Mi
THE
NAUTILUS
THE PILSBRY QUARTERLY
DEVOTED TO THE INTERESTS
OF CONCHOLOGISTS it^^""*^ ^
EDITORS AND PUBLISHERS
HORACE BURRINGTON BAKER
Professor Emeritus of Zoology, University of Pennsylvania
CHARLES B. WURTZ
Consulting Biologists, 1009 Commercial Trust Building
R. TUCKER ABBOTT
H. A. Pilsbry Chair of Malacology, Academy of Natural Sciences
MRS. HORACE B. BAKER
Philadelphia, Pennsylvania
PONY PRINTING, UPPER DARBY, PA.
April, 1960 nautilus iii
CONTENTS
Names of new genera, species, etc. in italics
Alaska 14
American Malacological Union 74, 160
Arion ater 36
Alabama 88
Atlantic, western 11, 23, 28, 39, 47, 76, 79, 103, 118, 119, 139
Beringius eyerdami Allyn G. Smith 5, 43
Berry, Lloyd E., obituary 117
California, marines 1, 43, 51, 65, 95
Calipyrgula circumstriata Leonard & Tong-Yun 125
Calipyrgula pecosensis Leonard & Tong-Yun 110
Campeloma integrum, sinistral 118
Canada 28
Carychium clappi Hubricht, for C. costatum 36
Clione limacina, stranding of 76
Cochliopa riograndensis, radula from Texas 125
Colorado 160
Conus, radulae of Puerto Rican species 119
Dates of the Nautilus 35
Decipifus gracilis McLean 10
Eulotidae 25
Eustomopsis polingi 63, 114
Family names in Pulmonata 114
Florida, inland 28, 37, 76, 137, 161
marines 11, 28, 79, 118
Fruiticicolidae 25
Green shells 63, 76, 1 14
Hatching, Liguus 37
Helminthoglyptidae 25
Honduras, British 129
Indiana 113
Indo-pacific 103
Introduction of unionids 36
of Zachrysia 76
Kentucky 57
Maine 113
Maryland 21, 28, 160
iv NAUTILUS Vol. 73 (Index)
Massachusetts 36, 113
Melampus 28, 47
Melongena corona, feeding & eggs 11, 77
Michigan 36
Mya arenaria on buoys 39
New Guinea 63, 114
New Jersey 28, 113, 133
New York 28
North Dakota 38
Ohio 1 13, 118
Oklahoma 37
Olivella biplicata, variation 65, 95
Oncomelania, production of eggs 147
Oyster populations, sub-tidal 139
Pacific, eastern 1, 9, 43, 51, 65, 95, 103
western 103, 160
Phyllonotus peratus Keen 105
Phyllonotus peratus decoris Keen 107
Pleistocene, inland 110, 125
marines 51, 65
Pliocene marines 65
Puerto Rico 152
Pulmonata, family names 114
Radulae, mounting minute 135
Relaxation of muricids 16
Rhodacmea kahawbensis, anatomy 88
Robertson, Roberts, appointment 160
Sampling, quantitative 131
Sinistral 118
Snails eaten by fishes 14
South Carolina 28, 1 1 3
Succinea pyrites Hubricht, for S. aurea, in part 113
Texas 28, 37, 110, 125
Thysanoteuthis rhombus 23
Truncatella 79
Virginia 21, 113
Watson, Hugh (obituary) 158
West Indies 28, 79
Wheatley collections 72
April, 1960 nautilus v
Wyoming 155
Xanthonychidae 25
Zachrysia auricoma Sc Z. provisoria from Florida 76, (4) iii
Zetek, James (obituary) 159
INDEX TO AUTHORS
Abbott, R. Tucker 160
Atmus, R 37
Baker, H. Burrington 25, 35, 76, 114
Baker Sc Stewart H. Jadis 158
Basch, Paul F 88
Beetle, Dorothy E 155, 160
Blake, John W. (Carriker &) 16
Branson, Branley A. 37
Branson & William R. Heard 14
Burch, Mrs. John Q 118
Carriker, Melbourne Romaine & John W. Blake 16
Chace, E. P 36
Clarke, Arthur H., Jr 36
Clench, William J 76, 114, 161
Dexter, Ralph W 118
Dineen, Clarence F. (Holle &) 28, 46
Erdman, Donald S. (Voss &) 23
Golley, Frank B 1 52
Gray, I. E. (Wells &) 139
Grimm, Wayne 21
Haas, Fritz ^ Alan Solem 129
Heard, William R. (Branson &) 14
Henrard, J, B 63
