, J. K. and W. R. Allen. 1964. The Mussel Fauna of the Upper Cumber-
land Basin Before Its Impoundment. Malacologia /(3): 427-459.
Ortmann, A. E. 1924. The Naiad-fauna of Duck River in Tennessee. American
Midland Naturalist. 9: 3-47.
1925. The Naiad-fauna of the Tennessee River System
Below Walden Gorge. American Midland Naturalist 9: 321-372.
1926. The Naiades of the Green River Drainage in Ken-
tucky. Annals of the Carnegie Museum 17: 167-188.
Simpson, C. T. 1914. A Descriptive Catalogue of the Naiades or Pearly Fresh-
water Mussels. Bryant Walker. Detroit, Michigan. 1540 pp.
Williamson, E. B. 1905. Odonata, Astacidae, and Unionidae Collected Along
the Rockcastle River at Livingston, Kentucky. The Ohio Naturalist 5(6):
309-312.
Wilson, C. B. and H. W. Clark. The Mussels of the Cumberland River and
Its Tributaries. U.S. Bur. Fisheries Doc. 781. 63 pp.
THE REPRODUCTIVE ANATOMY OF
TRYONIGENS REMONDl (TRYON, 1863):
HELMINTHOGLYPTIDAE
By Walter B. Miller
Department of Biological Sciences, University of Arizona
Tucson, Arizona 85721
Two shells of Tryonigens remondi (Tryon) were collected by
Auguste Remond, a French geologist, in the vicinity of Mazatlan,
62 NAUTILUS Vol. 85 (2)
Sinaloa, Mexico, and sent to George W. Tryon, Jr., who described
it as Helix Remondi Tryon (1863). The synonymy and distribu-
tion of T. remondi has been reviewed by Solem (1959).
A single living specimen from Manzanillo, Colima, was mailed
to Pilsbry (1905, p. 256) but unfortunately by the time it arrived,
it had been crushed, partly decayed, and the remains had dried
hard. Nevertheless, Pilsbry was able to make out certain details of
the lower genitalia as well as the serrated keel of the tail and an
Epiphragmophora-like radula. Based on these studies, he estab-
lished the genus Tryonigens Pilsbry (1927: 189-191).
The author and his son, Walter B. Miller III, collected several
specimens of Tryonigens remondi in December, 1962, and again in
December, 1963, in the vicinity of Mazatlan, Sinaloa. The locality
is a wooded, tropical ravine along the Mazatlan-Durango highway
at an elevation of about 2,600 feet. At that time of the year, the
snails were all hibernating. By diligent digging, however, the
10 mm
Figure 1. Lower genitalia, Tryonigens remondi (Tryon), No. 4556-A: ag,
albumin gland; ep, epiphallus; fo, free oviduct; go, genital orifice; pe, penis;
prm, penial retractor muscle; pro, prostate; sp, spermatheca; spd, spermathecal
duct; ut, uterus; va, vagina; vd, vas deferens; ve, verge. Drawing to scale indi-
cated, from stained whole mount.
October, 1971
NAUTILUS
63
author's son found a single live adult during each of the two visits.
In each case the animal had buried itself in loose dirt among the
rockslide and had developed a very thick and firm epiphragm. Dead
shells were relatively common on top of the ground or in shallow
crevices among the rocks, indicating that during its active season,
it is probably an active surface dweller rather than a deep rockslide
inhabitant.
The two animals (WBM Cat. 4397-A and 4556-A) were dis-
sected by the author and stained whole mounts of the genitalia
were prepared. The lower genitalia of No. 4556-A are shown in
Fig. 1; the details of the verges and epiphalli of both specimens
are shown in Fig. 2. Pilsbry's description of the lower genitalia is
verified, specifically the absence of a dart apparatus, the presence
of a large verge, the mode of insertion of the penial retractor, and
the absence of a visible epiphallic cecum (flagcllum). Other striking
characteristics stand out as follows:
1. The spermathecal duct is extremely short, measuring approxi-
mately 2 mm. or 2/5 of the length of the free oviduct.
2. There is no spermathecal diverticulum.
lue ep
5mm
Figure 2. Details of penis and epiphallus, Tryonigens remondi (Tryon).
A, No. 4397-A; B, No. 4556-A. ec, epiphallic cecum; ep, epiphallus; lue, lumen
of epiphallus; lup, lumen of penis; prm, penial retractor mu.scle; sdo, seminal
duct orifice; vd, vas deferens; ve, verge. Drawings to scale indicated, from
stained whole mounts.
64 NAUTILUS Vol. 85 (2)
3. The mantle has a light greyish-white color, with no other
color markings or spots of any kind.
Additional details of interest are: (1) the subterminal orifice of
the sperm duct in the verge, (2) the absence of a penial sheath,
(3) the variability in length and shape of the verge in the two dis-
sected specimens, and (4) the very short internal cecum on the
epiphallic duct (Fig. 3) at its junction with the vas deferens. This
latter cecum is not visible externally but is very probably a vestige
of an ancestral, larger epiphallic cecum (flagellum).
Pilsbry's question of the affinity or parallelism of Tryonigens
with Sonorella must remain in abeyance, at least as far as the com-
parative anatomy of the genitalia is concerned. Like Sonorella,
Tryonigens has a verge, no dart apparatus, and no spermathecal
diverticulum. In Sonorella, the epiphallic cecum (flagellum) is
minute or absent; in Tryonigens it is absent. In Sonorella, the
spermathecal duct is relatively long, while in Tryonigens it is
extremely short. Pilsbry considered that the serrated keel of the
tail showed a decided resemblance to Leptarionta and Lysinoe,
while the radula was similar to that of Epiphragmophora. It can
now be stated that the spermathecal duct is also like that of
Epiphragmophora.
Solem (1959) lists the range of Tryonigens remondi from Guer-
rero to Sonora at Guaymas, and to Chihuahua at Guasaremos, Rio
Mayo. The range of Tryonigens nowhere overlaps or comes in
contact with the range of Sonorella. An investigation of the south-
ern limits of Sonorella (Miller, 1967a) reveals a large area south
of latitude 29 °N in Sonora where neither Sonorella nor Tryonigens
are found.
The evidence at hand does not support a close relationship
between Sonorella and Tryonigens. Sonorella probably arose from
a marginal population of Sonorelix or Eremarionta (Miller, 1967b)
while Tryonigens probably arose from Leptarionta or a common
Leptarionta- like ancestor. This would represent an additional
instance of convergent evolution, with secondary simplification of
structures, as noted in the case of Mohavelix and other helmintho-
glyptid genera (Miller, 1970: 278).
Literature Cited
Miller, W. B. 1967a. Two new Sonorella from Sonora, Mexico, and notes on
southern limit of genus. The Nautilus 87(1): 1-9.
Miller, W. B. 1967b. Anatomical revision of the genus Sonorella (Pulmonata:
October, 1971 nautilus 65
Helminthoglyptidae). Univ. of Arizona doctoral dissertation #E979I.
Miller, W. B. 1970. A new species of Helminthoghjpta from the Mojave Desert.
The Veliger 72(3): 275-278.
Pilsbry, H. A. 1905. Mollusca of the Southwestern states, I: Urocoptidae; Hel-
icidae of Arizona and New Mexico. Proc. Acad. Nat. Sci. Phila. 57(1):
211-290, Pis. XI-XXVII.
Pilsbry, H. A. 1927. The structure and affinities of Humboldtiana and related
helicid genera of Mexico and Texas. Proc. Acad. Nat. Sci. Phila. 79.- 165-192,
Pis. 11-14.
Solem, A. 1959. Notes on Mexican mollusks. II. Occ. Papers Mus. Zool. Univ.
Mich., 677; 1-15.
Tryon, G. W. Jr. 1863. Descriptions of two new species of Mexican land shells.
Proc. Acad. Nat. Sci. Phila. 75(6) (Oct.-Nov., 1863): 281.
FLUORESCENCE IN MESODOJS CLAVSVS (SAY)
By Hugh C. Rawls and John M. Baum
Department of Zoology, Eastern Illinois University
Charleston, Illinois 61920
A recent paper (Rawls and Yates, 1971) reported the discovery
of fluorescence in the mucus of specimens of three genera of endo-
dontid snails exposed to long-wave ultraviolet light, and suggested
that it possibly would be found to occur in the remaining genera
of that family. We now wish to report the discovery of fluores-
cence in the mucus of specimens of Mesodon clausus (Say), appar-
ently the only polygyrid to exhibit this phenomenon.
As part of a study of the role played by bacteria in the production
of fluorescence in land snails, we had been conducting a survey of
the flora of mucus samples from various representatives when we
observed fluorescence in specimens of Mesodon clausus. Such an
observation was intriguing to us because we had not expected any
polygrid to exhibit this phenomenon. Our preliminary work with
various members of this family had led us to the assumption that
fluorescence would not be demonstrated in polygrids, and we had
therefore intended to use mucus samples from members of this
family as controls against which to compare mucus samples from
fluorescent snails. Our assumption, obviously, was wrong, and the
discovery of fluorescence in Mesodon clausus quickly caused us to
revise our thinking with regard to the occurrence of this phenom-
enon in the representatives of any given family. We now suggest
that fluorescence, whenever observed in land snails, will be found
to occur in all representatives of a given species but not necessarily
in all members of any higher group.
66 NAUTILUS Vol. 85 (2)
The discovery of fluorescent members in a family in which we
had not expected the phenomenon to be exhibited led us to carry
out a careful study of the response to ultraviolet light by specimens
of all available polygrids. We collected living representatives from
several widely separated localities in Illinois, and we utilized pre-
served specimens from numerous points throughout the range of the
family. Of all the polygrids which we subjected to ultraviolet light,
only Mesodon clausus displayed any fluorescence. The phenomenon
was observed to be uniformly characteristic in all representatives of
this species, the fluorescence being as bright and as distinctive in
preserved snails as in living specimens. The color of the fluorescence
is a greenish blue which is recognizably different from that observed
in endodontids. Spectrophotometric investigation of mucus samples
from living specimens reveals fluorescence peaks at 275 nm. and at
330nm., suggesting the presence of two fluorescent compounds in
the mucus.
The nature of the fluorescent compounds present in the mucus
of Mesodon clausus is still being investigated, as is true also of the
compounds in the mucus of fluorescent members of the Endodon-
tidae. We are not prepared now to present definitive data concern-
ing these compounds, but we do wish to comment on the presence
of certain bacteria in mucus samples from these and other snails,
because a relationship between these bacteria and the phenomenon
of fluorescence seems possible.
Of the several kinds of microorganisms which we have found in
mucus samples from land snails of various species, representatives
of the genus Pseudomonas are consistently present. These bacteria
are common in soil and water, and include representatives which
can produce fluorescent pigments ranging in color from green
through blue and violet to shades of red and yellow, according to
Breed and others (1957). For example, we have found that bac-
terial isolates from the mucus of Mesodon clausus, Mesodon thy-
roidus, Anguispira kochi and Anguispira alternata all produce
pigments which fluoresce under ultraviolet light. The association
of such bacteria with the phenomenon of fluorescence in land
snails of certain species would therefore seem to be entirely logical.
Further, the bacteria in the isolates prove to be members of the
genus Pseudomonas. Mesodon thyroidus, however, does not fluor-
esce, whereas the others do. We are thus faced with a relationship
October, 1971 nautilus 67
which at first seems valid, but which later does not appear to hold
true.
If pseudomonads are either directly or indirectly responsible for
the fluorescent properties of certain land snails, one thing seems
certain and that is that something in the physiology of these snails
permits the phenomenon to occur, whereas some factor of the
physiology of nonfluorescent snails prevents it. On the other hand,
if bacteria are not related to the phenomenon as it occurs in fluores-
cent snails, something in the physiology of these snails produces the
fluorescence. We are continuing investigations along several lines
in an attempt to resolve this problem, and we plan to report on our
findings in a subsequent paper.
Literature Cited
Breed, Robert S., E. G. D. Murray and Nathan R. Smith. 1957.
Bergey's Manual of Determinative Bacteriology. 7th ed., p. 90.
Williams and Wilkins Company, Baltimore, 1957.
Rawls, Hugh C. and Roger L. Yates. 1971. Fluorescence in Endo-
dontid Snails. The Nautilus 85(1): 17-20.
UNITED STATES RESEARCH TRENDS IN MALACOLOGY
By Dee S. Dundee
Department of Biological Sciences, Louisiana State University
New Orleans, Louisiana 70122
A recent survey was made by mc of the types of molluscan topics
appearing in 10 U.S. journals beginning in 1900. They were broken
into three periods: 1900-1920, 1921-1950, 1951-1969. It was nec-
essary to make arbitrary assignments of many papers, since their
categories were mixed or not clear-cut. The following journals were
analyzed and tabulated in figure 1.
Evolution Amer. Microscopial Soc. Trans.
Quarterly Review of Biology Comp. Biochemistry & Physiology
Ecology Jour. Experimental Zoology
Ecological Monographs Nautilus
American Midland Naturalist Malacologia
In addition, a similar analysis was made of the types of molluscan
research projects funded by the National Institutes of Health from
1960 through 1969 (see fig. 2).
68
NAUTILUS
Vol. 85 (2)
From this search it was found that research was being done over
the last ten years by less than 100 workers, a remarkable few in
view of the fact that mollusks represent the second largest group of
living animals. Until recent years, most papers dealt solely with
taxonomy and distribution. In recent years the emphasis has veered
toward ecology, behavior, cytology, and reproductive biology.
M 1900- -1920
E I921--1950
85 1951 --19)0
w »
Syttemjiics
Distcibatioii
Paleontology Sbell-Raduls Fbysjoioiiy
Ecology
Parasite
Related
J£
jSi
Embryology Mlicellaneooi Statistics
Reprodocfioa
Behavior Cytology
Figure 1. Percent of papers published on various aspects of malacology from
1900-1970. Based upon a survey of ten U. S. journals.
October, 1971
NAUTILUS
69
NO.
15
123-4 56789 123456789 123456789 123456789 12-3456789
ECOLOGY BEHAVIOR PHYSIOLOGY PATHOLOGY REPRO.-EMBRYOL.
10
123456739 123455789 123456789 123456789 123456789
NEUROPHYSIOL. CYTOL.-CELL PHYS. MOLE. BiOLrBIOCHEM. PARASITE RELATED GENETICS
Figure 2. Generalized types of malacology funded from 1961 through 1969
by National Institutes of Health.
Notes
Additions to the Cayman Islands Land Mollusks. — During
Nov. 1970, Lt. Col. Corinne Edwards of Coconut Grove, Florida
collected a few land shells on the western end of Grand Cayman
Island. Two of the few species collected are worthy of record.
These notes are an addition to a paper which was published for
these islands in 1964. (Clench, W. J. Occasional Papers on Mol-
lusks, Howard Univ. 2: 345-384, pi. 61-63).
SUCCINEIDAE
Succinea latior C. B. Adams
Succinea latior C. B. Adams 1845, Contributions to Conchology
pt. 3, p. 38 (Montego Bay, Jamaica); Clench 1964, Occ. Papers on
Mollusks, Harvard Univ. 2; 359, pi. 62, fig. 2.
Remarks: I had not seen this species from the Cayman Islands at
70 NAUTILUS Vol. 85 (2)
the time the Cayman Island report was published but listed it on
the authority of H. A. Pilsbry.
Sjyecimens examined: Georgetown, Grand Cayman.
POLYGYRIDAE
Praticolella griseola (PfeiflFer)
Helix griseola PfeiflFer 1841, Symbolae Historiam Heliceorum 1:
41 (Veracruz, Mexico).
Praticolella griseola (PfeiflFer). Pilsbry 1940, Monograph No. 3,
Acad. Nat. Sci. Philadelphia /: pt. 2, p. 690, fig. 425.
Remarks: This is the first Cayman Islands record of this intro-
duced species. It has been introduced also at Key West, Florida,
Habana Province, Cuba, and San Pedro de Macoris, Hispaniola.
It is endemic to Mexico.
Specimens examined: Georgetown, Grand Cayman. — William J.
Clench, Museum Comp. Zool., Cambridge, Mass. 02138.
MoLLUSKS OF Otsego Lake, New York — Ecological and limno-
logical studies are being undertaken on Otsego Lake, Otsego Co.,
N. Y. by researchers at the Stephen C. Clark Biological Field Station
at Cooperstown. The molluscan species, and their distributions
throughout the lake, have been determined:
Bivalvia Planorbidae
Unionidae Helisoma trivolvis (Say)
Lampsilis radiata (Gmelin) Helisoma anceps (Menke)
ElUptio complanata (Lightfoot) Helisoma campanulata (Say)
Anodonta cataracta (Say) Gyraulus parvus (Say)
Anodontoides ferussacianus (Lea) Promenetus exacuous (Say)
Strophitus undulatus (Say) Physidae
Alasmidonta undulata (Say) Physa heterostropha (Say)
Sphaeriidae Viviparidae
Pisidium correpressum (Prime) Viviparus georgianus (Lea)
Pisidium subtruncatum (Malm) Pleuroceridae
Sphaerium sulcatum (Lamark) Spirodon carinata (Bruguiere)
Gastropoda Valvatidae
Lymnaeaidae Valvata tricarinata (Say)
Lijmnaea humilis (Say) Valvata sincera (Say)
Lymnaea palustris (Miiller) Hydrobiidae
Lymnaea emarginata (Say) Amnicola limosa (Say)
Lym.naea columella (Say) Amnicola lustrica (Pilsbry)
A new state park has recently been completed at the northern
end of this body of water. Siltation from the creation of artificial
sand beaches has resulted in chronically turbid waters in that part
of the lake. I feel that this list will provide a valuable baseline for
future reference if radical changes in the fauna take place. Collec-
October, 1971 nautilus 71
tions were made over a 2 year period at more than 200 sites in from
0 to 50m of water. — ^Willard N. Harman, New York State Univer-
sity College, Oneonta, New York 13820.
Sphaertum lacustre in Oklahoma — On June 10, 1970 I col-
lected Sphaerium lacustre (Miiller) in the "Tin Horn" region of
Pennington Creek about 2.5 miles south of Tishomingo, Johnston
County, Oklahoma. Specimens were numerous in shallow side
channels but lacking elsewhere. The identification was verified by
Rev. H. B. Herrington, Westbrook, Ontario. This species has not
been reported before in Oklahoma.
PisiDiUM CoMPRESSUM IN MISSISSIPPI — On April 23, 1970 I col-
lected Pisidium compressum Prime in a small tributary of the
Strong River five miles northeast of the junction of highways 13
and 49 near Mendenhall, in Simpson County, Mississippi. The
clams were abundant in a small sand-bottomed pool about 75 feet
cast of the point where the stream passes under highway 13. The
specimens were identified by Rev. H. B. Herrington, Westbrook,
Ontario. This species has not been reported previously in Missis-
sippi. Both notes by William F. Gale, Ichthyological Associates,
RD 1, Box 306A, Berwick, Pa. 18603.
The Appearance of Pseudosuccinea columella (Say) in Ari-
zona— In Tucson, Arizona, this lymnaeid snail was collected in
November of 1970 on water hyacinth, Eichhornia crassipes (Mart.)
Solms, in a nursery which imported the plants from Florida. Of 18
specimens examined, the largest measured 12.5 mm. in length,
whereas the mean of all specimens was 10.5 mm. The relatively
small size, dark color, and heavy shell compare closely with these
characters in specimens from Florida (Baker, 1911, Special Publ. 3,
Chicago Acad. Sci., p. 170-171, discusses this small form from the
southern United States). At this point, this snail is not known to
have become established in the Tucson area, and it remains to be
seen if this species will spread from its initial introduction into
Arizona, or even if the initial colony will maintain itself. — Richard
H. Russell, Department of Biological Sciences, University of Ari-
zona, Tucson, 85721.
72 NAUTILUS Vol. 85 (2)
Veronicellids Still on the Move in the Gulf Coast —
Since 1960 the only Veronicellid slug found in the New Orleans
area in nature has been Veronicella ameghini Gambetta (Dundee,
Stutts, Hermann, 1965). From time to time Veronicella ftoridana
(Leidy) has been seen in nurseries in the area; it was only a matter
of time until they became introduced in nature. These nurseries
at v/hich V. fioridana were found demonstrated to us, by collecting
from incoming plant materials, that they were coming in on plants
from Florida!
Within the past 3 months reports of V. fioridana living in the
city have been reaching me. They are now definitely established
in two areas in the city with the possibility of others existing at the
present. Up to now the distribution of V. fioridana in nature in
the U.S. has been: various localities in south Florida, and Hodges
Gardens in western Louisiana.
Veronicella ameghini, v/hich has been in the New Orleans and
Mobile areas for ten years is also spreading. The latest finding of
it was in Lafayette, Louisiana (100 miles west) in a greenhouse and
immediately outside of it! Within the last ten years it has spread
from New Orleans and Mobile northward to about the middle of
the states of Alabama, Mississippi, and Louisiana, and now it is
very likely to become established westward also. — Dee S. Dundee,
Louisiana State Univ., New Orleans, La. 70122.
Reference Cited
Dundee, Dee S., B. S. Stutts, and P. W. Hermann. Preliminary
survey of a possible molluscan pest in the Southern United States.
Ecol. 46(1 & 2) 192-193 (1965). [For Umax and Milax, see The
Nautilus, 80(3): 108].
Strombus Range Extensions — A juvenile, 120.9 mm.-long speci-
men of S. raninus Gmelin was obtained in 1970 in 23 meters, about
69 miles east of Wilmington, N. C. (34°20'N., 76°52.3'W.) and a
juvenile, 80 mm.-long specimen of S. gigos Linne was obtained in
25 meters, about 38 miles east of Georgetown, S. C. (32°58.2'N.,
78°45.5'W.). Cruise No. E-28-70 of the R/V Eastward was sup-
ported by NSF grant No. GB 8189 to Duke University, under the
direction of Dr. F. John Vernberg. I wish to thank Dr. Robert
Robertson for identifying these specimens. — Alan H. Shoemaker,
Baruch Institute, University of South Carolina, Columbia, S. C.
29208.
October, 1971 nautilus iii
Publications Received
Starrett, William C. 1971. A sun'cy of the mussels (Unionacea)
of the Illinois River: a polluted stream. Bull. Illinois Nat. Hist.
Survey, vol. 30, art. 5, pp. 265-403 17 figs., 4 color pis. of 23
unios. Exhaustive and excellent. Obtained free from: Chief, 111.
Nat. Hist. Surv., Natural Resources Bldg., Urbana, 111. 61801.
Chcatum, E. P. and R. W. Fullington. 1971. The aquatic and land
Mollusca of Texas. Pt. 1, The Recent and Pleistocene members
of the gastropod family Polygyridae in Texas. Bull. 1, Dallas
Museum of Natural History, vi + 74, 10 pis., 15 maps. Full
monographic treatment, keys, excellent illus.
Jacobson, Morris K. and William K. Emerson. 1971. Wonders of
the World of Shells: Sea, Land and Fresh- water. 80 pp., illus.
Dodd, Mead and Co., N. Y. $3.95. Excellent little hardback for
young collectors.
Paget, O. E. 1971. List of European Malacologists. Gives addresses
and interests of 275 people. Obtained by sending to Dr. O. E.
Paget, Museum of Natural History, Burgring 7, Vienna A- 10 14,
Austria, 5 International Reply Coupons (purchased at your local
post office for 21 cents each).
Coan, Eugene V. 1971, July. The Northwest American Tellinidae.
Supplement to The Veliger, vol. 14, 63 pp., 12 pis., 29 text figs.
$5.75. Thorough and well-illustrated.
Shirai, Shohei. 1970. The Story of Pearls. 132 pp., 63 pis. (many
in color), 62 text photos. Beautifully illustrated account of Japa-
nese cultured pearls. $7.95. Japan Publ. Trading Co., San Fran-
cisco, 94103.
Russell, Henry D. 1971. Index Nudibranchia — a catalog of the
literature from 1554-1965. 141 pp., about 7,000 entries. $9.75.
Includes titles of papers, all taxa and subject cross-indices. Dela-
ware Museum of Natural History.
WILLIAM H. WEEKS SHELL COLLECTION: New price lists
of this famous collection, with full scientific data^ are in prepa-
ration. Many new additions of fine and rare species are also
included. To obtain free copies write:
George E. Jacobs, 853 Riverside Drive, New York, N. Y. 10032
INDEX NUDIBRANCHIA
A Catalog of the Literature from 1554 to 1965
by Henry D. Russell
7,000 entries: 2,400 titles, 3,400 taxa, 3,500 subject and
geographical cross-references are included in this new com-
prehensive index to 400 years of nudibranch literature. An
indispensable research source for malacologists and marine
biologists, assembled by a nudibranch specialist.
141 pages, 9^2 x I21/2 inches Plastic bound.. Price: $9.75
LIVING VOLUTES
A Monograph of the Recent Volutidae of the World
by C. S. Weaver and John E. du Pont
Hailed as the most complete, scientific and useful book on
the volutes, this work monographs 200 species and subspecies
with 79 colored plates, including color photographs of living
animals in their natural habitats. Distribution maps, anatom-
ical drawings and complete synonomies are given.
375 pages, 9 x 12 inches Text figures and maps
79 full-color plates Bound. Price: $55.00
Swainson's EXOTIC CONCHOLOGY
A Book Collector's Classic, now only $13.75 (formerly $30.00)
This sumptious and faithful facsimile of William Swainson's
1841 conchological work contains 48 beautiful, colored plates,
together with the original text and a modern analysis by R.
Tucker Abbott and Nora McMillan. Handsomely bound, gilt-
edged on 3 sides, and in large 9 x 12 inch page size.
All available from your favorite book or shell dealer, or
directly from the
Delaware Museum of Natural History
Box 3937, Greenville, Delaware 19807, U.S.A.
Vol. 85 JANUARY, 1972 No. 3
THE
NAUTILUS
THE PILSBRY QUARTERLY
DEVOTED TO THE INTERESTS OF CONCHOLOGISTS
EDITORS AND PUBLISHERS
R. Tucker Abbott, du Pont Chair of Malacology
Delawafe Museum of Natural History, Greenville, Del. 19807/Qq
Charles B. Wurtz, Consulting Biologist ' '
3220 Penn Street, Philadelphia, Pa. 19129
CONTENTS
Gastrocopta armifera (Say). By Leslie Hubricht 73
New gastropods from the Pleistocene of Illinois.
By A. Bryon Leonard 78
Observations on the gastropod, Charonia variegata, in Trinidad
and Tobago. By Peter L. Percharde 84
Chromosome number of the surf clam, Spisula solidissima.
By John W. Ropes 93
A new Fasciolaria from the northeastern Gulf of Mexico.
By William G. Lyons 96
Hemphillia dromedarius, a new arionid slug from Washington.
By Branley A. Branson 100
Winter distribution of Melampus bidentatus (Say) on a Cape
Cod salt marsh. By John W. Grandy IV 106
Notes 109
$5.00 per year ($5.75 to Foreign Countries) $1.50 a copy.
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JAN 3 1 1972
NAUTILUS:
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THE NAUTILUS
Vol. 85 January, 1972 No. 3
GASTROCOPTA ARMIFERA (SAY)
By Leslie Hubricht
4026— 35th Street
Meridian, Miss. 39301
Gastrocopta armifera (Say) which has been considered one var-
iable species, is in fact five distinct and easily recognized species,
once the distinguishing characters are understood. These species
can be readily divided into two groups by the size and shape of the
shell. The first group contains only G. armifera, and is distin-
guished by its larger, ovoid shell. The other group contains G.
abbreviata, G. similis, G. ruidosensis, and G. clappi. These four
species all have smaller, more cylindrical shells with more dome-
shaped spires and finer sculpture. All four species have been found
associated with G. armifera, and can be readily sorted by the differ-
ence in size and shape of the shell.
The graphs (figs. 1-4) show the relative height and diameter in
mm. of G. armifera and each of the four other species with which
it was found associated. Fig. 3 shows the size difference between
G. armifera and G. abbreviata where they were found living to-
gether, showing clearly that there are two species. While the dif-
ferences in the teeth would have made it possible to sort most of the
specimens, there were some in both species which were intermediate
in one or more characters. In the other species there are differences
in the teeth which are adequate to separate them from G. armifera
and from each other.
I am indebted to Dr. Juan J. Parodiz of the Carnegie Museum,
Pittsburgh, for the loan of material from the Sterki collection.
Key to the Species
1. Columellar lamella with a forward lobe 2
Columellar lamella without a forward lobe 3
2. Shell large, ovoid, columellar lamella usually with a distinct vertical lobe;
basal tooth usually small G. armifera
Shell smaller, more cylindrical, columellar lamella with vertical lobe weak
or wanting; basal tooth usually well developed G. abbreviata
3. Columellar lamella with a distinct angle near its center G. similis
Columellar lamella a flat projecting plate 4
4. Columellar lamella sloping backward G. ruidosensis
Columellar lamella sloping forward G. clappi
73
74 NAUTILUS Vol. 85 (3)
Gastrocopta armifera (Say)
Pupa armifera Say. 1821. Jour. Acad, Nat. Sci. Philadelphia,
2; 162.
Gastrocopta armifera (Say). Pilsbr)'. 1948. Land Mollusca of
North America (north of Mexico). Acad. Nat. Sci. Philadelphia,
Mono. 3, vol. 2, pp. 874-877, figs. 472: 1-4.
G. armifera varies in length from 3.7 to 5.0 mm., and in diameter
from 2.0 to 2.6 mm. The shell is usually ovoid, but may be cylin-
drical in elongate individuals. Such elongate shells are too large to
be confused with the other species of the complex. Angular and
parietal lamellae not always completely fused, as there is often a
shallow furrow separating them, and the angular lamella projects a
little beyond its junction with the parietal lamella. The columellar
lamella is shaped somewhat like an inverted 'Y', being branched
below. The lower palatal fold is rather thin, entering; the upper
palatal fold is similar but smaller. Supra-palatal tubercle small.
Basal fold usually very small.
G. armifera is found in calcarious areas over most of eastern
United States, ranging west to Colorado and New Mexico. It is
absent from the Piedmont and southern Atlantic Coastal Plain, and
from peninsular Florida. Its northern limits are not well understood
because of confusion with G. similis.
Gastrocopta abbreviata (Sterki)
Bifidaria armifera abbreviata Sterki. 1909. The Nautilus 23: 53.
Bifidaria armifera interpres Sterki. 1909. The Nautilus 23: 52.
Gastrocopta armifera abbreviata (Sterki). Pilsbry. 1948. Land
Mollusca of North America (north of Mexico). Acad. Nat. Sci.
Philadelphia, Mono. 3, vol. 2, p. 877, figs. 474: 1-3.
G. abbreviata varies in length from 3.3 to 4.2 mm., and in diam-
eter from 1.8 to 2.2 mm. The shell is more slender and generally
smaller than that of G. armifera. The angular and parietal lamellae
are more completely fused, the angular lamella does not project
beyond the junction with the parietal lamella. The vertical lobe of
the columellar lamella is quite short or wanting, the lamella being
shaped like an inverted 'U', or may be reduced to a horizontal fold.
The basal fold is well developed. G. abbreviata can not always be
distinguished from G. armifera by the teeth. The difference between
these two species is much like the difference between G. tappaniana
(C. B. Adams) and G. pentodon (Say), where differences in size
January, 1972 nautilus 75
and shape of the shell must be used to distinguish them.
G. abbreviata ranges from Illinois and Mississippi west to Col-
orado and New Mexico, and from North Dakota south to Texas. It
has been found associated with G. armifera at the following local-
ities: Missouri: St. Louis Co.: Prospect Hill. Mississippi: Oktibbaha
Co.: cedar glade, 1 mile southwest of Osborn (fig. 3). Oklahoma:
Haskell Co.: drift, South Canadian River, Whitefield. Texas:
Comel Co.: loess, near Guadalupe River, 1.5 miles north of Sattler
(fig. 4). Kerr Co.: loess just west of Ingram. The two lots labeled
interpres Sterki in the Sterki collection, Carnegie Museum, from
Fort Gibson, Oklahoma, and Wichita, Kansas, are mixtures of G.
abbreviata and G. armifera.
Gastrocopta siniilis (Sterki)
Bifidaria armifera similis Sterki. 1909. The Nautilus 23; 53.
Bifidaria armifera affinis Sterki. 1909. The Nautilus 23: 53.
Gastrocopta armifera (Say) form similis Sterki. Pilsbry. 1948.
Land Mollusca of North America (north of Mexico). Acad. Nat.
Sci. Philadelphia, Mono., 3, vol. 2, p. 877, fig. 472: 6.
Gastrocopta armifera (Say) form ajfinis Sterki. Pilsbry. ibid. p.
877, fig. 472: 5.
G. similis varies in length from 3.2 to 4.3 mm., and in diameter
from 1.7 to 2.0 mm. The shell is smaller and more slender than
that of G. armifera. The angular and parietal lamellae are com-
pletely fused. The columellar lamella lacks the forward lobe. The
vertical and backward lobes are higher and thinner than in G.
armifera. The lower palatal fold is usually very short, being tuber-
cular, often distinctly wider and more deeply immersed than in
G. armifera. The upper palatal fold is similar but smaller. The
basal fold is very small or wanting.
G. similis is a northern species, ranging from northern New York
and southern Canada, west to Minnesota and Kansas. It has been
found associated with G. armifera at the following localities: Mis-
souri: St. Charles Co.: loess, 2.5 miles northwest of St. Charles. St.
Louis Co.: talus. Cliff Cave; talus, Fox Creek Gap, 1 mile west of
Allenton; talus. Fern Glen (fig. 1). Illinois: Madison Co.: talus,
2 miles northwest of Alton.
Gastrocopta ruidosensis (Cockerell)
Bifidaria armifera var. ruidosensis Cockerell. 1899. The Nautilus
13: 36.
76 NAUTILUS Vol. 85 (3)
Columella tridentata Leonard. 1946. The Nautilus 60: 20, pi. 3,
figs. 1, 2.
Gastrocopta proarmifera Leonard. 1946. The Nautilus 60: 21,
pi. 3, figs. 3-5. Pilsbry. 1948. Land Mollusca of North America
(north of Mexico). Acad. Nat. Sci. Philadelphia, Mono. 3, vol. 2,
pp. 878-880, figs. 473: 3-4.
Gastrocopta tridentata (Leonard). Pilsbry. 1948. ibid. p. 880,
figs. 473: 1-2.
G. ruidosensis varies in length from 3.5 to 4.2 mm., and in diam-
eter from 1.8 to 2.0 mm. The shell is smaller and more slender
than that of G. armifera. The angular and parietal lamellae are
completely fused. The columellar lamella lacks the forward and
vertical lobes and the backward lobe is expanded into a broad plate.
The lower palatal fold is short and broad, the upper palatal fold is
similar but smaller. The basal fold is very small or wanting. The
columellar lamella and the lower palatal fold are deeply immersed.
G. ruidosensis is found living in a small area in central New
Mexico, but occurs as a Pleistocene fossil from western Kansas to
central Texas. It has been found associated with both G. armifera
and G. abbreviata at the following localities: Texas: Gomel Co.:
loess, near Guadalupe River, 1.5 miles north of Sattler (fig. 4).
Kerr Co.: loess, just west of Ingram.
On a wooded hillside, 7 miles east of Pryor, Mayes Co., Okla-
homa, associated with G. armifera, I found a single living shell that
agrees with the holotype of Columella tridentata Leonard in all
essential details. This is a rare aberration which might occur in any
species of the group.
Gastrocopta clappi (Sterki)
Bifidaria clappi Sterki. 1909. The Nautilus 22: 108, pi. 8, fig. 4.
Gastrocopta armifera clappi (Sterki). Pilsbry. 1948. Land Mol-
lusca of North America (north of Mexico). Acad. Nat. Sci. Phila-
delphia, Mono. 3, vol. 2, p. 878, figs. 472: 7-8, 472a.
Gastrocopta clappi (Sterki). Hubricht. 1962. Sterkiana 7: 1.
G. clappi varies in length from 3.5 to 4.3 mm., and in diameter
from 1.8 to 2.0 mm. The shell is smaller and more slender than
that of G. armifera. The angular and parietal lamellae are com-
pletely fused. The columellar lamella lacks the forward and back-
ward lobes. The vertical lobe is expanded into a thin, forward
sloping plate, and is placed low on the pillar. The lower palatal
January, 1972
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DIAIETER
Fig. 1, solid circles — G. similis, open circles — G. armifera, from Pleistocene
talus, Fern Glen, St. Lx)uis Co., Missouri. Fig. 2, solid circles = G. clappi, open
circles = G. armifera, from cedar woods, Tumhole Sink, Mammoth Cave Na-
tional Park, Edmonson Co., Kentucky. Fig. 3, solid circles = G. abbreviata,
open circles = G. armifera, from a cedar glade, 1 mile southwest of Osborn,
Oktibbaha Co., Mississippi. Fig. 4, solid circles = G. ruidosensis, large open
circles = G. abbreviata, small open circles = G. armifera, from Pleistocene
losss, near Guadalupe River, 1.5 miles north of Sattler, Gomel Co., Texas.
78 NAUTILUS Vol. 85 (3)
fold is short, the upper palatal fold is similar but smaller. The basal
fold is very small or wanting.
G. clappi ranges from southwestern Virginia to central Kentucky,
south to northern Alabama and Georgia. It has been found asso-
ciated with G. armifera at the following localities: Kentucky: Ed-
monson Co.: cedar woods, Turnhole Sink, Mammoth Cave National
Park (fig. 2). Tennessee: Moore Co.: cedar woods, 2.5 miles south-
west of Lynchburg. Georgia: Dade Co.: roadside, 3 miles southwest
of Wildwood.
NEW GASTROPODS FROM THE
PLEISTOCENE OF ILLINOIS
By a. Byron Leonard
The Illinois State Geological Survey
Urbana, Illinois 61801
In the course of studies of the molluscan faunas recovered from
sediments of Kansan age at eight localities and from more than
25 fossiliferous outcrops of Illinoian age in the Pleistocene rocks of
Illinois, six kinds of apparently undescribed gastropods were en-
countered. Formal descriptions of these mollusks follow.
Types and paratypes of these gastropods are catalogued in the
Paleontological Collections of the Illinois State Geological Survey,
except for suites of paratypes deposited with the Academy of Nat-
ural Sciences of Philadelphia.
Vertigo occulta, new species Fig. 8
Diagnosis: A small species of Vertigo, ovate-cylindrical in shape,
similar to Vertigo nylanderi and to V. briarensis from both of which
it differs in important details, principally with respect to the place-
ment of the lower palatal fold.
Type locality: Long Lake Section, Banner Formation, in humic-
stained sihs of Kansan age, situated in the Cen. NW14 NW'/4 sec.
18, T.2 N, R 2 E, Schuyler County, Illinois.
Description of holotype: An ovate-cylindrical shell of five obliquely
striate whorls; suture deeply impressed; aperture small, occupying
less than one-third the total length of shell; peristome slightly
everted and somewhat thickened, with furrow behind; outer lip of
peristome sharply indented at middle, forming clearly marked
sinulus; denticles six: a small, low, elongate angular, about one-
third the length of parietal, the latter elongate, heavy, deeply
January, 1972 nautilus 79
entering, and inclined above toward angular; small, nodular, col-
umellar lamella, not ascending inwardly; basal fold small, nodular,
subcolumellar in position; palatal folds elongate, elevated and thin,
the upper fold deeply entering from the angle of the sinulus, and
marked by a deeply impressed furrow without; lower palatal thin,
elevated, almost membranous, arising at columella and extending
almost at right angles to long axis of shell toward the inner end of
the upper palatal fold, where it bends slightly outward. This rela-
tion of the two palatals is evident also from their external impres-
sions. Total length of shell, 7.8 mm.; diameter, 1.0 mm.; diameter
of aperture, 0.6 mm., height, 0.5 mm.
Variations and comparisons: The large series of shells obtained
from seven Kansan and four early Illinoian localities vary little
except that the lower palatal fold is in some examples higher and
more membranous than in the holotype.
Vertigo occulta differs from V. nylanderi in that it is more robust;
the angular lamella is less deeply entering; the columellar lamella
does not ascend inwardly; and conspicuously by the placement of
the lower palatal fold, that is deeply situated within the aperture,
almost at right angles to the upper palatal. In V. briarensis the
angular lamella is wanting, the basal fold is extremely reduced or
wanting; and the lower palatal fold is more rugged, less deeply
immersed within the aperture, and subparallel to the upper palatal
fold.
Vertigo briarensis, new species Fig. 9
Diagnosis: Vertigo briarensis is small, the ovate-cylindrical shell
being always less than two millimeters in length, and having five
well-rounded whorls. It is similar to V. occulta, from which it
seems to have been derived.
Type locality: Briar Bluff South Section, in Petersburg Silts of
Illinoian age, in the SW corner NWI/4 sec. 21, T 1 N, R 1 E, Henry
County, Illinois.
Description of holotype: Shell ovate-cylindrical, having five ob-
liquely and finely striate whorls; suture deeply impressed; aperture
less than one- third total length of shell; peristome only slightly
everted, not thickened, and only vaguely indented, forming an
indistinctly outlined sinulus; denticles: an elongate elevated parietal
fold, deeply entering the aperture; a nodulose columella lamella;
and two palatal folds, deeply situated within the aperture, the lower
80 NAUTILUS Vol. 85 (3)
more so, and sub-parallel to the upper. Total length of shell, 1.75
mm.; diameter, 1.0 mm.; diameter of aperture, 0.5 mm., height of
aperture, 0.5 mm.
Variations and comparisons: Unlike V. occulta, V. briarensis has
not been recovered from Pleistocene deposits of Kansan age, but it
is widely distributed both geographically and stratigraphically in
deposits within the various substages of the Illinoian Stage. Shells
from eight localities are remarkably uniform, although there is
slight variation in size. None of the examples seems to intergrade
with V. occulta although they overlap stratigraphically, and V. bri-
arensis replaces V. occulta in deposits younger than early Illinoian,
Vertigo briarensis differs from V. occulta in that the angular
lamella is wanting and the basal almost so; the sinulus is poorly
defined; the palatal folds are both less developed and the lower
is subparallel to the upper, rather than being almost at right angles
to it.
Gastrocopta venusta, new species Fig. 7
Diagnosis: Shell small, less than two millimeters in length, having
five moderately inflated whorls, suture well -impressed; surface
sculpture granulate, except last two whorls both granulate and
obscurely striate; shell somewhat similar to Gastrocopta holzingeri
agna (Pilsbry and Vanatta) and G. ^alcis Leonard, but differing
from both in important details.