Holle, Paul A. & Clarence F. Dineen 28, 46
Hubricht, Leslie 36, 1 1 3
Jackson, Ralph W 1 60
Jadis, Stewart H. (Baker &) 158
Johnson, Richard 1 72
Keen, A. Myra 103
Leonard, A. Byron & Tong-Yun Ho 110, 125
McLean, James H 9
Merrill, Arthur S 39
VI
NAUTILUS Vol. 73 (Index)
Post, R. L 38
Rosewater, Joseph 57, 78
Roth, Ariel A. & Edward D. Wagner 147
Siekman, Lula B 118
Smith, Allyn G 1, 43
Solem, Alan (Haas &) 129
Sparks, B. W 158
Stohler, R 66, 95
Teskey, Margaret C 74
Tong-Yung Ho (Leonard &) 110, 125
Torre, Alfredo de la 79
Turner, Ruth D 11, 77, 135
Valentine, James W 31
Voss, Gilbert L. & Donald S. Erdman 23
Wagner, Edward D. (Roth &) 147
Warmke, Germaine L 119
Wells, Harry W. & I. E. Gray 139
Wilson, Druid 137
Wurtz, Charles B 131
REVIEWS
Italics means new taxons included originally
Abbott, R. T, Assiminea 38
Altena, C. O. v.R. Philomycidae & Limacidae (1) iii
Arias, C. S. (See Pain) (1) iii
Augustine, D. L. (See Chernin) 38
Basch, P. F. Gundlachia (2) iii
Carriker, M. R. Urosalpinx (2) iii
Chernin, E., Michaelson & Augustine, Planorbidae & Marisa 38
Clench, W. J. Admiralty 6- Bismarck, Solomons & Bahama
Is 77, 78, (4) iii
Deslandes, N. (See Paraense) 78, (3) iv
Hubendick, B. Ancylidae & Planorbidae (2) iii
Lopes, H. de L. Turhonilla 78
Lopes ^ Sa Cardosa. Solariella 78
Michaelson, Edw. H. (See Chernin) 38
Pain, T., & S. Arias C. Pomacea (1) iii
Paraense, W. L., Sc Deslandes, Taphius 78, (3) iv
April, 1960 nautilus vii
Reynolds, M. B. Pleistocene (4) iii
Riedel, A. Oxychilus (2) iii
Sa Caidosa, Paulo de (See Lopes) 78
Starabogatov, Ja. I. Planorbidae, Gyraulus (i) iii, 78
Stohler, R. Astraea & Macrarene (4) iii
Thompson, Fred G. Lithasiopsis (3) iv
Welch, d'A. A. Achatinella 38
Zilch, A. Euthyneura (3) iii
THE NAUTILUS
Vol. 73 JULY, 1959 No. 1
A NEW BERINGIUS FROM THE PACIFIC NORTHWEST
WITH COMMENTS ON CERTAIN DESCRIBED FORMS
By ALLYN G. SMITHi
During the last year and a half more than a dozen specimens
of a large Beringius have been obtained from Mr. Everett C.
Stiles of Bellingham, Washington; these were taken by trawl
fishermen operating off the Washington coast. Loan of these to
the California Academy of Sciences for study has prompted a
review of available specimens under the names Beringius cre-
bricostatus Dall, B. crebricostatus undatus Dall, and B. kenni-
cotti Dall. These include shells furnished by the U. S. National
Museum, among them the type of B. c. undatus, through the
courtesy of Dr. Harald Rehder; by the Stanford University De-
partment of Mineral Sciences from Dr. Myra Keen; by the San
Diego Society of Natural History from Mr. E. P. Chace; and sev-
eral specimens from the private collections of Messrs. Walter J.
Eyerdam, of Seattle, and John Q. Burch, of Los Angeles.
One obvious problem in studying shells of the genus Beringius
is the relative scarcity of material. One must, at present at least,
draw conclusions on but few specimens, often only a single one
from a given locality. Many are beach-worn or damaged, or are
not sufficiently full grown to show adult characters well. But
even under these difficulties it seems possible to draw some ten-
tative conclusions about certain described forms for the benefit
of others who may desire to pursue the problem further upon
acquisition of more and better material.