Type locality: Wildcat Creek Section, Petersburg Silt Formation of
Illinoian age, in die NEI/4 NEI/4 NEI/4 sec. 19, T 13 N, R 4 W,
Mercer County, Illinois.
Description of holotype: Shell small for the genus, having five
moderately inflated whorls; structure deeply impressed; first three
whorls having granulate sculpture; the last two whorls having
granulate surface overlain with faint striae; aperture small, nar-
rowly ovate, less than one third height of shell; peristome thin and
sharp, but with heavy crest behind; denticles 7: A fused angulo-
parietal as in the immersidens group, the two forked in front; the
angular the more conspicuous in front, extending to the peristome,
the fused denticle is high, strong, sinuous and deeply entering; the
columellar lamella arises deeply within the aperture on the colu-
mella, rapidly increases in height, and extending almost to the
parietal before decreasing in height and extending to the outer edge
of the peristome; the basal fold is broad in front and situated on the
January, 1972 nautilus 81
callus that bears the palatal folds, it narrows and increases in height
inwardly; the lower palatal fold is broad where it arises from the
callus but it narrows inwardly; the upper palatal fold is much
smaller, narrower and shorter than the lower; a small granular
suprapalatal fold is invariably present. Total length of shell, 1.75
mm.; diameter, 0.95 mm.; diameter of aperture, 0.55 mm.; height of
aperture, 0.5 mm.
Variations and comparisons: Gastrocopta venusta was recovered
from three localities, one in the Kansan and two Illinoian deposits;
in all the three localities yielded only a dozen specimens, among
which the only observed variation was a few tenths of millimeters
in length. G. venusta differs from G. holzingeri agna in that: the
angular lamella is more conspicuous; the columellar lamella extends
to the outer edge of the peristome; the basal and palatal folds are
broad below and narrow within; and the aperture is narrower and
more triangular.
In G. falcis the angulo-parietal fold is broadly forked in front,
the columellar lamella does not extend to the outer lip of the peri-
stome, and the suprapalatal fold is high and entering rather than
tubercular.
Ariniger exigua, new species Figs. 2, 3
Diagnosis: A minute species of Armiger, having two and one-half
to three depressed, carinate whorls, that slowly increase in diameter
toward the aperture. The nuclear whorl has finely granular sculp-
ture, while the remaining whorls exhibit irregular, subvertical
striations and less numerous carinae. There are no spiral striations.
The last whorl docs not descend toward the aperture, which is
roundly ovate in form.
Type locality: Briar Bluff South Section in Petersburg Silts of
early Illinoian age, in the SW corner of NW|4 sec. 21, T 17 N,
R 1 E, Henry County, Illinois.
Description of holotype: Shell piano-spiral with two and one-half
whorls that are flattened somewhat above, rounded below; the
periphery is somewhat above the middle of the last whorl which is
roundly angular; last whorl not descending toward the aperture;
aperture rounded-oval, and narrowly appressed to preceding whorl;
spire depressed but not below level of general dorsal surface; umbil-
icus impressed but revealing all the whorls; nuclear whorl having
granular sculpture that extends on to succeeding whorls where they
82 NAUTILUS Vol. 85 (3)
are overlain by coarse diagonal growth lines which on the last whorl
are interspersed with costae, of which there are about 18 on the
last whorl; there is no hint of spiral striae. Lip of peristome thin
and simple. Greater diameter of shell, 1.95 mm.; lesser diameter,
1.67 mm.; height of aperture, 0.7 mm.; width of aperture, 0.6 mm.
Variations and comparisons: Shells vary but little among the 53
paratypes available. The largest shell observed had two and three-
fourths whorls, and its greater diameter measured 2.2 mm.
Armiger exigua differs from A. crista (Linne) in that it is much
smaller, although having the same number of whorls; the nuclear
whorl is granulate rather than striate; there is no indication of spiral
striae; the last whorl does not descend toward the aperture; and the
aperture is roundly oval in shape.
Succinea exile, new species Fig. 1
Diagnosis: Shell similar in form to Succinea gelida Baker but uni-
formly smaller (total length always less than 5 mm.) more slender
in form, and having the aperture narrowly ovate above.
Type locality: Briar Bluff South Section, Petersburg Silts of lUi-
noian age, in the SW corner NWV4 sec. 21, T 1 N, R 1 E, Henry
County, Illinois.
Description of holotype: Shell small for the genus; the three and
one-half whorls rounded, the last proportionately small, separated
by deeply incised suture; nuclear whorl swollen but smaller (diam-
eter 0.5 mm) with granular sculpture, the remaining whorls having
delicate but rather irregular vertical striae; aperture pear-shaped,
rounded below, narrowed above; outer lip of peristome not greatly
curved inward above; columellar lip rounded below, angled above
toward the short callus spread against the parietal wall; aperture
length about one half that of shell. Total length of shell, 4.9 mm.;
diameter, 2.4 mm.; height of aperture, 2.4 mm., diameter, 1.7 mm.
Variations and comparisons: Among the 42 shells from three
Kansan and 4 Illinoian localities there is little variation except that
due to difference in age. Succinea exile differs from S. gelida which
it resembles in general form, in being uniformly smaller, and the
shell having a more slender form; the aperture is narrower above,
and the outer lip does not turn inward above as sharply as it does
in gelida.
January, 1972
NAUTILUS
83
Fig. 1. Succinea exile Leonard, holotype; X 6.5. Figs. 2, 3. Armiger exigua
Leonard, holotype; X 12.5. Spiral and umbilical views, respectively. Figs. 4, 5,
6. Punctum parvulum Leonard, holotype; X 18.8. Spiral, apertural and umbilical
views, respectively. Fig. 7. Castrocopta venusta Leonard, holotype; X 18. Fig. 8.
Vertigo occulta Leonard, holotype; X 19. Fig. 9. Vertigo briarensis Leonard,
holotype; X 19.
84 NAUTILUS Vol. 85 (3)
Punctuni parvulum, new species Figs. 4, 5, 6
Diagnosis: A minute, depressed, conoid, spiral shell, similar to
Punctum minutissimum (Lea) which it most resembles, but differ-
ing in being smaller in size and having a lesser number of whorls,
lacking spiral striae, and having a much narrower umbilicus.
Type locality: Long Lake Section, in silts of the Banner Forma-
tion of Kansan age, situated in the center of NW'/4 NWJ/4 sec 18,
T 2 N, R 2 E, Schuyler County, Illinois.
Description of holotype: Shell small, depressed, conoid, whorls
three and one-half, rounded; suture moderately impressed; aperture
lunate, peristome thin, simple; nuclear one and one-third whorls
granular, the granular sculpture extending onto the remaining
whorls where it is overlain by extremely delicate vertical striae, not
apparent without strong magnification; no indication of spiral lines
either above or below; umbilicus very narrow, its diameter no more
than one-sixth that of the shell. Diameter of shell, 1.3 mm.; height,
0.7 mm.; height of aperture, 0.5 mm., diameter, 0.5 mm.; diameter
of umbilicus, 0.22 mm.
Variations and comparisons: The 95 shells from seven Kansan
and two Illinoian localities are remarkably uniform except for age
variations. P. parvulum differs from P. minutissimum in that it is
slightly smaller and with fewer whorls; the nucleus is granular
rather than smooth; the striae are more delicate; spiral striae are
lacking, and the umbilicus is much narrower.
OBSERVATIONS ON THE GASTROPOD,
CHARONIA VARIEGATA, IN TRINIDAD AND TOBAGO
By Peter L. Percharde
Trinmar Limited, Point Fortin,
i Trinidad and Tobago
When the author came to Trinidad in 1958, underwater observa-
tions soon revealed that Charonia variegata was fairly common in
these waters. Many specimens have since been observed underwater
in their habitat, and observation on their feeding habits reveals that
they are voracious predators.
Forms and Habitat
Like the shells of many other species of tropical gastropods,
Charonia variegata presents two separate morphological forms. The
shallow water, coral reef form is characterized by the denser calcar-
January, 1972
NAUTILUS
85
eous shell, the lighter colouration of the shell pattern and the
tendency for the shell to be overgrown with algae, and the early
whorls to be badly pitted by the boring sponge Cliona sp. The deep
water form, which is found in 15 to 45 metres of water, lives on a
grit and muddy silt bottom with occasional rock outcrops, under the
eaves of which the mollusc finds shelter while resting. G. Warmke
and R. Tucker Abbott reported in "Caribbean Seashells," that divers
in Puerto Rico found Charonia variegata in underwater caves.
Fig. lA. Specimen of Charonia variegata whicii was observed on many occa-
sions feeding on echinoids, mainly Eucidaris tribuloides. Fig. IB. Specimen
tagged and observed over an extended period, feeding mainly on asteroids,
Echinaster sentus, and bivalve molluscs, Laevicardium lacvigalum and Chione
cancellata.
86
NAUTILUS
Vol. 85 (3)
Not one specimen has ever been found in a true marine cave in
Trinidad. However, most of the specimens observed have been
found under the eaves of large rocks or coral reefs, or out on the
open mud substrate at the foot of a reef or rocky outcrop. While
searching for a place to rest, even large specimens of Charonia
variegata manage to creep under rocks and wedge themselves into
the silt substrate.
The shallow water form appears to be fairly dormant during the
Fig. 2A. Specimen of Eucidaris tribuloides taken away from the light-coloured
Charonia in lig. lA. Fig. 2B. Specimens upon which the dark-coloured Charonia
was feeding {Laevicardium laevigatum. Echinastcr sentus and Chione cancellata) .
January, 1972
NAUTILUS
87
daytime, especially if there is strong sunlight and clear water. The
deeper water form is characterized by the much lighter construction
of the shell, strongly reminiscent of the thin but strong shells of
Tonna galea Linne 1758. It is significant that both these two
species belong to the same super family, the Tonnacea and possess
in some ways similar feeding habits, especially in the use of highly
acid saliva jetted at their prey in order to subdue same. The shell
pattern of the deep water form is also more colourful, being more
vivid yellow and orange. (See Feeding Habits).
The protoconch and the early whorls, of the deep water form are
more likely to be found in good condition and without pitting and
shell damage by fouling growth, etc. The deep form, quite often
avails itself of shelter in discarded man-made objects whilst extend-
ing the shell, between varices. The author has observed specimens
in the process of shell extension, sheltering in an old bucket, truck
tyres, paint or grease drums and sections of steel pipe.
Fig. 3. Map of locality records of Charonia variegata found by the author.
88 NAUTILUS Vol. 85 (3)
The shallow water form has not been observed in these strange
habitats, although in certain reef areas these objects are regrettably
very common. In shallow water, Charonia variegata seeks shelter
far inside holes in the coral and in rock reefs, specimens are often
found, wedged tightly in the crevices below large fallen rocks. The
finding and observation of these specimens often requires the use
of a powerful underwater light even in daylight hours.
Feeding Habits
After over fifty observations of feeding behaviour, it is considered
that Charonia variegata is one of the most voracious of gastropod
predators here in the waters of Trinidad and Tobago. The food
consists of a wide range of invertebrates, including the echinoderms,
Echinaster sentus, Eucidaris tribuloides, Tripneustes sp., Synapta
sp., Cucumaria sp.; the molluscs, Laevicardium laevigatum, Chione
cancellata, Fasciolaria tulipa, Cypraea zebra; and the crustaceans,
Panulirus argus and Panulirus guttatus.
The method of attacking the prey, employed by specimens ob-
served, was fairly constant. A creeping approach, with tentacles
being swept from side to side. This is followed by contact with the
prey by the tentacles and the extension of the large proboscis ready
to eject the acid saliva over the prey. This action rapidly causes
paralysis of the intended prey, followed by the subsequent wrapping
of the foot of Charonia variegata around the stunned animal.
The pleurembolic proboscis of Charonia variegata contains, in the
odontophore, a radula armed with strong rasping teeth. Two jaws
are also present, mounted laterally. These are thin chitinous plates
and are not very strong but possibly are used in die process of
opening bivalve molluscs, after the valves have started to gape,
subsequent to being jetted with acid saliva.
However, Charonia variegata is capable of piercing the tests of
echinoids and the author has observed specimens of Eucidaris trib-
uloides being attacked and wrenched out of their habitat in reef
crevices. The test is usually pierced either in the area of the madre-
pore plate or alongside the five-toothed mouth.
While engaged in feeding, the triton holds its prey firmly by the
foot and completely envelopes it in thick mucus. Once the test is
pierced, the extended proboscis enters the rasped hole and the entire
contents are devoured, leaving only the five-toothed mouth struc-
ture, or "Aristotle's lantern," which is often detached from the test.
January, 1972 nautilus 89
In Trinidad, during February, large numbers of the asteroid
Echinaster sentus congregate in sheltered areas of bays in these
waters. This behaviour is to assist in mating, the sexes being sep-
arate. While this mass grouping is taking place they are often
attacked by Charonia variegata.
The author has observed pairs of Charonia variegata methodically
driving dozens of specimens of this common orange-red asteroid up
a silty mud slope, attacking the outside members and instead of
completely devouring them as they would do under normal circum-
stances, just killing them and rasping a hole out of the central part
of their bodies. Specimens of Charonia variegata examined after
being engaged in one of these feeding "orgies," have their intestines
stuflFed full of digested asteroids.
The intestines and digestive glands are stained orange-red, and
a gritty mass in the intestines is composed of the calcareous particles
of decomposed dermal skeletons. Dietary preference may also play
a part in determining shell colour and patterns, as the author has
observed that specimens which favour a diet of the orange-red
Echinaster sentus are usually the specimens with the most pro^
nounced shell colouration and pattern, while specimens which
favour echinoids, are a pale golden colour with reduced shell pat-
tern (See plate).
Other gastropods are paralyzed, enveloped and devoured. The
powerful proboscis with radular teeth and acid saliva, soon removes
the entire animal from the shell.
The method of opening bivalves is similar. The triton emits a
quick jet of acid saliva, followed by the rapid extension of the
proboscis between the partially gaping valves. No observations have
been made on Charonia variegata eating buried bivalves. They eat
specimens caught on the surface of the substrate. The gasropod
FascioLaria tulipa often shares the same resting habitat with Char-
onia variegata, but Fasciolaria tulipa remains buried 5 to 10 centi-
metres below the resting place of Charonia variegata. Juvenile
specimens have been observed being eaten by Charonia variegata.
On two occasions juvenile spiney lobsters, Panulirus argus and
Panulirus guttatus have been observed being eaten by Charonia
variegata. The method of approach and attack, was not observed but
in the case of the juvenile Panulirus argus the animal was still mov-
ing, after the large mollusc rasped a hole through the skin at the
90 NAUTILUS Vol. 85 (3)
junction of the abdomen and carapace on the ventral surface. This
is a position usually favoured by Octopus vulgaris vi^hen feeding on
similar prey. Ten hours later, when re-observed, the abdomen and
carapace were found empty.
General Behaviour and Enemies
Initially, while attempting to determine movement patterns for
Charonia variegata, the author had been hampered by the fact that
while this species is not rare, it is very rarely found in any number
exceeding two, in any one area. Consequently, when six large
specimens were observed in various habitats off the south coast of
Gasper Grande Island, they were tagged, in the hope that they
could be observed over an extended period.
Previous experiments with tagging this species in a bay on the
south coast of Monos Island, had resulted in the the use of the con-
ventional plastic numbered tag carefully attached to the spire of
the shell with monel or stainless steel wire. The most useful devel-
opment however was the use of a small plastic marker float, approx-
imately five centimetres in diameter attached to the shell by a thin
nylon cord, one half metre in length.
The use of this type of marker enabled the author to relocate
specimens repeatedly for many months of observation. The tagging
experiment off the south coast of Gasper Grande was quite success-
ful and the work revealed that the deep water form moved con-
siderable distances in one week periods. One specimen moved down
the slope from 14.5 metres to 30.5 metres and laterally 200 metres,
in a seven day period.
The deep water forms moved more than the shallow water forms,
and while it was found that the deep water forms sometimes moved
up into shallow water, the shallow water forms were never observed
to venture below eleven metres. The shallow water forms were
found to move and feed mainly at night time, while the deep water
forms, also mainly nocturnal, were often observed moving and
feeding by day.
Eventually, after about two years, all the tagged specimens were
lost and could not be relocated but during the course of a deep
underwater search carried out one Sunday, the author was swim-
ming horizontally at a depth of thirty metres when one of the
largest of the tagged specimens was relocated in a coarse sand gully
between two large rock ridges. The animal was moribund and the
January, 1972 nautilus 91
operculum had fallen out of the aperture onto the sand. Part of the
dead mollusc still remained in the shell and by the stage of decom-
position, it had been dead for about a week. The cause of death
was not known but what surprised the author was the fact that in
this same gully, several other empty shells of Charonia variegata
were found; all large, fully adult, if not gerontic specimens. There
were no shells of any other species of molluscs in the area and a
careful search failed to reveal any signs of Octopus sp. or other
predators. Some of them had been laying in this location for many
years. There were no signs of damage to any of the shells and two
of them had opercula lying close to the shell on the sand substrate.
Two observations with regard to the enemies of Charonia vari-
egata have been made by the author. In the first instance, a
medium-sized specimen was picked up and eaten by a large ray,
Aetobatus narinari. The shell was crushed and dropped in small
pieces as the ray rose up from the bottom and swam away. In the
second instance a large Loggerhead turtle, Caretta caretta, was
observed crushing and eating a large specimen of the deep water
type of Charonia variegata. Local fishermen have also informed
the author that the Hawksbill turtle, Eretmochelys imbricata, often
eats "trumpet conks."
Reproduction and Dispersal of Larvae
In Trinidad and Tobago, Charonia variegata lays eggs in May
just before the onset of the rainy season. The egg cases are usually
laid at night time. The cases are clustered together in a large mass
and coloured a pale parchment yellow. Egg cases placed in shallow
water are hidden underneath fallen rocks or coral, but egg cases
placed in deep water are quite often laid out in the open.
Once again, man-made objects attract females about to lay in
deep water. As the author has observed, Charonia variegata egg
cases alongside old rubber tyres on several occasions. Only one
observation of copulation in this species has been made. Two speci-
mens were observed under fallen rocks on May 5, 1967. The male
was only half the size of the female. The female was a fairly old
specimen with a badly eroded shell spire. The penis of the male, is
capable of extension to reach into the mantle cavity of the female;
however, the degree of extensibility does not compare with other
large gastropods of these waters, for example — Strombus gigas and
S trombus costatus.
92 NAUTILUS Vol. 85 (3)
Conclusion
After several years of extensive, detailed observation, we found
that Charonia variegata is an extremely active and voracious preda-
tor, feeding on a v^ide variety of invertebrate prey.
This large mollusc is aided by its highly acid saliva vi'hich is used
most effectively in paralyzing the prey during an encounter. This
mollusc must play an important role in the ecological balance of the
extensive areas of its habitat.
It is also fairly well established that this species is found in two
separate forms. A shallow water, rock or coral reef form and a
deeper water form. The morphological difference with regard to
the shell of these two forms are quite easily discernable even to an
observer underwater.
The tagging of specimens for observation has been quite success-
ful, and especially the use of a small plastic float marker attached
to specimens by a half-metre nylon cord. This has enabled the
author to relocate specimens over considerable periods of time.
Acknowledgments
The author would like to express sincere thanks to John B.
Saunders and Dr. Hans G. Kugler for their constant encourage-
ment. To Errol G. Law for his help underwater and many hours
of diving together to collect the information for this work. To Dr.
Ailsa Clark of the B.M. (N.H.) for her kind assistance in identify-
ing the echinoderms. Also to Capt. R. Navarro for his great help
with marine transport.
Literature References
Clench, W. J. and Ruth D. Turner. 1957. The family Cymatiidae
in the Western Atlantic. Johnsonia 3; 189-244.
Hirsh, G. 1915. Ernahrungsbiologie fleischfressender Gastropoden.
Zool. Jahrb. Abt. Allg. Zool. Physiol. 35. 1917. Der Kalk. Zool.
Jahrb. Abt. Allg. Zool. Physiol. 36.
Houbrick, Joseph R. and Vera Fretter. 1969. Some aspects of the
Functional Anatomy and Biology of Cymatium and Bursa. Proc.
Malac. Soc. Lond. 38: 415-429.
Lebour, Marie V. 1945. The Eggs and Larvae of some Proso-
branchs from Bermuda. Proceedings Zoological Society London
7/4; 462-489.
Panceri, P. 1868. Nouvelles observations sur la salive des mol-
lusques gasteropodes. Annls. Sci. Nat. (Zool.) 10: 39-100.
Schroeder, R. E. 1962. Urchin Killer. Sea Frontiers 8: 156.
Sugar, J. A. 1970. Starfish threaten Pacific reefs. National Geo-
graphic /37(3): 340-353.
Warmke, G. and R. T. Abbott. 1968. Caribbean Seashells. Liv-
ingston Press, Wynnewood, Pa.
January, 1972 nautilus 93
CHROMOSOME NUMBER OF THE SURF CLAM,
SPISULA SOLIDISSIMA
By John W. Ropes
National Marine Fisheries Service
Biological Laboratory, Oxford, Maryland 21654
Ripe eggs from the surf clam, Spisula solidissima (Dillwyn, 1817),
an important commercial bivalve of the western North Atlantic
coast (Ropes, 1967), were collected and prepared for chromosome
counts. An examination of chromosome numbers was undertaken
to learn more of the clam's biology after a study of its reproductive
cycle (Ropes, 1968). Recent examinations of pelecypods have
shown an agreement in chromosome numbers for species within a
genus or family (Rosenfield, 1963; Menzcl and Menzel, 1965;
Menzel, 1968; Longwell and Stiles, 1968), and my results are iden-
tical to reported counts for other species in the family Mactridae.
Surf clams were obtained from commercial vessels at Ocean City,
Md., during July and August 1969. The clams were brought to the
laboratory, held overnight in a refrigerator at 7°C, and on the next
day each clam was placed in a basin of standing sea water at room
temperature (20°C). This simple thermal stimulation induced
spawning. Ova were pipetted into beakers of sea water and a dilute
sperm suspension was added. A microscopic examination at low
power to observe chromosomal activities preceded transferring the
cells onto slides, flooding them with a 1% orcein in 45% acetic acid
solution, and adding a coverslip. The slides were warmed by passing
them over the flame of an alcohol lamp to intensify the reaction of
the stain with the chromosomes. After a 30-minute staining period,
the slides were pressed between sheets of bibulous paper to squash
the eggs and spread the chromosomes. A ring of adhesive painted
on the edges of the coverslip and storage of the slides in a refrig-
erator extended the use of the preparations for several days. Egg
examinations and photographs were made with a Zeiss Photomicro-
scope and lOOX Planapochromat objective.
The unfertilized eggs of S. solidissima, and most other mollusks
(Raven, 1958), are normally blocked from completing maturation
at metaphase of the first meiotic division; after sperm penetration,
maturation continues (Allen, 1953). Thus, fertilized eggs were used
to obtain reduction division. Fertilized eggs held at room tempera-
ture for a 30-minute period before staining contained the lowest
94 NAUTILUS Vol. 85 (3)
and most constant number of chromosomes. The N or haploid
chromosome number of 18 was observed for S. solidissima at meta-
phase of meiosis (Fig. lA). Eggs found at anaphase of meiosis
revealed that the2N or diploid chromosome number is 36 (Fig. IB).
The chromosome numbers for S. solidissima given above are com-
parable with those found for other species in the family Mactridae.
Kostanecki (1904) reported N= 18 and 2N = 36 for Mactra sp. col-
lected from the Mediterranean near Naples and Trieste. The species
used by Kostanecki were, according to Allen (1953), M. stultorum
and M. helvacea. Menzel (1968) found the same haploid and
diploid numbers (18 and 36) for the channeled duck clam, Labiosa
plicatella (Lamarck), from local Florida waters and for the dwarf
surf clam, Mulinia lateralis (Say), from Virginia.
The family Mactridae is widespread in the world. As many as
50 nomenclatorially valid generic names have been listed for the
family (Vokes, 1967), which includes over 200 species (Ropes,
Chamberlin and Merrill, 1969). Agreement in the number of
chromosomes of the four species reported here is too little evidence
to conclude that they are representative of the family. Examina-
tions of the chromosome numbers of other Mactrid species might
add more evidence of agreement within the family.
Literature Cited
Allen, R. D. 1953. Fertilization and artificial activation in the egg
of the surf-clam, Spisula solidissima. Biol. Bull. /05(2): 213-239,
Kostanecki, K. 1904. Cytologische Studien an kiinstlich partheno-
genetisch sich entwickelnden Eiern von Mactra. Arch. Mikroskop.
Anat. 64: 1-98.
Longwell, A. C. and S. S. Stiles. 1968. Fertilization and completion
of meiosis in spawned eggs of the American oyster, Crassostrea
virginica Gmelin. Caryologia 2/(1): 65-73
Menzel, R. W. and M. Y. Menzel. 1965. Chromosomes of the
species of quahog clams and their hybrids. Biol. Bull. i29(l):
181-188.
Menzel, R. W. 1968. Chromosome number in nine families of
marine pelecypod mollusks. The Nautilus 82(2) : 45-58.
Raven, C. P. 1958. Morphogenesis: The Analysis of Molluscan
Development. Pergamon Press, London, 311 p.
Ropes, J. W. 1967. Surf clamming — a growing fishery. Fish. News
Int. 6(8): 58-60.
Ropes, J. W. 1968. Reproductive cycle of the surf clam, Spisula
solidissima, in offshore New Jersey. Biol. Bull. 735(2): 349-365.
Ropes, J. W., J. L. Chamberlin and A. S. Merrill. 1969. Surf clam
fishery, p. 119-125. In: Firth, F. E. (ed.) The Encyclopedia of
January, 1972
NAUTILUS
95
Marine Resources. Van Nostrand Reinhold Co., N. Y.
Rosenfield, A. 1963. Some cytological and chemical characteristics
of the Ostreidae. Nat. Shellfish. Ass. 55th Annu. Conv., Summ.
Tech. Papers, p. 25-26.
Yokes, H. E. 1967. Genera of the Bivalvia: a systematic and biblio-
graphic catalogue. Bull. Amer. Paleont. 5/(232): 111-394.
'^^:*-y' '■'»■'' -A-^-
4l * * «*'!?/-»
% . " .4' v£> w^ j^'^x ■ V-.i
^
1
Fig. 1. Chromosomes of S. soUdissima. A, haploid number at metaphase;
B, diploid number at anaphase.
96 NAUTILUS Vol. 85 (3)
A NEW FASCIOLARIA FROM THE
NORTHEASTERN GULF OF MEXICO
By William G. Lyons
Florida Department of Natural Resources
Marine Research Laboratory*
St. Petersburg, Florida 33731
From August 1965 through November 1967 the Florida Depart-
ment of Natural Resources Marine Laboratory conducted a sys-
tematic sampling program, Project Hourglass, on the West Florida
Shelf (Lyons, 1968; Joyce & Williams, 1969). Collections of benthic
animals were taken along two east-west transects off Egmont Key
near St. Petersburg and off Sanibel Island. Stations on each tran-
sect ranged in depth from 6 to 73 m. Two specimens of Fasciolaria
collected at Hourglass station E (73 m) represent an undescribed
species. Searches of the mollusk collections of the U.S. Museum of
Natural History, Washington, D. C, the Museum of Comparative
Zoology, Cambridge, Massachusetts, and the Academy of Natural
Sciences of Philadelphia, Pennsylvania, failed to uncover any other
material. Two additional specimens were provided by Mr. Harvey
R. Bullis, Jr. from collections made off the Florida west coast by the
U.S. Fish and Wildlife Service exploratory fishing vessel Oregon.
Fasciolaria bullisi, new species Fig. 1
Description: Shell thin, fusiform, slender; length/width ratio of
the 3 unbroken specimens increasing with shell length from 2.62 to
2.87; largest specimen with nearly 8 whorls including nucleus;
background color pale yellow with large patches of deeper yellow
and orange. Embryonic whorls 1%, first smooth except for two faint
spiral threads on lower end (on unworn specimen), final ^/^ whorl
with 16-18 moderately strong axial riblets. First post-embryonic
whorl with five incised, equidistant spiral lines, replaced by five thin
brown bands on later whorls. Large specimens with 10-12 primary
brown bands and 6-9 secondary bands on body whorl. Aperture
ovo-elongate; outer lip thin, simple, finely lirate within. Columella
straight to arcuate, with two shallow, oblique anterior plicae;
siphonal canal long, slender, oblique, of a rich amber, deepening to
brown at tip. Operculum thick, corneous, ovo-elongate, attenuated
obliquely at anterior end. Periostracum on dried specimens tan,
very thin.
* Contribution No. 182
January, 1972
NAUTILUS
97
Figure 1. Fasciolaria bullisi Lyons, new species. Holotype, off Egmont Key,
Florida, 73 m. Length: 134.1 mm.
Type depositories: The holotype and one paratype are in the col-
lection of the U.S. Museum of Natural History, Smithsonian Insti-
tution (USNM), Washington, D. C. Additional single paratypes
are in the collections of the Museum of Comparative Zoology
(MCZ), Harvard University, and the Florida Department of Nat-
ural Resources Marine Research Laboratory (FSBC), St. Peters-
burg, Florida.
Material examined: Holotype: USNM 706880. Length 134.1 mm,
width 46.8 mm (living). Hourglass station E, 27°37'N, 84°13'W,
73m; bottom temperature 19.0°C; June 7, 1966. Paratypes: USNM
706881. Length 117.0 mm, width 43.0 mm (living). Oregon station
1024, northwest of Dry Tortugas, 25°13'N, 83°55'W, 119 m; bot-
tom temperature 20.0°C; April 19, 1954.— FSBC I 7294. Length
59.3 mm, width 22.6 mm (dead shell). Hourglass station E; bottorn
temperature 19.5°C; May 12, 1967.— MCZ 261430. Length 119.0
mm, width 49.1 mm (dead shell, siphon broken). Oregon station
98 NAUTILUS Vol. 85 (3)
1254, south of Pensacola, 29°43'N, 87°18'W, 164 m; bottom tem-
perature 17.2°C; March 1, 1955.
Remarks: The species is named for Harvey R. Bullis, Jr., Asso-
ciate Director, U.S. Bureau of Commercial Fisheries, Washington,
D. C, who graciously loaned his specimens for study.
The dead specimen with damaged siphon from oflf Pensacola
(MCZ 261430) was apparently more globose than unbroken speci-
mens. Although the spire is 4 mm shorter than that of the holotype,
the width of the body whorl is 2.3 mm greater, suggesting a lower
length/width ratio. Color, banding, nuclear sculpture, and absence
of a presutural ridge, however, indicate that it should be assigned
to Fasciolaria bullisi sp. nov.
It is difficult to place Fasciolaria bullisi within the currently
accepted systematic scheme. Hollister (1957) erected the subgenus
Cinctura to contain Fasciolaria lilium Fischer von Waldheim, F.
lilium tortugana Hollister, F. hunteria (Perry), F. branhamae
Rehder and Abbott, and F. apicina Dall. This group was distin-
guished from Fasciolaria s.s. (type species F. tulipa Linne) by a
prominent spiral ridge extending onto the parietal wall from within
the posterior portion of the aperture. The validity of Cinctura is
questionable; specimens of F. tulipa with a swollen or raised "hump"
on this area of the shell are common, though I have seen none as
strongly expressed (ridge-like) as those borne by species of Cinc-
tura. Fasciolaria bullisi lacks any indication of this ridge.
Primary and secondary banding of later whorls of F. bullisi is
weak, often interrupted like that of F. tulipa, but unlike the strongly
expressed bands of Cinctura species. Distinct axial riblets on the
last embryonic whorl readily separate it from F. hunteria, which
lacks such sculpture. However, F. bullisi has the longer spire and
more slender outline of certain Cinctura and, like the latter, lacks
the incised spiral lines found on later whorls of F. tulipa. Also, like
Cinctura, F. bullisi bears only traces of the rough, presutural wrin-
kles of F. tulipa.
The radula of the holotype of F. bullisi is similar to that of F.
hunteria, both in the subtriangular cusps of the median tooth and
in the number and shape of cusps of the lateral teeth. Cusps of the
median tooth of F. tulipa are longer and more slender; those of the
laterals are much more numerous and are also more slender. Rad-
ulae of F. lilium and F. branhamae have not been described.
January, 1972 nautilus 99
It is possible that F. bullisi is the eastern Gulf analogue of F.
branhamae, which occurs from Texas to Campeche in moderate
depths. Fasciolaria branhamae has an axially sculptured nucleus
and elongate siphon like F. bullisi, but differs in color, strength and
number of bands, possession of a strongly-defined presutural ridge,
and in having a more globose body whorl. It occurs sympatrically
with F. lilium, the western Gulf analogue of F. hunteria. Certain
similarities of sculpture, banding and outline between specimens
from separate localities suggest that F. branhamae may be merely
a subspecies of F. lilium. It was described as a subspecies of F.
distans Lamarck (= lilium Fischer von Waldheim) by Rehder and
Abbott (1951), but Hollister (1957) elevated it to specific rank. No
such similarities have been noted between F. bullisi and any other
western Atlantic Fasciolaria.
The possibility that F. bullisi represents a hybrid of F. hunteria
and F. tulipa, the two other eastern Gulf species, seems untenable.
No other specimens of F. bullisi are known from North Carolina to
the Mississippi Delta, the region where F. hunteria and F. tulipa
occur sympatrically. The deepest record I have seen for F. tulipa is
73m (Work, 1969:674) from off Suriname. In Hourglass collec-
tions, F. tulipa decreased in numbers markedly with increased
depth; only two specimens were collected in 73 m from 110 tows at
this depth. Fasciolaria hunteria showed a similar decrease in abun-
dance, but was still fairly common at 73 m stations. However, spec-
imens of the latter from 73 m had remarkably heavy shells with
very short siphons quite unlike those of F. bullisi. The deepest-col-
lected F. hunteria I have seen is a juvenile from 77 m off northwest
Florida. Specimens of F. bullisi seem to occur only at the extreme
edge or beyond the bathymetric range of their hypothetical parents.
It appears best at present to treat F. bullisi as a species distinct from
other western Atlantic Fasciolaria.
Acknowledgments
I thank Mr. Bullis for the loan of his specimens, and the many
workers at the above museums for their assistance. Dr. R. T. Ab-
bott, R. C. Bullock, H. R. Bullis, and D. K. Serafy critically read
an early draft of the manuscript. Most of their comments were
incorporated into the final copy. Mrs. S. D. Kaicher photographed
the type.
100 NAUTILUS Vol. 85 (3)
References Cited
Hollister, S. C. 1957. On the status of Fasciolaria distans Lamarck.
Nautilus 70(3): 73-84; 1 pi.
Joyce, E. A., Jr. and J. Williams. 1969. Rationale and pertinent
data. Memoirs of the Hourglass Cruises, Vol. I, Part I. Fla.
Dept. Nat. Resources Mar. Res. Lab.: 50 p.
Lyons, W. G. 1968. Mollusks of Project Hourglass. Am. Malacol.
Union Ann. Rpts. for 1968: 34-35.
Rehder, H. A. and R. T. Abbott. 1951. Some new and interesting
mollusks from the deeper waters of the Gulf of Mexico. Rev. Soc.
Malac, Habana 8(2): 53-66, 2 pi.
Work, R. C. 1969. Systematics, ecology and distribution of the mol-
lusks of Los Roques, Venezuela. Bull. Mar. Sci. /9(3): 614-711.
HEMPHILLIA DROMEDARWS, A NEW ARIONID
SLUG FROM WASHINGTON'
By Branley A. Branson
Eastern Kentucky University
Richmond, Kentucky 40475
During a survey of the terrestrial Gastropoda of the Olympic
Peninsula, Washington, several specimens of the peculiar arionid
slug genus Hemphillia were secured, including an apparently new
species. The genus exhibits a somewhat circumscribed distribution
in Idaho and Alberta, Canada (H. camelus Pilsbry and Vanatta)
(LaRocque, 1953; Pilsbry and Vanatta, 1898; Smith, 1943), Mon-
tana (H. danielsi Vanatta), Oregon, Washington, and British Col-
umbia (Henderson, 1929; Pilsbry, 1948). Pilsbry's (1917) H. mal-
onei, described from a single formalinized specimen collected near
Mt. Hood, Oregon, remains problematic.
Hemphillia and Binneya comprise the subfamily Binneyinae, a
complex of slug species morphologically intermediate between nor-
mally coiled, testaceous snails and shelless slugs in possessing an
exposed shell (partially coiled in Binneya) and short visceral cavity
confined to a dorsal hump or pouch-like arrangement of the body
(Pilsbry, 1948; Webb, 1961). In Hemphillia, the platelike shell is
only slightly attached to the mantle at its edges and, contrary to
Pilsbry's (1948) observation that "in life the shell is usually almost or
quite covered," usually exposed, even at rest. The foot is undivided.
' Supported in part by a Sigma Xi-RESA grant; in part by an Eastern Kentucky
University faculty grant
January, 1972 nautilus 101
Hemphillia burringtoni Pilsbry Fig. lb, d
Pilsbry (1948: pp 741-742, Fig. 397a, b, c, d)
In elevating this form to full species' rank, I was guided by the
consistent diflFerences of genitalia and external pigmentation pat-
terns. Hemphillia glandulosa Bland and Binney, in which Pilsbry
(1948) placed H. burringtoni as a subspecies, exhibits a papillose
mantle in contrast to the smooth one of H. burringtoni; a rugose
stimulator in contrast to a smooth one; and a considerably different
external color pattern. In H. burringtoni, the sides of the foot (Fig.
Id) bear spaced black lines which terminate in round black spots
(between the granules); this pattern is lacking in H. glandulosa.
Holotype and paratypes. Academy of Natural Sciences of Phila-
delphia (ANSP 182093); type locality: Rialto Beach, Clallam
County, Washington.
Distribution: Hemphillia glandulosa occupies mainland Wash-
ington west of the Cascades and adjacent British Columbia west-
ward to the Olympic and Grey Wolf mountains and southward to
northern Oregon. Hemphillia burringtoni is restricted to the Olym-
pic Peninsula of Washington State.
Collecting sites for Hemphillia burringtoni: 1, S17, RllW, T23N,
Macafee Quadrangle, 545 feet mean sea level (MSL), Grays Harbor
County, Washington, 5 July 1969; 1, Bush Pacific State Park, near
Bay Center, Pacific County, Washington, 5 August 1969; 1, S4,
R9W, T27N, rain forest of Mt. Tom Quadrangle, 1,000 feet MSL,
Clallam County, Washington, 11 July 1969; 1, three miles up trail
to Enchanted Valley from Dosewallips Campground, Olympic Na-
tional Park, Mt. Christie Quadrangle, 780 feet MSL, Jefferson
County, Washington, 3 July, 1969; 2, Deer Park Road, 10.3 miles
after leaving U.S. 101, Mt. Angeles Quadrangle, 2,460 feet MSL,
Clallam County, Washington, 7 July 1969; 1, Cox Valley, R5W,
T29N, Mt. Angeles Quadrangle, 3,435 feet MSL, Clallam County,
Washington, 13 July 1969; and 1, North Point Lookout, R2W,
T27N, Mt. Walker, 2,625 feet MSL, Mason County, Washington,
26 June 1969.
The following descriptions were secured from living specimens,
but the measurements are from relaxed, alcoholized individuals.
The jaw is dark-brown in most individuals and bears 10 central
striations, the lateral margins being non-striate. The body is
strongly laterally compressed, almost keel-like, behind the posterior
102 NAUTILUS Vol. 85 (3)
tip of the mantle, and likewise deeply incised (from a lateral view:
Fig. Id) to bear the visceral pouch. Posteriorly, the moderately
developed hornlike protrubrance above the caudal mucus gland is
bluntly rounded behind, but is rather triangular in lateral view.
Colorwise, the posterolateral margins of the mantle are mostly pig-
mentless, except for a distinct band which sends a series of very thin
lines toward the shell to produce a reticulum. The anterior and
dorsal portions of the mantle are much-speckled and blotched with
dark grey and black, and the head and tentacles are black. The
sole is pale yellowish-white and immaculate, whereas the foot im-
mediately above the pedal furrow, which is broken up into 17 to 23
cell-like granules, bears a single row of round, very black spots that
are contacted by thin, oblique lines. The sides of the body below
the dorsal hump are sooty gray to yellowish white, diagonally
marked by 7 to 9 rather broad, dark gray bands. The shell tapers
sharply caudad, the anterior quarter being yellow and the posterior
three quarters greenish-gray; the shell is farther subdivided (in
appearance) by a dense black accumulation of pigment beneath its
middle. The secretory groove, located at the posterior end of the
mantle, is directed nearly straight downward; in life this groove
stands open most of the time. The pneumostome is located just
posterior to the middle of, or as far back as the posterior one-third,
of the mantle. There are a few granules on the mantle caudal to the
shell. Proportional measurements: total length 13.07 mm (8.0-19.3);
width of foot/ total length = 0.20 (0.15-0.24) (in small specimens,
this percentage is larger); posterior end of mantle to pneumos-
tome/total length = 0.37 (0.35-0.40); width of shell/length of
shell = 0.68 (2.0-2.8/2.2-4.5 mm); width visceral pouch/length
visceral pouch — 0.51 (2.5-5.0 mm/5.0-10.4 mm); width back
behind pouch/height behind pouch = 0.39 (0.8-1.4 mm/ 1.8-3.5
mm); length body behind mantle/total length = 0.33 (1.5-9.0
mm/8.0-19.3 mm).
Key to species of Hemphillia
1 a. Body behind pouch at first depressed to receive the visceral
mass then forming a high, compressed keel 2
b. Body behind pouch neither depressed nor forming a com-
pressed keel 3
2 a. Visceral pouch bearing numerous papillae; penial stimulator
rugose within Hemphillia glandulosa
January, 1972
NAUTILUS
103
b. Visceral pouch nearly smooth; penial stimulator smooth within
Hemphillia burringtoni
3 a. Penis narrow, with an accessory sac; color yellowish-gray to
whitish with black markings Hemphillia daniehi
b. Penis broad, lacking an accessory sac; color ashy-gray, bluish-
black to black 4
4 a. Tail with a conspicuous horn-like protrubrance above meeting
of the pedal grooves Hemphillia dromedarius
b. Tail lacking a horn-like protrubrance above meeting of pedal
grooves Hemphillia camelus
Hemphillia dromedarius, new species Fig. la, c
Description of holotype. The head is dark-gray, the tenacles
somewhat lighter. A pair of shallow, pale yellowish white grooves
occupy the head immediately behind the tentacles. The mantle,
including the visceral pouch, is mottled gray, but much whiter on
the sides. Below the visceral pouch, the sides of the body are white
flecked with gray. Posterior to the pouch, the back is moderately
narrowed and rounded rather than keeled, and dark gray on the
midline, lighter below and tending to white flecked with gray along
the sides. The sole and sides of the foot are pale yellowish, the cell-
like granules above the pedal groove being delineated by gray, the
gray streaks coalescing posteriorly to color the edge of the foot gray.