The shells now before me, with others seen and studied, pre-
sent a puzzling array. Identification of them with described and
figured species is difficult and can be only tentative in view of
the wide variation in sculptural characters and in the shape
aspects of the shells. There appear to be four more or less dis-
tinct categories represented by Beringius shells with heavy spiral
ribbing as follows:
iResearch Malacologist, California Academy of Sciences.
1
2 NAUTILUS Vol. 73 (1)
1. Fairly long-spired shells with strong spiral ribs and relatively
little or no evidences of transverse ribbing or plications. Num-
bers of major spirals range from 8-14 on the body whorl and
3-6 on the earlier postnuclear whorls. B. crebricostatiis
2. Fairly long-spired shells with more numerous and less strong
spiral ribs and prominent transverse ribs or plications. On the
body whorl the spirals may be continuous over the entire area,
or may be strong only below the periphery and weak or sub-
obsolete above it. Numbers of major spirals are 8 or more on
the postnuclear whorls. The transverse ribs or plications range
in number from 12-15 per whorl. B. kennicotti
3. Shells similar to "2," preceding, but transverse ribbing much
weaker and sometimes obsolete. B. undatus
4. Shells wdth spiral sculpture similar to "2," preceding, but
with shorter spire, more tabulate whorls, extremely deep sutures,
and a more tumid body whorl. B., new species
In all specimens seen the canal is short and, if not worn or
broken, has a series of weaker, closely spaced ribs on it ranging
from 5-9 in number. Although a well-marked fasciole is some-
times present, generally it is weak or absent.
The line of demarcation between B. crebricostatiis and B. ken-
nicotti is fairly distinct; but between B. kennicotti and B. undatus
it is much less so. In fact, with more and better material than
is now available, these latter two species possibly can be merged
into a single variable species. Based on the shells at hand, B.
crebricostatus undatus Dall, 1919, seemingly is wrongly placed
and should stand, at least for the time being, as a separate species
more closely related on sculptural criteria to B. kennicotti than
to B. crebricostatus.
Beringius crebricostatus (Dall), 1887. PI. 1, fig. 1
This is the type species of the genus and has been well figured.
Sculpture consists of heavy, flat-topped, spiral ribs, separated by
deeply channeled interspaces. The ribs are "swaged" at the sum-
mits so as to overhang the interspaces slightly. While the original
description and figure indicate 3 of these revolving ribs occur on
the penultimate and earlier postnuclear whorls, a beach-worn and
damaged specimen in the Stanford Collection (No. 7213) from
Simeonof Island, Alaska, has 6 revolving ribs but is typical other-
wise. Another Stanford shell from Unalaska, the type locality, is
illustrated on plate 1, fig. 1. It has 4 prominent spiral ribs on the
penultimate whorl with a 5th weak one just below the upper
July, 1959 NAUTILUS 3
suture; the earlier whorls have 3 ribs, with a weaker one just
above the lower suture. In this shell, the channeled interspaces
are much wider than the spiral ribs. The approximate number
of spiral ribs on the body whorl is given as 14 in the original
description but on the two Stanford specimens it is 13 and 14,
respectively.
Related to B. crebricostatus is a fine adult specimen collected
alive off British Columbia in 238 fathoms (U.S.N.M. No. 210299,
U.S.F.C. Sta. 2862) . This has 6 spiral ribs on the early post-
nuclear whorls and about 19 on the body whorl and canal. In
this shell, the ribs are much narrower than the channeled inter-
spaces and are less prominent overall than in the typical form.
In B. crebricostatus, evidences of broad transverse ribbing
across the whorls are slight, and this feature seems not to apply
to the species when compared with B. kennicotti. Evidently one
also must allow for considerable variation in the number of spiral
ribs, their relative prominence, and their widths compared with
the corresponding widths of the adjacent channeled interspaces.
In general, the species has a comparatively long-spired shell, the
length of the spire being about equal to or greater than the length
of the aperture and canal. Such measurements should, of course,
make proper allowance for missing nuclear whorls as these are
often lost or damaged.
If the above allowances for sculptural variation are correct,
the range of the species is extended from the Aleutians south
at least to a position off the British Columbia coast. Thus it
overlaps the ranges of other forms to be considered.
"COLUS" PERISCELIDUS Dall, 1891.