The integument of the back is broken up into diagonal rows of low
1 3. 5 m tn
Fig. 1. A. Holotype, Hemphillia dromedarius, dorsal view. Staircase Falls,
Olympic National Park, Washington. B. Hemphillia burringtoni, dorsal view.
Mt. Christie Quadrangle, 3 miles up Enchanted Valley Trail from Dosswallips
Camp, Olympic National Park, Washington. C. Hemphillia dromedarius, left
lateral view of holotype. D. Hemphillia burringtoni, left lateral view.
104 NAUTILUS Vol. 85 (3)
ridges which emanate from the midline and extend dorsoventrad
and posteriad. The pneumostome, on the right in the posterior one-
half of the mantle, is surrounded by a narrow, white halo. The
shell, located on the posterior one-half of the visceral pouch, is pale
yellowish-horn in color, is transparent, and is approximately twice
as long as wide; its position causes the surrounding integument to
be thrown into five concentric grooves and furrows. The external
surface of the shell shows a series of fine growth ridges. From a
point on the midline of the dorsal pouch, immediately behind the
shell, a definite secretory groove extends posterioventrad slightly
toward the left. A deep caudal mucus pit occurs near the posterior
tip of the tail, and there is a definite horn-like process above it. A
white groove extends to the tip of the tail from the pit. The brown-
ish jaw possesses 20 plates, and the genital pore is located near the
base of the right tentacle. Total length 29.8 mm; width of foot
(4.0 mm) is 13% the length; the measurement from the posterior
tip of the visceral pouch to the pneumostome (6.5 mm) comprises
22% of the length, and the length of the foot posterior to the vis-
ceral pouch (12.5 mm) 42% of the length. The visceral pouch,
7.0 mm/ 13.0 mm, is slightly more than twice as long as wide
(54%). Width of the back behind the dorsal pouch (3.5 mm) is
70% of the depth of the back (5.0 mm.), and the width of the
shell (5.5 mm) is 69% of the length (8.0 mm). Holotype: USNM
577690; type locality (28 June 1969): Staircase Rapids, Staircase
Campground, Olympic National Park, Mt. Steel Quadrangle, Mason
County, Washington; 645 feet MSL.
Specific epithet from that of the Arabian or one-humped camel,
Camelus dromedarius.
Collecting localities for paratypes: 1, 3 miles up trail to Enchanted
Valley from Dosewallips Campground, Olympic National Park, Mt.
Christie Quadrangle, 780 feet MSL, Jefferson County, Washington,
3 July, 1969; Delaware Museum of Natural History (DMNH
43029); 2, slope between the junction of main road and Obstruc-
tion Point Road, Olympic National Park, 4,710 feet MSL, Mt.
Angeles Quadrangle, Clallam County, Washington, 14 July 1969;
1, S32, T25N, RlOW, Queets Campground, Olympic National
Park, 284 feet MSL, Salmon River Quadrangle, Jeflferson County,
Washington, 8 July 1969; 1, R9W, T23W, extreme southeast corner
of Kloochman Rock Quadrangle, 400 feet MSL, Jefferson County,
January, 1972 nautilus 105
Washington, 2 July 1969, Field Museum of Natural History
(FMNH 173022); 1, 5 miles above Flapjack Lakes trail head,
Olympic National Park, Mt. Steel Quadrangle, 3,353 feet MSL,
Jefferson County, Washington, 20 July 1969; 2, 0.3 mile downgrade
from the second station listed above, 4,620 feet MSL, Mt. Angeles
Quadrangle, Jefferson County, Washington, 14 July 1969; 1, North
Point Lookout, Mt. Walker, 2,586 feet MSL, Mason County, Wash-
ington, 26 June 1969.
Corroborative description and proportional measurements. The
mantle color pattern varies from nearly black or blue-black through
dark gray in front of the shell (slightly lighter behind it) to bluish-
black maculated with yellowish or yellowish densely spotted with
blue-black and gray, and is mostly devoid of papillae. Sometimes
the apex of the dorsal hump is creamy orange-yellow, but this is
probably associated with the reproductive period. The sides of the
body below the pouch range from white to yellowish white and
immaculate. The sides of the body behind the pouch bear numerous
gray to blackish maculations. The rather wide edge of the foot is
light gray, and there arc 54 to 56 cell-like granules above the pedal,
groove. The head varies from nearly white to light gray, which
allows the intensely black optic tracts to show through the integu-
ment. The strongly arcuate jaw is striated to its margin, and bears
18 to 20 striae. The secretory groove, located posterior to the shell,
is variable in position: directed directly caudad, slightly to the left,
or slightly to the right. The inflated penis does not bear an acces-
sory sac. Total length 28.5 mm (24.0-31.0); width foot/total
length = 0.13 (2.5-4.0 mm/24.0-31.0 mm); posterior end visceral
pouch to pneumostome/ total length = 0.24 (4.5-8.2 mm/24.0-30.6
mm); shell width/shell length = 0.59 (2.5-6.0 mm/5.0-9.0 mm);
width pouch/length pouch = 0.56 (5.5-7.5 mm/9.9-16.2 mm);
width back behind pouch/height behind pouch = 0.68 (3.3-6.0
mm/5.0-6.0 mm).
Comments on natural history. The oval, semi-opaque eggs, 50 to
60 of which are deposited in wet to moist decaying wood, average
3.3 mm (3.0-3.5) in length and 2.5 mm (2.7-2.7) in diameter. At
rest, this slug is coiled counterclockwise so that the tip of the tail
touches the head. From this position, the animal is capable of
quickly recoiling, even to the extent of "jumping" an inch or so,
a feature which has been previously recorded in the literature
106 NAUTILUS Vol. 85 (3)
(Hemphill, in Pilsbry, 1948, and Pilsbry, Loc. Cit.; Smith, 1943).
This is, to my way of thinking, a very definite anti-predation
startle reaction. The histology and musculature of Hemphillia
needs thorough investigation.
Diagnosis: Hemphillia dromedarius is an arionid slug most closely
related to H. camelus Pilsbry and Vanatta of Idaho and adjacent
Canada but which is distinguished from that species by being
darker and more boldly marked (H. camelus is mostly ashy-gray
with a tendency to produce lateral bands on the visceral pouch,
whereas H. dromedarius tends toward blue-black), by having a less
keeled tail and in possessing a definite caudal horn (Fig. 2) (lack-
ing in H. camelus). It differs from H. danielsi in matters of colora-
tion and in possessing the inflated penis and in lacking an accessory
stimulator
Literature Cited
Henderson, J. 1929. The non-marine Mollusca of Oregon and
Washington. U. Colorado Stud. 17: 45-190.
La Rocque, A. 1953. Catalogue of the Recent Mollusca of Canada.
Bull. Nat. Mus. Canada 129: 1-406.
Pilsbry, H. A. 1948. Land Mollusca of North America (north of
Mexico). Acad. Nat. Sci. Philadelphia Mongr. 3, II (2): i-xlviii;
520-1113.
Pilsbry, H. A. 1917. A new Hemphillia and other snails from near
Mt. Hood, Oregon. The Nautilus 30: 117-119.
Pilsbry, H. A. and E. G. Vanatta. 1898. Revision of the North
American slugs: Binneya, Hemphillia, Hesperarion, Prophysaon
and Adenulus. Proc. Acad. Nat. Sci. Philadelphia 50: 219-261.
Smith, A. G. 1943. Mollusks of the Clearwater Mountains, Idaho.
Proc. California Acad. Sci. 23: 537-554.
Webb, G. 1961. The phylogeny of American land snails with
emphasis on the Polygyridae, Arionidae and Ammonitellidae.
Gastropodia /; 51-52.
WINTER DISTRIBUTION OF
MELAMPVS BIDENTATVS (SAY) ON A
CAPE COD SALT MARSH'
By John W. Grandy IV
National Parks and Conservation Association
1701 18th St. N.W., Washington, D.C. 20009
Salt marsh snails (Melampus hidentatus Say) sometimes comprise
much of the food for wintering black ducks {Arms ruhripes) (Addy,
A portion of the author's Ph.D. dissertation in the Department of Forestry and
Wildlife Management, University of Massachusetts.
January, 1972 nautilus 107
1945). Despite recent studies on Melampus (Apley, et al., 1967;
Russel-Hunter and Apley, 1966), however, the winter distribution
of these snails has not been documented. This paper reports the
results of a two-year study on the winter distribution of salt marsh
snails in Nauset Marsh (a salt marsh), Eastham, Massachusetts.
Salt Marsh. Dexter (1947, pp. 285-290) divided a New England
salt marsh into high and low marsh. The plant species listed by
Dexter that are common in Nauset Marsh are:
High Marsh — salt meadow grass (Spartina patens); sea lavendar
(Limonium sp.); salt grass (Distichilis spicata); glasswort (Sali-
cornia sp.)
Low Marsh — salt marsh cordgrass (Spartina alternifiora); algae
(Fucus sp.); algae (Anscophyllum sp.)
Salt meadow grass is the most common high marsh species in
Nauset Marsh, and salt grass is the most uncommon. Kerwin
(Unpub. Ms.), working in a Virginia salt marsh, recognized a zone
of short Spartina alternifiora (referred to in this paper as short
Spartina marsh). In Nauset Marsh, the short Spartina marsh occurs
between the high and low marsh, and is pronounced when the high
and low marsh are connected by a gently sloping surface. The
ribbed mussel (Modiolus demissus) occurs in the short Spartina
marsh and locally in the high marsh.
Methods. On October 1, 1969, I established 2 plots, 4.5 inches in
diameter, in the high marsh and 2 plots in the short Spartina marsh.
Melximpus movements through these plots were monitored until
mid-November 1969.
Between September 20 and December 27, 1970, I established 20
plots (diameter either 4.5 or 12 inches) in the high marsh or short
Spartina marsh, and marked the snails within with white, quick-
drying enamel paint. These plots were checked periodically to
determine movements of snails to wintering areas. Much of the
information in this report was obtained during more than 80 hours
of observations in fall and winter 1969 and 1970.
Results and Discussion. Plots established in 1969 and before
December, 1970, were of little value. Counting in 1969, and mark-
ing and recounting in 1970 disturbed the habitat, allowing the
marsh surface to dry and apparently forcing the salt marsh snails
to move. Relocating marked snails was hampered by approaching
the marking site, with the chance of stepping or kneeling on marked
108 NAUTILUS Vol. 85 (3)
individuals. In contrast, marking Melampus snails in their winter-
ing locations was valuable, and indicated that the snails remained
sedentary during winter and that they adopted winter habitat in
late November 1970. Major wintering locations for Nauset Marsh
were:
1. High marsh around the bases of salt meadow grass and salt
grass, with shells in upright position with spires from 1/16 inch
below to V4 ir^ch above the peat surface, in clusters up to 20.
(Potential concentrations must depend on Melampus density; hence,
concentrations listed for different wintering areas should be viewed
as being relative). Salt meadow grass provided Uie most consistent
wintering location.
2. Beneath an empty shell (i.e. quahog, Venus mercenaria; ribbed
mussel) on the high marsh wherever the empty shell touches the
marsh surface. Melampus snails lay imbedded slightly in the marsh
surface with their length against the side of the empty shell. Con-
centrations are highly variable, but 23 were found beneath one
empty ribbed mussel shell on January 30, 1971.
3. Around the base and along the sides of the living ribbed
mussel, in upright position and slightly imbedded in the marsh, in
highly variable concentrations up to 30.
4. Around the base of short Spartina alterniflora and within the
sheathing leaves, in upright position, with no more than five win-
tering in each plant.
5. Around the base of Limonium sp. and Salicornia sp., in up-
right position, with no more than one or two Melampus wintering
around each stem.
6. In holes (about the diameter of Melampus) in the high marsh
surface, disassociated from the stems of vegetation but beneath the
salt meadow grass or salt grass, in upright position, with no more
than one snail per hole. Certainly, larger holes, where present,
might contain more snails.
Kerwin (Unpub. Ms.) working in the fall on a Virginia salt
marsh, found Melampus bidentatus associated primarily with salt
grass and salt meadow grass, and secondarily with short Spartina
alterniflora. Dexter (1945, pp. 137-138, and 1947, pp. 288-290)
working in summer on a Massachusetts salt marsh found Melampus
associated with salt grass and salt meadow grass. Hence, Melampus
inhabits the same general marsh areas throughout the year. The
January, 1972 nautilus 109
only apparent differences in summer and winter ecology are the
lack of movement and the tendency to cluster in suitable winter
habitat.
Melampus snails are not always found in each of the locations
noted. Research is needed to determine why, for instance, one area
of salt meadow grass contains no wintering snails while another
area, less than 10 feet away, may contain many snails.
Acknowledgments
I thank Mr. J. A. Hagar for his encouragement and support. Dr.
M. L. Apley furnished information concerning Melampus, and Dr.
John Teal, Woods Hole Oceanographic Institution, reviewed the
manuscript for errors in terminology used to describe a salt marsh.
This paper is a contribution of the Massachusetts Cooperative Wild-
life Research Unit, (Bureau of Sport Fisheries and Wildlife, Massa-
chusetts Division of Fisheries and Game, Wildlife Management
Institute, and University of Massachusetts).
Literature Cited
Addy, C. E., 1945. A preliminary report on the food habits of the
black duck in Massachusetts. Mass. Dept. Conservation, Res.
Bull. no. 6, pp. 1-11.
Apley, M. L., W. D. Russel-Hunter, and R. J. Auslizi, 1967. Annual
reproductive turnover in the salt-marsh snail, Melampus biden-
tatus. Biol. Bull. 133: 455-456. •
Dexter, R. W., 1945. Zonation of the intertidal marine mollusks at
Cape Ann, Massachusetts. The Nautilus 58: 135-142.
, 1947. The marine communities of a tidal inlet at Cape Ann,
Massachusetts: a study in bio-ecology. Ecol. Mono. 17: 261-294.
Kerwin, J. A., Distribution of the salt marsh snail {Melampus bid-
entatus Say) in relation to marsh plants in the Poropotank River
Area, Virginia. Unpub. Ms. on file, Patuxent Wildlife Research
Center, Laurel, Maryland, pp. 1-9.
Russel-Hunter, W. D., and M. L. Apley, 1966. Quantitative aspects
of early life history in the salt-marsh snail, Melampus bidentatus.
Biol. Bull. 133: 392-393.
NOTES
The Western Society of Malacologists' fifth annual meeting
will be held at Redlands University, June 18 to June 21. Symposia
on Pelecypoda led by Dr. Vida Kenk and on Opisthobranchia led
by Mr. Wesley Farmer and papers on other malacological topics will
be presented.
no NAUTILUS Vol. 85(3)
Inquiries may be sent to the secretary, Mrs, Edith Abbott, 1264
W. Cienega Ave., San Dimas, Ca 91773. Applications for member-
ship in the W.S.M. should be sent to the Treasurer, Mr. Ralph O.
Fox, Department of Invertebrate Zoology, California Academy of
Sciences, Golden Gate Park, San Francisco, Ca 94118. Dues are
$2.50 for Regular Members and $1.00 for students.
Executive Board members for the year are: President Mrs. Bea-
trice L. Burch; First Vice-President Mrs. Twila Bratcher; Second
Vice-President Dr. James H. McLean; Secretary Mrs. Edith Abbott;
Treasurer Mr. Ralph O. Fox; Members-at-Large Mr. Anthony
D'Attilio and Mr. Hans Bertsch; the three most recent Past Presi-
dents are Dr. William K. Emerson, Dr. A. Myra Keen and Dr.
Eugene V. Coan.
Locomotion of Marginella olivaeformis Kiener — Members of
the Marginellidae usually have been observed to move by crawling
in the ordinary gastropod fashion. However, at Point of Almadies,
north of Dakar, Senegal, in April 1971, Clover observed Marginella
olivaeformis Kiener, 1834, to progress through loose sand under-
water by a swimming-like motion. The animals were found at
lowest tide level in coarse sand and coral debris among green-slime-
covered rocks.
The foot of M. olivaeformis is broadened anteriorly into two ovate
lobes which are alternately flexed up and down, propelling the
animal forward. In their most strongly reflexed position the lobes
are appressed to the mantle-edge which is held extruded about
halfway around the shell. Thereafter the lobes flap downward with
some force. In a pan of seawater the animals were also seen to
crawl normally on the hard surface, sometimes using the swimming-
like motion in an attempt to scale the sides of the pan.
A broad anterior foot-margin is characteristic of some other Mar-
ginellidae also (notably M. aurantia Lamarck, 1822, and M. hyalina
Thiele, 1913), but its function in these species has yet to be ob-
served.—Barry Roth, 1217 Waller St., San Francisco, Calif. 94117,
and Phillip W. Clover.
WILLIAM H. WEEKS SHELL COLLECTION: New price lists
of this famous collection, with full scientific data^ are in prepa-
ration. Many new additions of fine and rare species are also
included. To obtain free copies write:
George E. Jacobs, 853 Riverside Drive, New York, N. Y. 10032
Vol. 85 April, 1972 No. 4
THE
NAUTILUS
THE PILSBRY QUARTERLY
DEVOTED TO THE INTERESTS OF CONCHOLOGISTS
EDITORS AND PUBLISHERS ^ ., .. ~^^-- — : , ' "'
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Delaware Museum of Natural History, Grebnville, ^^Jp^^/^^lt
Charles B. Wurtz, Consulting ^iologist M W ?l 1^72
3220 Penn Street, Philadelphia, Pa. 19129 • ' '
CONTENTS
Woods Hole, Mais.
Important Notice to Subscribers and Contributors vii
The spawn, early development and larvae oiCyphoma gibbosum
(Cypraeacea). By J. N. Gather and M. E. Crovo Ill
Reef mollusks of South Carolina. By Alan H. Shoemaker .... 114
Somatogyrus alcoviensis, new gastropod species from Georgia
(Hydrobiidae). By K. A. Krieger 120
A second ovoviviparous Nassarius. By Sally Diana Kaicher . . 126
Greggelix, a new genus of autochthonous land snails
(Helminthoglyptidae) from Baja California.
By Walter B. Miller 12^
Shell growth in the gastropod Littorina irrorata.
By Frasier O. Bingham 136
A mollusk new to Lake Birket Qarun, Egypt.
By Kenneth D. Rose 141
Notes and News 144 Publications received .... iv
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THE NAUTILUS
Vol. 85 April, 1972 No. 4
THE SPAWN, EARLY DEVELOPMENT AND LARVAE
OF CYPHOMA GIBBOSVM (CYPRAEACEA)
By James N. Gather
Department of Zoology
University of Michigan, Ann Arbor 48104
and
M. Ellen Crovo
2915 S.W. 102 Avenue
Miami, Florida 33165
During the summer of 1964 while working at the Bermuda Bio-
logical Station for Research, Dr. R. Tucker Abbott pointed out that
the egg capsules and developmental stages of the flamingo tongue,
Cyphoma gibbosum Linne had not been described in the literature.
On July 13 three specimens of C. gibbosum were collected on the
north reef and isolated in a battery jar aquarium with running sea
water and a screen cover. A basal section of the normal prey of
Cyphoma, the rough sea whip Muricea muricata Pallas, was in-
cluded as food.
Between 2000 hrs on July 13 and 0800 hrs on July 14, three egg
capsules were laid in a slightly curving row, with the attachment
plates touching, on the wall of the glass aquarium.
The capsules (fig. 1 ) were hat-shaped and were identical in form
and surface pattern so that each ridge and groove were identical in
shape and position even though there was some variation in size.
The similarity of surface pattern indicates that the capsules were
laid by a single female, for as Ankel (1936) noted in Nassarius and
Philbertia each female has a characteristic "fingerprint"-like pat-
tern in the sculpture of the egg capsules. The same characteristic
is true of Nassarius vibex, Ilyanassa obsoleta, Cypraea caputserpentis,
C. moneta, Conus leopardus, C. pennaceus and C. mindanus. The
basal attachment disc of Cyphoma' s egg capsule varied from 3,12
to 3.39 mm while the egg capsule proper were from 2.37 to 2.62 mm
and 1.50 mm wide. A well-formed operculum covered about half
of the dorsal side of the capsule (fig. 1).
The capsules contained 11, 18 and 19 eggs which were 0.260 mm
in diameter and spherical when first observed at 0800 hours. The
eggs were the same bright orange color as the adult snail's mantle.
Ill
1 12 NAUTILUS Vol. 85 (4)
There were no membranes around the individual eggs, which were
surrounded by the viscous capsular albumen. The lack of indi-
vidual egg membranes in those mesogastropods and neogastropods
with firm multi-egg capsules rather than gelatinous masses has been
verified with the electron microscope (Weymouth et al., 1961;
Taylor and Anderson, 1969) as well as with the light microscope.
The formation of individual egg membranes described by Ghiselin
and Wilson (1965) can only be considered artifactual due to their
admittedly defective histological material. The role they ascribe
to the membrane gland requires re-evaluation.
When the eggs were first observed, both polar bodies had been
extruded. The first cleavage occurred 3 hours and 10 minutes later
at 25°C and was markedly unequal with the smaller AB blastomere
only one-fifth the size of the CD blastomere. The first cleavage was
completed at 4 hours.
We were unable to rear the isolated eggs further, but have since
found that similar large eggs can be cultured in pasteurized sea
water (Costello et al., 1957) with methyl cellulose added to give
the consistency of the capsular albumen, for support of the large
eggs, which will otherwise flatten against the container and cytolyse.
Sodium Sulfadiazine (50 mg/1) and Streptomycin Sulfate (50
mg/1) were added to prevent bacterial growth.
Further observations were made in Miami, Florida, in November
1969. After 11 days in an aquarium, on November 19, one of
four specimens was observed cleaning an area of 8 sq cm of the
glass wall about 3 cm below the water surface. On the morning
of November 20 one of the snails deposited 23 capsules in an
irregular mass on the cleaned area. The snail returned to the egg
mass each morning and added more capsules to a total of 73 by
late evening of November 24. The eggs were initially white but
became light flesh pink by the tenth day, after which they became
progi'essively darkened to a deep rose as the veligers matured. The
embryos were motile by the fifth day and on the fifteenth day the
first veligers escaped from the capsule through a slit in the capsule
top. One average sized capsule contained 1,270 veligers.
In spite of removal of capsules for observation and of predation
by a Cypraea, the mother snail would return each day to the cap-
sule mass. When the veligers hatched the capsules were invaded
by the microfauna and maternal care ceased. Protection and clean-
April, 1972 nautilus 113
ing of egg masses is well known in the Cypraecea (Ostergaard,
1950; Crovo, 1971) but in the cowries the female seldom if ever
leaves the egg mass.
A second snail spawned on November 27 and over a period of four
days produced 78 capsules in a manner similar to that described.
There are some significant differences between the Bermuda and
Florida spawns although all specimens were carefully identified and
the egg capsules are as near identical as is expected within the
normal range of variation of a species. In the Florida forms there
are large numbers of white eggs which hatch rapidly into veligers
while in the Bermuda form the eggs are large and develop slowly
and are bright orange in color.
At present we must attribute these differences to geographic vari-
ation although seasonal variation cannot be excluded. We hope to
obtain more material from a broader geographical range for further
studies.
We wish to thank Martha B. Lackey for her excellent drawing
of the egg capsule, and Axel A. Olsson for encouragement and for
reading the manuscript, and Dr. Alan Kohn for identifying Conus
mindanus.
Literature Cited
Abbott, R. T. 1955. American Seashells. D. Van Nostrand,
Princeton.
Ankel, W. E. 1936. Prosobranchia. In Grimpe, G. and E. Wagler:
Die Tierwelt Nord- und Ostsee. IXbl. Akademische Verlagsge-
sellschaft, Leipzig.
Fig. 1. TTie egg capsule of Cyphoma gibbosum. (a) The basal attachment
disc is rather flat with the body of the capsule shaped like the crown of a hat
with a well formed operculum on top. The line represents 1 mm. (b) A cross
section of the capsule, where indicated.
114 NAUTILUS Vol. 85 (4)
Costello, D. P., Davidson, M. E., Eggers, A., Fox, M. H., and Hen-
ley, C. 1957. Methods for Obtaining and Handling Marine Eggs
and Embryos. Marine Biological Laboratory, Woods Hole.
Crovo, M. E. 1971. Cypraea cervus and Cypraea zebra in Florida
— One species or two? The Veliger, 73; 292-295.
Fioroni, P. 1967. Quelques aspects de I'embryogenese des Proso-
branches (Mollusca, Gastropoda). Vie Milieu, 18: 153-174.
Fretter, V. and Graham, A. 1962. British Prosobranch Molluscs.
Ray Society. London.
Ghiselin, M. T. and Wilson, B. R. 1966. On the anatomy, natural
history, and reproduction of Cyphoma, a marine prosobranch
gastropod. Bull. Mar. Sci., 16: 133-141.
Hyman, L. H. 1967. The Invertebrates: vol. VI, Mollusca I. Mc-
Graw-Hill, New York.
Kay, A. 1960. The functional morphology of Cypraea caputser-
pentis L. and an interpretation of the relationships among the
Cypraeacea. Int. Rev. ges. Hydrobiol., 45: 175-196.
Lebour, M. V. 1932. The larval stages of Simnia patula. J. Mar.
Biol. Ass. U.K., 18: 107-110.
Lebour, M. V. 1945. The eggs and larvae of some prosobranchs
from Bermuda. Proc. Zool. Soc. London, 114: 462-489.
Ostergaard, }. M. 1950. Spawning and development of some
Hawaiian marine gastropods. Pacific Sci., 4: 75-115.
Peile, A. J. 1926. The molluscs of Bermuda. Proc. Malac. Soc.
London, 17: 71-98.
Pelseneer, P. 1935. Essai d'Ethologie Zoologique. Palais des
Academies, Brussels.
Taylor, G. T. and Anderson, E. 1969. Cytochemical and fine struc-
tural analysis of oogenesis in the gastropod, Ilyanassa obsoleta.
J. Morph., 729:211-247.
Weymouth, R., Gather, J. N. and Bigelow, W. C. 1961. A study
of ultrastructure in the mosaic embryo of the snail, Ilyanassa
obsoleta. J. Applied Physics, 32: 1642.
REEF MOLLUSKS OF SOUTH CAROLINA
By Alan H. Shoemaker
Belle W. Baruch Coastal Research Institute
University of South Carolina
Columbia, South Carolina 29208
Collections off Beaufort, North Carolina have verified the exist-
ence of calcareous reefs and associated reef-building corals, Solen-
astrea hyades and Siderastrea siderea (Macintyre and Pilkey, 1969).
The distribution of these reefs appears to coincide with the inner
edge of the Gulf Stream southward, occurring intermittently with
April, 1972 nautilus 115
rocky outcroppings at least as far south as Charleston, South Caro-
lina. Merrill and Petit (1965) found oflf the South Carolina coast
mollusks previously unrecorded or rare north of the Caribbean
Province and south Florida region and they hinted at the possibility
of a much greater occurrence of southern mollusks in Carolina
waters. Later dredging by Menzies et al (1966) recorded over 130
species of mollusks in the Beaufort area, many of which were,
indeed, unknown north of the Bahamas and Florida Keys. Merrill
and Petit (1969) further added to this list of known molluscan
species oflf South Carolina and pointed out the need for more
research in this area. In October, 1970, a survey was undertaken
aboard the R/V Eastward of the Duke University Marine Labora-
tory under the direction of F. John Vernberg of the Belle W. Baruch
Coastal Research Institute, University of South Carolina, to verify
the occurrence of these reefs off South Carolina. Since broken
patches of rocky outcroppings and coral reefs were found off
Charleston and Georgetown, South Carolina, later dredgings in
March, 1971 were undertaken to examine the faunal diversity.
This paper will deal with the molluscan assemblage found on the
March, 1971, cruise.
This cruise was made possible by NSF grant GB- 17545 to Duke
University.
Physical Characteristics of the Reef
The reefs which lie parallel to the Gulf Stream occur in the area
off Georgetown as raised benches, rounded humps, or ridges. Other
areas within the vicinity are not raised but are still hard, rocky,
and contain coral, as evidenced by large numbers of byssally
attached Area zebra. The reefs and flat rocky areas are interrupted,
however, by sandy patches. Typical sand bottom bivalves, such as
Glycymeris americana and Macrocallista maculata, are found wher-
ever these patches occur. The area surveyed is roughly 32°55'N
and 78°45'W. Stations 16621 and 16624 are deeper reefs located
off Charleston and contain no living coral. Only the shallower
reefs of 25m to 30m depth contain living, reef-building corals, while
the first two deeper reefs supported a much smaller faunal com-
munity, molluscan or otherwise.
Method of Collection
Stations not included in this listing, as evidenced by gaps in the
numbering sequence, represent sand bottoms or other shipboard
116 NAUTILUS Vol. 85(4)
Table 1 - Location of the reefs.
Station
16621
16624
16633
16634
16635
16642
16643
16646
events not important to the present paper. Each station required
30 minutes to complete in order to fill the dredge. A rock dredge
was used at all stations and all material was collected in a heavy
chain link bag trailing the dredge. Although very small mollusks
may be lost in this dredge, a fine mesh dredge which could be used
on soft bottoms would be damaged or lost on rocky bottoms like
these. Material to be kept alive comes aboard in better condition
when collected in a chain bag which has enough flexibility to allow
for abrasion against other incoming materials.
BlVALVIA
At first glance, all of these bivalves appear to be southern or
tropical species. Very few are found north of Cape Hatteras, North
Carolina, where colder currents seem to keep them from extending
their range. The occurrence of Crassostrea virginica, Noetia pon-
derosa, Mercenaria mercenaria, and Dinocardium robustum prob-
ably marks the area as a shallow marine or estuarine habitat in
Pleistocene times, since all of these shells were dead and appear to
be subfossil remains. All other records are from living or, in a
few instances, freshly dead individuals. Macrocallista maculata and
Glycymeris americana are typically sand bottom species and, as
mentioned previously, are found on patches of sand which occur
intermittently among the reefs and rocky areas. Area zebra, listed
April, 1972
NAUTILUS
117
Table 2 - Mollusc*
Sample stations
as abundant at most stations, was present in great numbers and
probably paves the rocky bottoms, with many attached to each
other by their strong byssal threads. They also provide a substrate
for the equally common Chama congregata, Chama maeerophylla,
Pseudochama radians, and Crepidula aeuleata, and many corals,
bryozoans, and tunicates. The boring mussels Lithophaga aristata
118
NAUTILUS
Vol. 85 (4)
raoie i - cone.
Sample stations
Rsrare,l individual;C=coinmon,2-4 individuals; and A=abundant,over 4 individuals.
and Lithophaga bisulcata were common, found in tunnels dug in
coral heads and in burrows within the shells of Spondylus ameri-
canus. The other species were much less common, occurring only
once or twice, possibly because their habitat is truly limited.
April, 1972 nautilus 119
Gastropoda
Of all the gastropods collected, few species were found repeatedly.
Credidula aculeata and Cantharus tinctus were found in large
numbers on coral and other mollusks, but since most of the other
species are voracious predators, their numbers are quite limited by
available food supply, intraspecific competition, low reproduction
rates, and other natural population controls. While Fasciolaria
hunteria is probably common on the reef as a whole, the Murex
family also represents an abundant predator group. Some of these,
Phalium granulatum and Conus sozoni, are also found on sandy
bottoms which may indicate that their environmental requirements,
such as food supply, are not necessarily reef dependent. Only three
Murex dilectus were found on the cruise, all alive, with one indi-
vidual quite large and spinous. This species has not been recorded
north of southeast Florida or the Caribbean (Clench and Abbott,
1944; Abbott, 1954). All of the M. dilectus were of the frilly variety
and varied in color from tan in the largest one to dark brown in
the smallest one. Turritella exoleta, although found only as solitary
specimens on this cruise, was found in large numbers within one
sample on an earlier cruise, which may be indicative of a gregarious
or colonial mode of life. In general, gastropods represent only a
fraction of the total numbers of individuals of the molluscan
assemblage, with Area zebra alone comprising over 75% of the total
number of individual mollusks, and only a small fraction repre-
sented by the largely predaceous gastropods.
Conclusion
The reefs off the Carolinas need to be more thoroughly sampled,
both for mollusks and for other phyla with tropical representatives.
This particular cruise was cut short by an untimely snow storm.
Reef locations south of Charleston are unknown as yet, and although
they presumably follow the Gulf Stream all the way to Florida,
further work is needed to accurately plot them.
Literature Cffed
Abbott, R. Tucker. 1954. American Seashells, D. Van Nostrand
and Co., N.Y. 541p.
Clench, William J. and Farfante, I. Perez. 1944. The Genus Murex
of the Western Atlantic, Johnsonia, vol. 1, Museum of Compara-
tive Zoology, Cambridge, Mass.
Macintyre, Ian G. and Pilkey, Orn H. 1969. Tropical Reef Corals:
120 NAUTILUS Vol. 85 (4)
Tolerance of Low Temperatures on the North Carolina Conti-
nental Shelf, Science, 166: 374-375.
Menzies, R. J.; Pilkey, O. H.; Blackwelder, B. W.; Dexter, D.;
Ruling, P.; and McCloskey, L. 1966. A Submerged off North
Carolina, Int. Revue ges HydrobioL, 51: 3, 393-431.
Merrill, Arthur S. and Petit, Richard S. 1965. Mollusks New to
South Carolina, Nautilus, 79: 58-65.
Merrill, Arthur S. and Petit, Richard E. 1969. Mollusks New to
South Carolina: II, Nautilus, 82: 117-122.
SOMATOGYRUS ALCOVIENSIS, NEW GASTROPOD SPECIES
FROM Georgia (Hydrobiidae)
By K. A. Krieger
2652 Williamsburg Drive, Decatur, Ga. 30034
Much of the interior of Georgia has never been fully explored by
invertebrate zoologists, and until recently the Mollusca have been
particularly neglected. Family Hydrobiidae, which includes those
prosobranch gasti'opods with basocones on the central tooth of the
radula (Thompson, 1968), is represented by many genera through-
out the Southeastern States. Somatogyrus, itself widely distributed,
belongs to subfamily Hydrobiinae, in which a single functional
duct, the vas deferens, discharges at the tip of the penis (Morrison,
1949; Thompson, 1968). The present paper extends the known
range of Somatogyrus into the headwaters of the Altamaha River
system of Georgia.
Materials and Methods: Specimens were collected by hand or
sieve in shallow water on 5 September 1968, April and May 1970,
and 4 January 1971. The soft anatomy was described from living
specimens. Permanent radular mounts were prepared after dissec-
tion and mercurochrome staining. The verge was sketched from
animals preserved in sixty per cent ethyl alcohol.
SomatogjTus alcoviensis, new species (Figs. 1-4)
Description: The shell is small, thin, and globose, reaching 5.1
mm. in length and 4.6 mm. in greatest diameter (Fig. 1). The three
whorls are strongly convex, smooth, and unsculptured with numer-
ous fine transverse striations. The apex is often eroded. The sutures
are shallow. The aperture is large and widely ovate but acutely
angular posteriorly, presenting a white interior. The outer lip is
thin, not reflected; the inner lip is discontinuous, becoming a thin
callus along a wide attachment to the body whorl. The umbilicus
is reduced to a very narrow chink or is imperforate. Although the
April, 1972 nautilus 121
shell may be coated with a black deposit, the shell itself is yellow-
green.
Paratypes of Clappia umbilicatus and C. cahabensis revealed a
mean obesity index (width/height) of 1.01 (0.92-1.04) and 0.95
(0.87-1.00), respectively. Fifty specimens of Somatogyrus alcov-
iensis each from the Yellow River and Alcovy River populations
revealed respective mean obesity indices of 0.94 (0.86-1.06) and
0.95 (0.85-1.04).
The operculum (Fig. 2) is chitinous, ovate, and spiral with about
2.2 whorls. Externally, several strong growth lines and many finer
striations radiate from a prominent nucleus, which is situated about
one third the distance between the base and the apex and between
the anterior and posterior margins. The inner opercular surface is
featureless and smooth, but occasionally reflects the major external
features.
The head is black; the rest of the animal is gray, the ventral foot
and the verge being lighter. Tentacles are slender. Arising beneath
the mantle collar behind the right tentacle, the verge is simple,
compressed dorsoventrally, gradually tapers distally to a point, and
lacks any secondary features such as papillae or glands. When
relaxed the verge extends anteriorly then recurves posteriorly to the
left over the neck; the penis is indistinguishable from the verge
(Fig. 3)
The radula (Fig. 4) possesses the standard hydrobiid formula
2.1.1.1.2. There are about 65 rows of teeth. The central tooth
arises from a broad base with four ectocones on each side of a
larger mesocone, and with four basocones on each side of the ante-
rior face. A rather rectangular projection extends slightly anteriorly
from the lower face. The lateral teeth possess an extended, narrow
shaft which bends outward about halfway along its length; the
reflection contains four to six sharply pointed ectocones, four simi-
larly pointed entocones, and a larger, blunt mesocone. A very large,
bluntly pointed cusp projects infero-anteriorly from the lower face
of the tooth. The marginal teeth curve gradually inward from the
base, culminating in thirty to forty small, pointed cusps; the shaft
of the inner marginal is slightly flanged.
The eggs are laid singly or in small, irregular clusters. The
embryo is visible through a clear matrix, which occasionally is
partially coated with the black deposit.
122 NAUTILUS Vol. 85 (4)
Dimensions: Holotype: 3.0 mm. in length, 2.9 mm. in greatest
diameter; largest para type (Yellow River): length 5.1 mm., diam-
eter 4.6 mm.; smallest paratype (Alcovy River): length 1.5 mm.,
diameter 1.6 mm.
Type Localities: Holotype and paratypes are from Cedar Shoals
in Yellow River about 1.0 km. S. of Porterdale, Newton County,
Georgia. Additional paratypes are from Newton Factory Shoals in
Alcovy River immediately above Jackson Lake in Newton County,
Georgia. The species was named after its geographical location.
Type Depositories: From Yellow River: Holotype, Museum of
Comparative Zoology, Harvard University No. 277838; twelve para-
types in each of the following: Harvard University No. 277839; U.S.
National Museum, No. 701914; Florida State Museum, University
of Florida, Gainesville, UF No. 21453; Delaware Mus. Nat. Hist.
No. 41741. Additional paratypes from Alcovy River: Museum of
Comparative Zoology, No. 277840; USNM No. 701915; UF No.
21454; Delaware Mus. Nat. Hist. No. 41742. All specimens were
collected alive by the author.
Discussion: Somatogyrus alcoviensis possesses the prominent cusp
projecting infero-anteriorly from the face of the lateral tooth upon
which Walker (1909) primarily erected the genus Clappia. How-
ever, in few other respects does the present species resemble Clappia.
Comparison with paratypes of C. clappi Walker 1909 ( = S. umbili-
catus Walker 1904; see Goodrich, 1944) and C. cahnbensis Clench
1965 revealed that, while S. alcoviensis possesses an extremely
reduced or entirely imperforate umbilicus, the umbilicus of C. um-
bilicatus and C. cahabensis is much larger and deep. In addition,
S. alcoviensis presents a discontinuous columellar lip which is suc-
ceeded by a thin callus, although in C. umbilicatus and C. caha-
bensis the columellar lip is continuous and is attached to the body
whorl only at its upper margin. Thus, the characteristics of the
umbilicus and columellar lip preclude placing the present species
within genus Clappia, because "In shell characters, Clappia differs
from Somatogyrus in the conspicuous deep umbilicus, the straight,
thin inner lip without any callus thickening, which is entirely
separate from the body whorl, except for a very short distance at
the upper extremity" (Walker, 1909).
The present species is assigned to genus Somatogyrus because it
possesses shallow sutures, a white callus across the columellar mar-
April, 1972 nautilus 123
gin of the aperture, and four basocones on each side of the central
tooth of the radula (Thompson, 1968).
Somatogyrus alcoviensis is very similar to S. (Walkcrilla) tenax
recently described from Broad River, Elbert County, Georgia
(Thompson, 1969), differing only in a more globose shell, more
slender tentacles, somewhat lighter pigmentation, and a different
number and arrangement of cusps on the teeth. The entire animal
of S. alcoviensis is generally lighter in color than S. tenax; the
mantle collar of S. alcoviensis remains light gray, although that of
S. tenax is light grayish-orange. The central tooth of S. alcoviensis
possesses four basocones and only four ectocones on each side, not
three basocones or up to six ectocones on each side as in S. tenax.
The lateral tooth bears four to six ectocones but only four ento-
cones in S. alcoviensis, while in S. tenax there are either five or six
ectocones and only three entocones. The marginal teeth possess
about twice as many cusps as in S. tenax.
Ecology: Somatogyrus alcoviensis is restricted to shoals, where it
crawls on boulders, gravel, and vegetation, being absent from silt
and bottom sediments. Locally it may number in the thousands per
square meter, particularly in the dense mats of Podostemum cera-
tophyllum (riverweed) which cover submerged rocks in rapid water.
By its habits and radular form, S. alcoviensis probably is an auf-
wuchs and detritus feeder. It serves as food for the sunfish, Lepomis
sp., and is closely associated with several other mollusks, including
Pisidium sp., Sphaerium fabalis, Elliptio hopetonensis, E. productus,
and Oxytrema suturalis (Krieger, 1969). The eggs are deposited on
Podostemum stems and on the shells of living Oxytrema and
Somatogyrus.
April and May 1970 collections of S. alcoviensis from Yellow
River yielded specimens considerably larger (up to 5.1 mm. long,
4.6 mm. wide) than specimens collected concurrently from Alcovy
River (up to 4.1 mm. long, 3.9 mm. wide), and larger than speci-
mens collected at any other time from Yellow River (September
1968, January 1971). It was at this time, also, that eggs were
obtained from the Yellow River population. Apparently S. alcov-
iensis is confined to a life cycle of only one year, rapidly attaining
its greatest size in early spring and dying a few weeks after deposi-
tion of the eggs. Failure to obtain eggs or specimens of a similar
size from Alcovy River may indicate a later chronology of the life
124 NAUTILUS Vol. 85 (4)
cycle due to different stream conditions, such as cooler tempera-
tures, in Alcovy River. The pH and oxygen content of both local-
ities, however, are extremely similar (Krieger, 1969). It is, of
course, entirely possible that the Somatogyrus in Alcovy River reach
maturity without attaining a comparable size due to genetic or
environmental factors. In all other characteristics observed, both
populations appear to be identical.