The National Museum type lot consists of two fine specimens,
No. 122643, U. S. Fish Commission Station 2842 off Akutan
Islands, Aleutians. As has long been suspected these shells, which
are now before me, have the characters of a diminutive Beringius
belonging to the group of B. crebricostatus. Placement in the
genus Coins is questionable. The two specimens in the type lot
are almost identical in measurements but only one has a perfect
lip. McConnell's drawing in the Proc. U. S. Nat. Mus. 17, 1894,
pi. 27, fig. 6, is an excellent representation. Another somewhat
smaller specimen in the collection of the U. S. Geological Survey
4 NAUTILUS Vol. 73 (1)
(No. D 397) from Lash Bay, Tanaga Island, Andreanof Group,
Aleutians, has the tops of the spiral cords "swaged" so as to
overhang the adjoining channels, as in typical shells of B. crebri-
costatus.
Beringius kennicotti (Dall) , PL 1, figs. 2 and 3.
Two specimens that conform to the general requirements of
this species are in the Stanford University Collection. One (No.
929-1) comes from Petersburg, Alaska; the other (No. 7215, old
no. 929-2) from Kodiak Island, Alaska. These are illustrated on
pi. 1, figs. 2 and 3. Both have about the same numbers of spiral
ribs on the body whorl and canal (23 and 26, respectively); each
has 10 spirals on the earlier whorls. The transverse rib count on
the last three whorls, starting with the body whorl is 12-15-15
for the Petersburg shell and 13-14-13 for the specimen from
Kodiak Island. These transverse ribs or plications are strong
above the periphery of the body whorl but become gradually
weaker below it in both specimens. The major difference be-
tween the two specimens lies in the fact that the spiral ribs on
the body whorl are strong throughout on the shell from Kodiak
Island, whereas on the shell from Petersburg these are extremely
weak from the summit of the whorl to a point just below the
periphery where they become equal in strength to those on the
Kodiak shell. This difference in sculpture is liable to be con-
fusing unless allowance is made for it in relating single speci-
mens to this particular species.
B. kennicotti incisiis Dall, 1907, has not been seen by the writer
but is believed to be another sculptural variant.
Beringius undatus Dall, 1919. PL 1, figs. 4 & 5, pi. 2, figs. 1 & 2.
The type specimen is figured here for the first time on pi. 1,
figs. 4 and 5. It is U.S.N. M. No. 223031 and comes from 160
fathoms, mud, off Cygnet Inlet, Boca de Quadra, southeast
Alaska (U.S.F.C. Sta. 4224) . As stated, it is a relatively young
specimen and contains the operculum. The number of transverse
plications on the body whorl is 17, which is larger than average,
the range in other specimens studied being 12-14.
The transverse plications in some specimens of B. undatus are
relatively weak and difficult to count with accuracy. Because
these do not appear to be a major sculptural feature of the spe-
July, 1959 NAUTILUS 5
cies there seems good reason to make a generic distinction be-
tween it and B. kennicotti. Considering the possible limits of
sculptural variation in shells of the genus, however, the relation-
ship between the two nominal species is admittedly close and
we may be dealing with a single polymorphic species. Specimens
studied, that are considered to belong to B. undatus, are as
follo^vs:
1. A single specimen dredged to the north of Unimak Island,
Aleutians, in 41 fathoms, sand (U.S.N. M. No. 122718, U.S.F.C.
Sta. 3259) . This is a fairly large, heavy-textured, thick-lipped
shell 115 mm. long, taken alive. Spiral ribbing is strong through-
out, there being 21 on the body whorl and seven each on the
two preceding whorls. Transverse plications are weak, the num-
bers on the last three whorls being 11-12-13, respectively,
starting with the body whorl.
2. The type specimen from Boca de Quadra, southeast Alaska,
and another smaller specimen from the same general locality and
depth (U.S.N.M. No. 222589, U.S.F.C. Sta. 4225) .
3. A single large, full grown specimen, 143 mm. long, from
off Masset, British Columbia, in the Stanford Collection (No.
7214). This shell is illustrated on pi. 2, figs. 1 and 2. Numbers
of spiral ribs on the last three whorls are 18-9-9, starting with
the body whorl. The canal has a series of 9 ribs. The channeled
interspaces are not as deeply cut as in most of the other shells
of this species that have been studied. Transverse plications are
fewer than average and number 12-12-11 on the last 3 whorls.
The shell is well preserved but not collected alive.
4. Two specimens in the Stanford Collection (No. 7214) from
off the San Juan Islands, Puget Sound, Washington, in 25-30
fathoms. These are labeled "B. crebricostatus var. undatus Dall
(1919)" in Ball's handwriting. One is an imperfect, full grown
"dead" shell; the other is a young one with the nuclear tip
complete.
5. A single young shell, dredged off Flat Point, Lopez Island,
Puget Sound, in the California Academy of Sciences Paleo. Col-
lection (No. 34789), illustrated on pi. 2, fig. 6.
Beringius eyerdami, new species PI. 2, figs. 3, 4; PI. 3, figs. 1-4.
The specimens from Messrs. Stiles and Eyerdam differ mark-
edly from other species with heavy spiral ribbing and seem
worthy of a new name.
Holotype: Shell an adult specimen, large (about 41/2 inches
long) , of fairly heavy texture, globose, creamy-white, covered in
places with a thin, golden-brown periostracum. Nuclear whorls
missing; postnuclear whorls about 41/2, well-rounded, tabulate
6 NAUTILUS Vol. 73 (1)
at the summits, with fairly deep sutures. Body whorl tumid, con-
stricted at the base to form a short but wide, relatively straight
siphonal canal. Axial sculpture faint, consisting of a series of
broad, low irregular, widely-spaced, undulating plications, more
evident on the upper portions of the postnuclear whorls but
fading out almost entirely on the body whorl. Spiral sculpture
prominent and strong over all postnuclear whorls, consisting on
the first of 6, on the next of 7, and on the body whorl of 15
heavy, square-cut, revolving ribs bounded by narrower but deep,
square-cut channels that are concave at their bottoms; both
spiral ribs and their interspaces widen gradually as they approach
the outer lip of the aperture, which is crenulated inside to cor-
respond with the heavy outside spiral sculpture. Outside of canal
worn, devoid of ribbing in the holotype. Over-all microsculpture
consisting of close, irregularly placed, growth riblets that con-
tinue over the spiral ribs and into the channeled interspaces.
Major spiral ribs on the body whorl generally cut by one, cen-
trally placed, weak, incised line and sometimes by more still
weaker ones. Aperture capacious, subovate, of a pinkish color
inside; outer lip thick, not flaring; inner lip appressed, consist-
ing of a heavy wash of callus; columella slightly curved. Canal
wide, short and relatively straight. Just inside the mouth of the
shell on the upper side of the inner lip is a broad, low, rounded,
raised area or boss. Operculum thick and coarse, normal for the
genus. Animal (in alcohol) a female, the upper portion of the
mantle yellowish-white, lightly dusted with irregular, light gray
maculations; edge of mantle incised and ribbed corresponding
with the outside sculpture of the shell, the ribs marked with red-
brown for a distance of about 5 mm. back of the edge. Dimen-
sions of shell in mm.: length, 113.9; maximum diameter, 74.8;
maximum width of aperture, 37.7; length of aperture and canal,
75.0; length of canal alone, approximately 23.5 mm.; number of
postnuclear whorls, 41/2-
Locality and disposition of specimens: Dredged at various
times in 1957 and 1958 by the trawlers "Cooledge II," "Karen,"
"Northern Light," and "Paul L." in approximately 100 fathoms
on La Perouse Bank about 40 miles off Cape Flattery opposite
the entrance to the Strait of San Juan de Fuca, Washington.
Holotype in the Calif. Acad. Sci. Paleo. Type Coll. (C.A.S. no.
36318). A total of 14 paratypes from the same general locality
have been designated as follows: 2 in Calif. Acad. Sci. Paleo.
Type Coll.; 8 in the collection of Mr. E. C. Stiles; one in the
collection of Mr, Walter J. Eyerdam; 3 in the collection of Mr.
John Q. Burch; and one in the collection of the San Diego So-
ciety of Natural History.
Geographical Range: Chignik, Alaska (Norberg) to the coast
of Washington in about 100 fms. off the Strait of San Juan de
July, 1959 NAUTILUS 7
Fuca (Stiles) ; Puget Sound, Washington (Eyerdam) .
Remarks: Because so many specimens of a large, deepwater
Beringius are available for study at one time, somewhat detailed
comments on the range in variation of the shells seems pertinent.