Acknowledgments
The author is grateful to Drs. W. D. and Madeline P. Burbanck,
Department of Biology, Emory University, for encouraging this
study and for generously providing use of their facilities. Appre-
ciation also is extended to Dr. James E. Sublette, Department of
Biology, and to Dr. A. L. Gennaro, Curator, Natural History
Museum, Eastern New Mexico University, as well as to the staff
of the 27th Tactical Hospital, Cannon AFB, N. M., for the use of
their facilities. Paratypes of Clappva clappi and C. cahabensis were
loaned by Dr. Kenneth J. Boss, Museum of Comparative Zoology,
Harvard University. Dr. Fred G. Thompson, Florida State Museum,
University of Florida, reviewed the manuscript and contributed
valuable information concerning the systematics of this species.
Literature Cited
Clench, William J., 1965. A new species of Clappia from Alabama.
The Nautilus 75(1), pp. 33-34.
Goodrich, Calvin, 1944. Certain operculates of the Coosa River.
The Nautilus 58(1), pp. 1-10.
Krieger, K. A., 1969. Factors controlling the distribution of Oxy-
trema suturalis Haldeman in the Yellow River of Georgia. Mas-
ter's Thesis, Emory University.
Morrison, J. P. E., 1949. The cave snails of eastern North America.
News Bull. Ann. Rept. Am. Malac. Union, 1948, pp. 13-15.
Thompson, Fred G., 1968. The aquatic snails of the family Hydro-
biidae of peninsular Florida. Univ. Florida Press, Gainesville,
268 p.
— 1969. Some hydrobiid snails from Georgia and Florida. Quart.
Jour. Florida Acad. Sci., vol. 32(4), pp. 241-265.
Walker, Bryant, 1909. New Amnicolidae from Alabama. The
Nautilus 22(9), pp. 85-90.
April, 1972
NAUTILUS
125
Fig. 1. Views of two paratypes of Somatogyrus alcovicnsis Krieger new species.
Lengtfi 3.0 mm., greatest diameter 2.9 mm. (Delaware Museum Natural His-
tory, no. 41741).
Figs. 2-4. Somatogyrus alcoviensis Krieger, new species. Fig. 2. Outer surface
of operculum. Fig. 3. Verge in relaxed position; broken line near base indicates
the degree of dorsoventral compression. Fig. 4. Radula, frontal view: (1-r)
central tooth, lateral tooth, inner and outer marginal teeth.
126 NAUTILUS Vol. 85 (4)
A SECOND OVOVIVIPAROUS ISASSARIVS
By Sally Diana Kajcher
30 160th Avenue
Redington Beach, Florida 33708
Nassarius muelleri (Maltzan, 1884), from West Africa has been
known for some time to be ovoviviparous (Knudsen, 1956), but
this reproductive process has not been shown in any other member
of the genus. Two specimens of Nassarius albus (Say, 1826),
(? Buccinum ambiguum Pulteney, 1799, non Solander, 1766) en-
countered during my current investigation of western Atlantic
Nassarius disclosed this shallow water species to be ovoviparous,
also.
A beach-drift specimen of N. albus (figure 2a) from Bermuda
collected by Mr. Arthur Tucker Guest contained the dried animal.
It had withdrawn so far into the shell that it was necessary to cut
a small hole in the dorsal side of the shell to dislodge the remains.
Six embryonic shells ranging from 1.2 to 1.5 mm in diameter were
found within. Further probing produced the dried remains, includ-
ing the radula, of the adult.
A second, slightly larger specimen (figure 2b) was collected alive
by Dr. Emily Yokes in less than six inches of water during an
extremely low tide in December, 1970, approximately 12 miles
south of Champoton, Campeche, Mexico. This female, preserved
in alcohol, carried 18 embryos as well as four incompletely devel-
Fig. 2. Shells of female Nassarius albus (Say). 2a, from Bermuda. 2b, from
Campeche, Mexico.
April, 1972
NAUTILUS
127
oped eggs. Although the soft parts broke as they were being
extracted, enough remained intact to clearly show the embryos in
the fragile oviduct (figure la). The embryos are about 1 mm in
diameter and wcU-dcvclopcd, with eyes clearly visible through the
transparent shell (figure lb). It seems possible that the four
incompletely developed eggs (figure Ic) are undevoured nurse eggs.
The shells of the two specimens, collected from widely separated
Bermuda
Figs. 1 and 3. Nassarius albus (Say). Fig. la, soft parts of Bermuda female,
lb, embryos. Ic, incompletely developed eggs. Fig. 3a. radular row, from Ber-
muda. 3b, radular row from Campeche, Mexico.
128 NAUTILUS Vol. 85 (4)
localities, differ somewhat in form and sculpture, a common occur-
rence in this genus. Both fall within the range of shell variation
I have observed for the species. The radulae (figure 3) are prac-
tically identical.
I am indebted to Mr. Arthur Tucker Guest of Crawl, Bermuda,
and Dr. Emily Yokes of Tulane University for the specimens, and
to Mr. William Lyons, Florida Department of Natural Resources
Marine Laboratory, St. Petersburg, Florida, for his advice and
friendly criticism during the preparation of this paper.
Literature Cited
Knudsen, J. 1956. Marine Prosobranchs of Tropical West Africa
(Stenoglossa) — Atlantide Rep. 4, pp. 53-54.
GREGGELIX, A NEW GENUS OF AUTOCHTHONOUS
LAND SNAILS (HELMINTHOGLYPTIDAE) FROM
BAJA CALIFORNIA
By Walter B. Miller
Department of Biological Sciences, University of Arizona
Tucson, Arizona 85721
Helix lohrii Gabb, 1868, was described from a dead specimen
(ANSP #58106) collected "from the higher table lands near
Moleje" (i.e. Mulege of modern maps), Baja California Sur, Mexico;
in accordance with Articles 27 and 32 (c) (i) of the International
Code of Zoological Nomenclature (pp. 29, 35-36), the trivial name
lohrii should be written loehri as will be done here.
Sonorella lohrii (sic) lioderma Pilsbry, 1904, was described from
several dead specimens found "near Moleje, Lower California."
Subsequently, Pilsbry (1916) raised S. lioderma to specific rank and
figured the holotype. He then also synonymized Helix steganella
Mabille, 1895, with ''Sonorella" loehri. Other workers (G. D.
Hanna and A. G. Smith, 1968) have placed loehri in the genus
Micrarionta Ancey, 1880. However, thus far, the lack of data on
the reproductive anatomy of loehri and liodernia has prevented
their positive generic identification.
I am pleased to report that in December 1970, my son William
Nixon Miller and I succeeded in collecting a large number of
live lioderma specimens from two separate localities in San Jose
Comondu and near La Purisima, Baja California Sur. Dissection
April, 1972 nautilus 129
of these specimens reveals that they belong to a new genus of
helminthoglyptids, described below. Their positive identification
was made possible by comparison with two paratypes of S. lioderma
kindly provided by the Academy of Natural Sciences of Phila-
delphia. A paratype of lioderma provided by the ANSP is shown
in Plate 1, along with the shell of one of my dissected specimens,
#5313-X, from La Purisima. The holotype of Helix loehri Gabb
(ANSP #58106) was also provided by the ANSP for examination;
it had the characteristic granular sculpture of the body whorl as
described for that species.
In 1966, G Dallas Hanna visited the Paris Museum d'Histoire
Naturelle and examined and photographed some of Mabille's types
of Baja California mollusks. Based on his observations, he and
Allyn G. Smith (1968) then concluded that Micrarionta peninsu-
laris (Pilsbry, 1916) was a synonym of Helix indigena Mabille,
1895; that Helix steganella Mabille, 1895, and H. invecta Mabille,
1895, were synonyms of Micrarionta lohrii (sic) Gabb 1868); and
that Helix digueti Mabille, 1895 was a nomen inquirendum because
the type material was not found.
I subsequently visited the Paris Museum in August 1971, and
Mr. Henry Chevallier kindly showed me Mabille's types exactly as
they had been set aside for Hanna, with the addition of the types
of Helix digueti. It was immediately apparent that Helix indigena
Mabille, 1895, p. 64, Helix digueti Mabille, 1895, p. 65, and
Sonorella lioderma Pilsbry, 1904, are all the same species, with the
name indigena Mabille, having priority. Helix invecta Mabille,
1895, p. 65, and Helix steganella Mabille, 1895, pp. 64, 65, are very
heavily granulated shells very close to H. loehri Gabb. However, in
the type of H. loehri, the granulations are located on the radial
striae whereas in H. invecta and H. stega7^ella the granulations are
so thick as to obliterate any underlying striation. I have several
lots of specimens sent to me by Munroe Walton and collected in
1970 by Charlotte Church (nee Walton) from localities near San
Javier, near Loreto, which completely match the sculpture of H.
invecta and H. steganella but not that of the type of H. loehri. At
present, no population with the characteristic sculpture of H. loehri
has been rediscovered.
Sonorelix peninsularis (Pilsbry) is not a synonym of Helix indi-
gena Mabille, and is a perfectly valid name.
130 NAUTILUS Vol. 85 (4)
Pending the procurement of live animals of H. loehri and H.
invecta, it is premature to speculate on the specific or subspecific
standing of these names. It appears most probable, however, that
they are congeneric with indigena, which I place in the new genus
described below.
Greggelix W. B. Miller, gen. nov.
Type species: Greggelix indigena (Mabille, 1895)
I am pleased to name this new genus in honor of Wendell O.
Gregg, friend and colleague, who introduced me to the snails of the
desert southwest.
Diagnosis: The new genus Greggelix is distinguished from other
genera in the family Helminthoglyptidae by the following anatomi-
cal characteristics of the reproductive system: No dart, dart sac, or
mucus glands; a short verge (penis papilla); a very long epiphallic
caecum and a very long spermathecal diverticulum, each from
nearly as long to decidedly longer than the spermathecal duct.
Description of the reproductive anatomy of the type species,
Greggelix indigena (Mabille, 1895) (Figure 1) is as follows:
The penis is short and saccular, equipped with a short, bulbous
somewhat spherical verge. The epiphallus is about twice as long
as the penis. The extremely long epiphallic caecum is compressed
into about twenty tight coils which first wind distally from the
epiphallus, then double back proximally and finally reverse again
with the distal end lying loosely, uncoiled, in the body cavity. The
coils are held tightly together by a blood vessel and connective
tissue. A penial retractor muscle inserts on the epiphallus about
midway along its length and attaches to the floor of the lung near
the mantle collar.
The vagina is about equal in length to the penis. The sperma-
thecal duct lies uncoiled along the uterus-prostate complex bound
by connective tissue. The spcrmatheca is located in its usual posi-
tion for helminthoglyptids, namely just posterior of the ventricle,
held tightly appressed by connective tissue and a loop of the
anterior aorta. A very long spermathecal diverticulum originates
near the proximal end of the spermathecal duct and lies bound
against the uterus in loose coils, with its distal end just anterior of
the albumin gland. The albumin gland, hermaphroditic duct, and
ovotestes are typically helminthoglyptid.
All specimen ts #5313 are from La Purisima Canyon, in a lava
April, 1972
NAUTILUS
131
diverticulum is noticeably shorter than in the other specimens but
rockslide 3.0 mi. cast of San Isidro along the road to Canipole;
specimen #5309 is from San Jose Comondu in a rockslide at the
edge of town along the road to San Javier. It is to be noted that
in one of the La Purisima specimens (5313-C) the spermathccal
Plate 1. A— C. Greggelix indigena (Mabille), WBM #531 3-X, La Purisima.
Baja Calif. Sur. D— F. Greggelix indigena (Mabille) PARATYPE, ANSP
#88367, "near Moleje, Lower California." Scale in mm.
132
NAUTILUS
Vol. 85 (4)
April, 1972 nautilus 133
is still as long as the spermathecal duct. The San Jose Comondii
specimen (5309) has a somewhat longer penis than found in the
La Purisima specimens, while its spermathecal diverticulum is
slightly shorter than the spermathecal duct. Since it was the only
live adult from that locality, no conclusions can be reached on
whether these differences are consistent and significant.
Live specimens of both Comondu and La Purisima populations
had the mantle collar pigmented greenish-yellow in contrast to the
usual orange of most Sonorelix and Sonorella. This pigmentation
is not considered a generic characteristic, but it is usually consistent
at the subspecific or specific level.
Explanation to Opposite Page
Fig. I. Greggelix indigena (Mabille, 1895). Lower genitalia. All drawings in
figures 1-4 made from projection of stained whole mounts. Abbreviations used:
Fig. 2. Sonorelix (Sonorelix) borregoensis (Berry, 1929). Genitalia. Fig. 3.
Sonorelix (Herpeteros) peninsularis (Pilsbry, 1916). Lower genitalia. Fig. 4.
Micrarionta (Xerarionta) areolata (Pfeiffer, 1845). Lower genitalia.
134 NAUTILUS Vol. 85 (4)
Discussion
This genus appears to be most closely related to Sonorelix Berry,
1943, Sonorella Pilsbry, 1900, Mohavelix Berry, 1943, and Tryon-
igens Pilsbry, 1928, in that all of these genera have lost the dart
apparatus. Direct phylogenetic relationship among these dart-less
genera is not to be inferred, however, since evidence tends to indi-
cate that the loss of the dart apparatus has occurred more than
once in the evolution of the helminthoglyptids (Miller 1970).
In order to compare the reproductive anatomy of Greggelix with
that of Sonorelix, drawings were prepared from stained whole
mounts of the genitalia of Sonorelix (Sonorelix) borregoensis
(Berry, 1929) (Fig. 2) and Sonorelix (Herpeteros) peninsularis
(Pilsbry, 1916) (Fig. 3). It can be seen that in both subgenera of
Sonorelix the epiphallic caecum and the spermathecal diverticulum
are much shorter than the spermathecal duct. There are other
salient differences. The verge of S. (Herpeteros) is relatively enor-
mously large and bulbous, a characteristic of the subgenus (Berry,
1947) found not only in S. (H.) peinsularis but also in S. (H.)
angelus Gregg, 1948 and S. (H.) inglesiana (Berry, 1928) not fig-
ured here). The penis and vagina of Sonorelix s. s. are both un-
usually long and a muscular vaginal node is present; this is the
case not only in S. (S.) borregoensis but also in S. (S.) rixfordi
(Pilsbry, 1919), S. (S.) melanopylon (Berry, 1930) and S. (S.)
avawatzica eremita (Pilsbry, 1939) all of which were also examined
but not figured. Whether Sonorelix s. s. and Sonorelix (Herpeteros)
are truly congeneric remains to be determined.
It is interesting to note that the unusually long epiphallic caecum
and spermathecal diverticulum of Greggelix are found elsewhere
only in certain species of Helminthoglypta and in Micrarionta
(Xerarionta) (Fig. 4). A prolific population of M. (X.) areolata
(Pfeifler, 1845) occurs all along the Magdalena Plain of Baja Cali-
fornia Sur, separated from the Comondu Canyon population of
Greggelix by not more than ten miles. While one cannot infer
phylogeny simply on size of spermathecal diverticulum and epi-
phallic caecum or on geographical proximity, it is tempting to
hypothesize an ancestral Xerarionta population attempting to sur-
vive an increasingly drier climate in the rockslides of the Sierra
de la Giganta and losing the dart apparatus through genetic drift
and saltational speciation (Miller, 1970). An extensive survey of
April, 1972 nautilus 135
chromosome numbers and karyotypes of many of the helmintho-
glyptid genera and subgenera has now been undertaken by Noor-
ullah Babrakzai at the University of Arizona and it is possible that
this will yield information useful in determining phylogeny more
precisely.
Acknowledgments
I wish to thank the following for invaluable assistance in the
preparation of this article: The Academy of Natural Sciences of
Philadelphia for the loan of the holotype of Helix loehri and two
paratypes of Sonorella lioderma; Mr. Henry Chevallier, Paris Mu-
seum d'Histoire Naturelle, for making available Mabille's types of
Helix indigena, H. digueti, H. steganella, and H. invecta; Dr. W. O.
Gregg for the loan of whole mounts of several species of Sonorelix
s. s.; Munroe Walton and his daughter Charlotte Church for several
lots of Greggelix shells from many localities in Baja California Sur;
my laboratory assistant, Laura Emmett, for photographs of the bor-
rowed types; my son, Nick, for his assistance and fantastic collec-
tor's "luck" in obtaining live specimens; and Dr. J. C. Bequaert for
his steady encouragement and criticism of this article.
Literature Cited
Berry, S. S. 1947. On the generic relationships of certain Lower
Californian helicoid snails. Leaflets in Malacology 1 (3): 9-12.
Gabb, W. M. 1868. Description of new species of land shells from
Lower California. American Journal of Conchology 3: 235-238,
pi. 16.
Gregg, W. O. 1948. A new and unusual helicoid snail from Los
Angeles County, California. Bull. So. Calif. Acad. Sci. 47 (3) :
100-102.
Hanna, G. D. and A. G. Smith. 1968. The Diguet-Mabille land
and freshwater mollusks of Baja California. Proc. Calif. Acad.
Sci. 30 (18): 381-399.
Mabille, J. 1895. Mollusques de la Basse Californie. Bulletin
Societe Philomatique Paris 8 (7): 54-76.
Miller, W. B. 1970. A new species of Helminthoglypta from the
Mojave Desert. The Veliger 12 (3) : 275-278.
Pilsbry, H. A. 1904. A new Lower Californian Sonorella. The
Nautilus 18 (5) : 59.
Pilsbry. H. A. 1916. Helices of Lower California and Sinaloa. The
Nautilus 29 (9): 97-102, pis. 2-3.
136 NAUTILUS Vol. 85 (4)
SHELL GROWTH IN THE GASTROPOD
LITTORIJSA IRRORATA
By Frasier O. Bingham
Rosenstiel School of Marine and Atmospheric Science
University of Miami, Florida 33149
Littorina irrorata (Say, 1822), a rather globose, light gray snail
known as the Salt Marsh Periwinkle, is common in the salt marshes
from New York to the Rio Grande River, excluding those of
southern Florida. Presented here are observations on several char-
acteristics of the shell of L. irrorata. Behavior of the species has
been treated in a separate article (in press).
Growth
Several hundred specimens of L. irrorata of various lengths were
collected at random from a salt marsh near Panama City, Florida,
and sorted into size classes. Four of the classes were replaced in the
marsh on March 8, 1969, and collected five months later on August
9, 1969. The classes were handled somewhat differently in order
to prevent breakage of smaller shells and the space requirements of
the large snails. Separate plastic screen cages were used to retain
classes 1 and 2 (Figure 1). The exterior surface of the shell lip of
each specimen in class 3 was painted with white enamel. A notch
filed in the shell lip was used in the determination of growth in
class 4 (Figure 2). All measurements were made with vernier
calipers accurate to 0.1 mm. The data on the four size classes of
snails are presented in figure 3.
It may be taken from figure 3 that as the snails increase in size,
a smaller percentage increase and a larger actual increase in length
Fig. 1. Growth cage situated in salt marsh. A — wooden stake; B — plastic
screen cage; C — specimens; D — marsh floor. Fig. 2. Recaptured specimen from
class 4. A — shell added during study; B — filed notch.
Contribution No. 1475 from the University of Miami, Rosenstiel School of
Marine and Atmospheric Science.
April, 1972
NAUTILUS
137
is effected. See figures 4 and 5 for graphic representations.
The present study was carried out entirely during the warmer
months of the year and so no data are available on growth during
the winter. The length of time required for an individual to
approach its maximum size may, then, only be estimated. If the
growth rate in the winter is similar to that in the summer, which
may very well be the case as no distinct growth lines appear on
the shells before maximum size is approached, the growth incre-
ments of size classes 1, 2, and 3 may be combined to form a
hypothetical growth curve. Such a curve (Figure 6) indicates that
the snails could approach maximum size in 20 months. Sexual
maturity is reached before this time as will be discussed in another
paper.
Geographic Variation
Bequaert (1943) wrote of L. irrorata, "It is remarkably uniform
throughout its range, in shape as well as in color."
The inspection of nine groups of 50 specimens each taken from
Corpus Christi, Texas, to Beaufort, North Carolina, confirms
Bequaert's statement, as no differences in color, and only minor
ones in shell shape and number of body whorl ridges were noted
from area to area (Figure 7).
Sexual Dimorphism
Several hundred large specimens were collected in the salt marsh
and measured with vernier calipers for shell length. Fifty-nine
specimens measured between 19.1 and 19.5 mm. Of these, 33 or
Fig. 3. Growth in shell length of L. irrorata.
138
NAUTILUS
Vol. 85 (4)
-I — I — I — I — I — I — I — I — r-
45 6 7 8 9 10 II H 13 M 15 16 17 18 19 20
Original Shell lenqth (mm)
is I 7 S 9 10 11 13 13 M 15 16 17 18 19 20
Original Shell length (mm)
Fig. 4. Increase in shell length in Fig. 5. Increase in shell length in
L. irrorata (%) March-August. L. irrorata (mm) March- August
Fig. 6. Hypothetical growth cnjrve for L. irrorata.
A — extrapolation; B — increase in length, class 1; C — increase in length, class 2;
D — increase in length, class 3.
April, 1972
NAUTILUS
139
56% were females. Thirty-nine specimens measured between 19.6
and 20.0 mm and 28 or 72% of these were females. Forty-three
specimens measured between 20.1 and 20.5 mm. Of these, 27 or
63% were females. Nine specimens were longer than 20.6 mm. All
of these were females.
Sexual dimorphism of shell size, with the female being larger than
the male, has been observed in several other species of Littorina
(e.g. Moore, 1937, in Littorina littorea (Linne); Linderking, 1951,
in Littorina angulifera (Lamarck); Struhsaker, 1966, in Littorina
pintado (Wood); Littorina picta (Philippi) and Littorina scabra
(Linne)). The size difference is not related to protrandry and is
thought by Moore (1937) and Linderking (1951) to be a conse-
quence of a faster growth rate which they found in females.
Phenotypic Characters
The anterior region of the shell lip of L. irrorata collected in the
salt marsh was noted to possess an indentation (Figure 9) not seen
in other species of Littorina observed by the author, and not men-
Fig. 7. Variation in shell morphology of L. irrorata in southeast Atlantic and
Gulf states.
140
NAUTILUS
Vol. 85 (4)
tioned by Bequaert (1943) in his thorough description of the shell.
On the other hand, a large group of the species not showing the
indentation was found living on limestone boulders near St. Marks,
Florida. Fifty juvenile specimens from this group were collected
and marked with white enamel on the shell lip and shortly there-
after released in the salt marsh. Fifty specimens from the marsh
were marked in the same manner, released on the limestone boul-
ders and subsequently lost during a storm. After five months
growth, 41 of the specimens were recaptured in the marsh and
observed for changes in lip contour. All 41 of the recovered ani-
mals, which five months earlier had possessed no indentation,
possessed one upon recapture (Figure 9). Since specimens of all
sizes in the salt marsh exhibit the indentation and since specimens
not exhibiting the indentation develop one soon after being trans-
ferred to the salt marsh, the indentation of the shell lip is an
environmentally induced character rather than a genetic one.
The portion of the shell lip at which the indentation occurs is
that at which the snail is often attached to the rounded leaves of
the marsh grass, Juncus roemarianus. The mantle is not extended
when the snails are attached to the grass, so no special restriction
on the laying down of new shell exists in the area of attachment.
Abrasion of fragile newly laid down shell at this point by the grass,
therefore, is possibly responsible for the indentation.
Literature Cited
Bequaert, J. C. 1943. The genus Littorina in the Western Atlantic.
Johnsonia 1 (7): 1-27.
Linderking, R. E. 1951. Observations on Littorina angulifera Lam.
from Biscayne Key, Florida. Quart. J. Fla. Acad. Sci. 14: 247-250.
Moore, H. B. 1937. The biology of Littorina littorea. Part 1.
Fig. 8. Dimensions used in length-width ratio computations. L — length;
W — width. Fig. 9. Specimen removed from limestone boulder and placed in
the marsh for five months. A — edge of shell lip when removed from Limestone
Boulder; B — paint; C — shell lip indentation.
April, 1972 nautilus 141
Growth of the shell and tissues, spawning, length of life, and
mortality. J. mar. biol. Ass. U. K. 21: 721-742.
Struhsaker, J .W. 1966. Breeding, spawning, spawning periodicity
and early development in the Hawaiian Littorina: L. pintado
(Wood), L. picta (Philippi) and L. scabra (Linne). Proc. malac.
Soc. Lond. 37: 137-166.
A MOLLUSK NEW TO LAKE BIRKET QARUN, EGYPT
By Kenneth D. Rose
Department of Geology and Geophysics
Yale University, New Haven, Connecticut 06520
Birket Qarun is a large brackish-water lake situated in the Fayum
Depression on the eastern edge of the Sahara Desert, about sixty
miles southwest of Cairo, Egypt. The modern lake is about 25
miles long and four miles wide, but in Pliocene and Pleistocene
times it apparently covered a much greater area. The lake has been
steadily diminishing in size since its formation (probably in the
late Miocene), causing increasing salinity which has resulted in the
current bracking state. Although there is some doubt as to how the
lake was formed, there clearly has been no connection with the sea
in recent geologic time. Birket Qarun is very shallow, nowhere
reaching a depth of more than six meters. Its surface is about 45
meters below sea level.
The molluscan fauna is noticeably limited in diversity, although
not at all in absolute numbers. Blanckenhom (1901) was one of
the first to mention the lake itself. In his review of Egyptian geol-
ogy and paleontology he listed eight mollusk species found there.
E. A. Smith (1908), however, was the first to describe the Birket
Qarun mollusks, listing nine gastropod species and two pelecypods.
His gastropod list included all those published later by Gardner
(1932) except for two species of Planorbis. The two pelecypods
recorded by Smith are both species of Corbicula. All eleven fonns,
as noted by Smith, are typical lacustrine or fluviatile (Nilotic)
forms. Pallary (1909, 1924) surveyed the entire Egyptian mollus-
can fauna, discussing many of the species found in the lake; how-
ever, he found no additions to the fauna.
Less than twenty years after Smith's publication, Gardner (1937)
reported the discovery of two species previously unknown in Birket
142 NAUTILUS Vol. 85 (4)
Qarun. There were the pelecypods Scrobicularia cottardi (Pay-
raudeau) and a variety of Cardium edule Linne. Neither species
had been found in any of the older Fayum lake beds, or in any of
the recent lakes or streams of the area. Even more significantly,
neither species had been recorded by Blanckenhorn or Smith. Con-
sidering the abundance of these two forms, especially Cardium
edule, it is very improbable that they could have been overlooked
by the earlier workers. Both are brackish Mediterranean forms,
and the only suitable explanation for their sudden appearance is
that they were introduced to the lake as eggs or tiny young carried
in mud on the feet of shore birds. Blanckenhorn was the first to
suggest this method of introduction (in reference to the gastropod
Hydrobia peraudieri Bourguignat), and Gardner (1927, 1932) sup-
ported this hypothesis.
The most comprehensive study of the Fayum mollusks (and the
most recent known to this author) was published by Gardner in
1932. Fifteen species, including four pelecypods and eleven gastro-
pods, were reported from Birket Qarun (those preceded by an
asterisk were also recorded by E. A. Smith in 1908):
Cardium edule Linne, 1758
*Corbicula africana Krauss, 1848
*Corbicula consobrirw, (Cailliaud), 1827
Scrobicularia cottardi Payraudeau, 1826
*Bulinus truncatus (Audouin), 1827
*Cleopatra bulimoides (Oliver), 1804
*Hydrobia peraudieri Bourguignat, 1862
*Lanistes carinatus (Olivier), 1804
*Melanoides tuberculatus (Miiller), 1774
*Planorbis ehrenbergi Beck, 1837
Planorbis mareoticus (Letourneux), 1884
Planorbis planorbis (Linne), 1758
*Theodoxus niloticus (Reeve), 1841
*Valvata ni/oiica Jickeli, 1874
*Viviparus unicolor (Oliver), 1804
Recently (November 1968 through January 1969), this writer
had the opportunity to study the molluscan fauna of Birket Qarun.
My survey resulted in the new record of Pirenella conica (Blain-
April, 1972 nautilus 143
ville, 1829), (Potamididae, Cerithiacea), a form which was con-
spicuously absent from previous faunal lists. This species is now
the commonest gastropod in the lake. Together with Cardium edule
and Hydrobia peraudieri, this snail forms thick shell banks all along
the southern shore of Birket Qarun. Pirenella conica, a typical
Mediterranean brackish form, has been discovered in many north
African brackish lakes. Like the two species discovered by Gardner
(1927), is it highly doubtful that it could have been overlooked by
previous students; therefore, it must have been introduced since the
last faunal study (1932), also possibly by birds.
It is significant to note that Cardium edule is at least as successful
today as when Gardner first recorded its presence. Furthermore, it
is by far the commonest molluscan species in the lake. All speci-
mens observed were noticeably small and thin-shelled as compared
to typical Mediterranean specimens. This would be expected in an
environment of low salinity. Many individuals were pale-yellow
in color.
References Cited
Blanckenhorn, M. 1901. Neues zur Geologic und Palaontologie
Aegyptens. Zeitsch. Deutschen Geol. GeselL, vol. 53, pp. 307-502
(p. 463).
Gardner, E. W. 1927. The Recent Geology of the Northern Fayum
Desert. The Geological Magazine, vol. 64, no. IX, pp. 386-410.
Gardner, E. W. 1932. Some Lacustrine Mollusca from the Faiyum
Depression: A Study in Variation. Mem. Inst, d'^gypte, vol. 18,
pp. 1-123, pis. 1-8.
Pallary, Paul. 1909. Catalogue de la Faune Malacologique de
I'figypte. Mem. Inst. d'Egypte, vol. 6.
Pallary, Paul. 1924. Supplement a la Faune Malacologique Ter-
restre et Fluviatile de I'figypte. Mem. Inst. d'Egypte, vol. 7, fasc.
I, pp. 1-61, pis. 1-4.
Smith, E. A. 1908. On the Mollusca of Birket el Qarun, Egypt.
Proc. Malac. Soc. London, vol. 8, pp. 9-11.
144 NAUTILUS Vol. 85(4)
NOTES
Records of Introduced Mollusks: New Mexico and Western
Texas — ^The mollusks reported are judged to have become estab-
lished as a result of introductions by man. Specimens have been
placed in collections of the U.S. National Museum. Anodonta
grandis Say. (USNM 681635 — cataloged as A. stewartiana Lea).
This mussel was collected in a pond in the western part of the city
of El Paso, El Paso County, Texas, in 1968 and 1971. The pond
is locally known as "Cement Lake" and belongs to the Southwestern
Portland Cement Company. It was constructed in 1910. Although
water is sometimes pumped into the pond from the Rio Grande,
establishment of the mussels from this source seems unlikely because
( 1 ) nearest records known from the Rio Grande drainage are about
700 miles to the southeast in the lowermost part of the system, (2)
usually the Rio Grande is dry or merely a trickle between El Paso
and Presidio, Texas, seemingly precluding establishment of perma-
nent populations of unionid mussels in that section of the river and
(3) no unionids of any kind, either fossil or living, have been seen
by us or have been reported from the Rio Grande drainage above
Presidio. We conclude, therefore, that this population has probably
resulted from stocking with fish that bore the glochidia. According
to Mr. O. P. Kroeger, Plant Superintendent (written comm., Feb.
22, 1971) "During the 1930's the lake was stocked with fish: Cat,
Bass, Perch, etc., by a local group as a private fishing club." Taylor
(1966. Veliger, 8:198) suggested that occurrence of an Anodonta
in a lake in Arizona had resulted from glochidia carried by fishes.
He noted that in western states ". . . this is the only known case in
which a mussel has been introduced along with the fishes."
Specimens have low umbones and a slightly sinuous hinge line,
lacking the inflated umbonal area and the straighter hinge line of
Anodonta stewartiana Lea, which occurs to the east in central Texas.
Corbicula fluminea (Miiller). Identification is by Dr. J. P. E.
Morrison, U.S. National Museum. This clam was reported as
Corbicula manilensis Philippi by Metcalf (1966. The Nautilus,
80: 16-20), At that time the clam was known only from two irriga-
tion drainage ditches in the Mesilla Valley of Doiia Ana County,
New Mexico, and El Paso County, Texas. In subsequent years it
has spread northward into Sierra County, New Mexico, where it
occurs in the Rio Grande and in two of its impoundments, Elephant
April, 1972 nautilus 145
Butte and Caballo Reservoirs. Seemingly dispersal of the clam into
these waters has taken place in the years 1966-1970, as it was not
observed there during surveys made between 1963 and 1965.
(USNM 706608, Elephant Butte Reservoir; USNM 706607, Caballo
Reservoir; USNM 706606, Rio Grande at Percha Diversion Dam,
5 mi. S Caballo Reservoir Dam).
Radix auricularia (L.) s.l. Specimens of this Palearctic lymnaeid
snail have been taken in Lake Roberts on the Gila River, Grant
County, New Mexico, by Carl Olsen in 1966 (USNM 706609) and
from a ranch pond, located eight miles southeast of Deming, Luna
County, New Mexico, by Kay S. Samson and G. L Wilson in 1970
(USNM 706610). USNM specimens are cataloged as Radix
japonica Say. — Artie L. Metcalf and Richard Smartt, Department
of Biology, The University of Texas at El Paso, El Paso, Texas
79968.
CORBICULA MANILENSIS (PhILIPPI)iN OKLAHOMA. This SpCCicS
has been found recently in Lake Overholser near Bethany, Okla-
homa City. The specimens were collected by Mrs. Louise Cassel on
August 25, 1969, and she stated in her letter that "the little clam
variety seems to be new to Lake Overholser." — -William J. Clench,
26 Rowena St., Dorchester, Mass. 02124.
The Type Locauty of Stagnicola montanensis (Baker) 1913
— This species was described from "Hayes Creek, near Ward, Mon-
tana, altitude 3825 feet. . . . collected by Mr. L. E. Daniels in April,
1912" (F. C. Baker, 1913, The Nautilus, 27.- 115-116). No town of
Ward now exists, but by making local inquiry it was learned that
shortly after the turn of the century (circa 1910) there was a post
office by that name seven miles south of Hamilton, Ravalli County,
Montana. The old brick building is still standing approximately
one-half mile north of the Jim Ward Ranch on Hayes Creek. This
ranch (at an elevation of 3780 feet) is located at the middle of the
south 1/2 of sec. 3, T.4N., R.21W. (U.S.G.S. Topographic Sheet
N4607.5, Hamilton South Quadrangle, 1:24,000, 1964). Taylor,
et al. (1962, Malacologia, 1(2) : 240) were correct in suggesting that
this area most likely represents Daniels' type locality for this snail.
Stagnicola montanensis has been found in abundance in the boggy,
spring-seepage areas on both sides of Hayes Creek near and above
this ranch. — Richard H. Russell, Department of Biological Sciences,
University of Arizona, Tucson, Arizona 85721.
146 NAUTILUS Vol. 85 (4)
Reproduction of Amblema costata (Rafinesque) in Moose
River, Minnesota — Dennis 1971:86) states that Amblema costata
(Rafinesque) "is often found in coarse gravel in strong currents
seldom in silt or mud." Many specimens of this species were found
in Moose River, Minnesota near the town of Sturgeon Lake. Large
adults, at least 15 years of age, were found in a strictly soft, thick,
mud substrate. This rather different habitat for the species from
that indicated by Dennis might be explained by host-fish dispersal
patterns. Such an explanation is encouraged by the following obser-
vations which indicate that the Moose River population depends on
very occasional good year-classes of shed glochidial mussel larvae
from fish: On November 3, 1969, 31 juvenile Amblema costata
were collected with a scraper net with a mesh fine enough to
include one-year-old mussels. All of these specimens were 2-year-
old Amblema costata. There were no younger or older Amblema
costata (other than tlie adults). A year later a similar collection
turned up only 3-year-old Amblema costata juveniles. Juveniles of
other species in the same bottom varied in age from 1-6 or more
years, thus indicating that the chance dispersal of host-specific fish
rather than change in bottom conditions from year to year was
responsible for the presence of just one year-class of Amblema
costata. All of the juveniles collected were donated to the Ohio
State Museum, Columbus, Ohio for investigators who wish to
examine this problem in further detail. — Marc J. Imlay, National
Water Quality Lab., Environmental Protection Agency, Duluth,
Minn. 55804.
Literature Cited
Dennis, Sally D. 1971. Pennsylvania Mussel Studies. Final report.
Center for Aquatic Biology, Dept. of Biol., Eastern Michigan
University, Ypsilanti, Michigan. 138 p.
NEWS
The American Malacological Union's 38th annual meeting
will be held in Galveston, Texas, from July 9 through July 14, 1972.
President Arthur Merrill promises an interesting meeting that will
be hosted by the Galveston Shell Club. AMU members will be
sent registration information in a few weeks. Others may obtain
details from Mrs. Katy W. Fowler, Titus Harris Clinic, 200 Uni-
versity Boulevard, Galveston, Texas 77550.
April, 1972 nautilus iii
E>R. John Teng-Chien Yen, paleontologist, died in Villanova,
Pa., on February 4, 1972, at the age of 68. He had suffered a stroke
a few years previously. He was born in Canton, China, on Feb-
ruary 15, 1903. He received his B.Sc. at the National University of
Nanking, and his Ph.D. from Berlin University in 1939. He pub-
lished on Chinese mollusks and on Mesozoic and Tertiary inverte-
brates. Dr. Yen taught geology at Villanova University, Pa., from
1956 to 1966.
The Collection of Karl Mayer-Eymar. — Karl Mayer-Eymar,
stratigrapher and paleontologist, published numerous papers, mainly
on fossil mollusks, from 1853 until 1906. Many of his articles are
based on material he collected himself during his extensive travels
through Europe and Egypt, or on material that was sent to him
for identification and age assignment. Thus Mayer-Eymar assem-
bled large collections of Tertiary mollusks during his long career.
He is the author of several stage names of the European Tertiary,
and he described more than a thousand new species. Therefore his
collections contain the mollusk faunas upon which those stages
were based and numerous type specimens.
The collections of Mayer-Eymar used to be housed at the Depart-
ment of Geology of the Eidgenossische Technische Hochschule in
Zijrich. It may be of interest to paleontologists dealing with Ter-
tiary mollusks to know that these collections are now deposited at
the Naturhistorisches Museum Basel, by which institution they are
also administered. — Peter Jung, Naturhistorisches Museum, Augus-
tinergasse 2, 4051 Basel, Switzerland.
Dr. Leo G. Hertlein, Curator Emeritus of Geology at the Cali-
fornia Academy of Sciences, died on January 15, 1972, following a
very brief illness. Biographical information was published in vol.
84, no. 2, of The Nautilus (October 1970).
The Hall of Shells, consisting of thirty exhibits on mollusks, at
the Delaware Museum of Natural History, Kennett Pike, route 52,
6 miles north of Wilmington, will open to the public on May 13,
1972.
iv NAUTILUS Vol. 85 (4)
PUBLICATIONS RECEIVED
Carter, J. L. and R. C. Carter. 1970. Bibliography and Index of
North American Carboniferous Brachiopods (1898-1968). Mem-
oir 128, Geol. Soc. Amer., 382 pp. $8.00. G.S.A., Box 1719,
Boulder, Colorado 80302.
Wilson, Barry R. and Keith Gillett. 1971. Australian Shells. 168
pp., 106 pis. in color, 34 text figs. A. H. and A. W. Reed Co.,
Sydney, and Chas. E. Tuttle Co., Rutland, Vermont 05701. U.S.
$21.50. A stunning, popular book on gastropods, featuring identi-
fications and living mollusks. Highly recommended.
Brost, F. B. and R. D. Coale. 1971. A Guide to Shell Collecting in
the Kwajalein Atoll. 157 pp., 33 figs. Common shells of the
Marshall Islands. Chas. E. Tuttle Co., Rutland, Vermont 05701.
Habe, T. 1971. Shells of Japan. 140 pp., color illus. Color Books,
no. 25, Hoikusha Publ. Co., Osaka. Translated by T. I. Elliott.
Useful and attractive little hand guide. Paperback.
Harman, Willard N. and C. O. Berg. 1971. The Freshwater Snails
of Central New York — with illustrated keys to the genera and
species. Search, (Cornell Univ. Agriculture, Ithaca), vol. 1, no.
4, 68 pp. Excellent coverage of 42 species.
Jacobson, M. K. and W. K. Emerson. 1971. Shells from Cape Cod
to Cape May. Paperback, 152 pp. Dover Publ., 180 Varick St.,
N.Y. 10014. $2.00. An up-dated and improved version of "Shells
of the New York City Area" of 1961.
Dance, S. Peter. 1971. Seashells. 159 pp., color illus. Hamlyn
House, London. Excellent introduction to marine shells.
Kuroda, T., Tadashige Habe and Katura Oyama. 1971. The Sea
Shells of Sagami Bay. xvi + 1,279 pp., 121 pis. (105 in color).
Maruzen Co., Tokyo. $58.00. 1,121 species and subspecies are
covered, including 30 new genera and subgenera, 104 new species
and subspecies.
Graham, Alastair. 1971. British Proscobranchs and other Opercu-
late Gastropod Molluscs. 112 pp., 118 drawings. Synopses of the
British Fauna, no. 2. Academic Press, Oak Tree Rd., Palisades,
N.Y. 10964. $4.00. Paperback.
Aiken, D. W. and K. J. Fuller. 1970. The Living Volutes of Africa.
70 pp., line drawings of 31 species. Published by R. E. Petit, P.O.
Box 133, North Myrtle Beach, S.C. 29582. $3.00.
Pownall, Glen. 1971. New Zealand Shells and Shell Fish. 87 pp.,
colored illus. $4.95 N.Z. Seven Seas Publ. Pty., Wellington. 150
species in color; recipes.
Andrews, Jean. 1971. Sea Shells of the Texas Coast, xvii + 298
pp., illus. $17.50. Good for Texas coast; collecting methods; 221
species well illustrated in black-and-white.
MBI. WHO! I IBRARY
UH 17XU Y