The total number at hand or otherwise known is as follows:
The holotype and 14 paratypes, from the type locality, ob-
tained by Mr. Stiles. The paratypes include 3 kindly loaned by
Mr. and Mrs. Burch and one supplied by Mr. Eyerdam, who
also furnished the holotype. An additional specimen (topotype)
is stated to be in the collection of Mrs. Elizabeth Phelps, Delray
Beach, Florida, has not been available for study.
A single shell (paratype) in the museum of the San Diego
Society of Natural History (No. 12917) from off Vancouver Is-
land, British Columbia.
Two shells obtained by Mr. Ingvar Norberg at Chignik, Alaska,
one of which is in Mr. Eyerdam's collection and the other said
to be in the Tromsoe Museum, Norway. The latter of these has
not been studied.
A single shell collected by Mr. Eyerdam during seine opera-
tions for herring in 1945 at Raspberry Island, Kodiak Island
Group, Alaska.
A single full-grown but "dead," worn specimen collected by
Mr. Eyerdam in Puget Sound at Restoration Point, Bainbridge
Island, Kitsap Co., Wash.
Comments on these follow in the order listed.
Of the 15 shells from La Perouse Bank obtained by Mr. Stiles,
which are at hand, 9 are fine specimens taken alive and have
opercula. Two, including the holotype, have the animals pre-
served in alcohol; both are females. The remaining 6 are "dead"
shells, somewhat worn; one of these is a poorly preserved, half-
grown specimen. Thirteen of these shells have the heavy spiral
sculpture described for the holotype and are remarkably constant
in this outstanding feature, although there are minor variations.
Several have all, or nearly all of the smooth nuclear whorls
present, which may number from 2 to 21/2; they are all some-
what worn, however. Where these are broken off the animal
plugs the open hole with shell material, a condition present in
the holotype. In several of the better preserved shells, the spiral
ribbing continues over the outside of the canal but in diminish-
ing strength, the canal being sculptured with a series of about
9 of these weaker ribs, which are closely spaced and bordered
by wider, shallow interspaces. The incised spiral lines on top of
the major ribs vary considerably and are absent on some shells.
8 NAUTILUS Vol. 73 (1)
Although a flaring lip seems not to be a feature of the species,
in two specimens there is a marked recurving toward the outer
edge and in one of these the lip is much thickened and doubled.
Coloi' also varies somewhat. The thin periostracum when present,
is light brown in some specimens, without the golden hue. One
unusually fine fresh shell is a beautiful pinkish-brown color over
all. In such fresh shells the color inside the aperture ranges from
pink toward a light purple, which evidently changes to lighter
flesh or salmon pink with age or exposure to light.
The other two shells in the type lot deviate remarkably from
the normal heavily ribbed form. On the the most striking one
of these, a "dead" shell illustrated on pi. 3, fig. 2, there is no
spiral ribbing v*^hatever on the upper whorls; toward the base of
the body whorl, however, a series of four strong ribs bordered
by quite narrow channels appears abruptly; the usual series of
weaker ribs occurs on the outer canal. Axial sculpture consists
of broad, very irregular undulations of varying prominence.
Over-all microsculpture appears as the usual closely set growth
riblets but these are finely beaded in an axial rather than in a
spiral direction. The shell is shorter than normal, with a stubby
spire, the summits of the tabulate whorls descending into the
sutures, a feature not exhibited in the normal heavily ribbed
specimens.
While there is a possibility that this single specimen may be
a pathologic variant, the other of the two with aberrant sculp-
ture and other aspects seems perfectly normal. It was collected
alive and has the longer spire of the heavily ribbed form from
the area; it is illustrated on pi. 3, fig. 3. The heavy spiral ribs are
only 7 in number and begin abruptly well below the periphery
of the body whorl, with the usual series of weaker ribs on the
outer canal. The upper part of the body whorl is sculptured with
6 or 7 weak spiral cords, widely spaced, and bordered by wide,
shallow interspaces. At the summit of this whorl is a maze of
similarly weak cording, which originates at the suture and extends
diagonally to the area where the spiral cords begin. (This fea-
ture is suggested also in the figure of B. marshalli Dall, 1919, in
Bull. 112, U. S. National Museum, p. 91, pi. 9, fig. 3.) The sum-
mits of the whorls also show a descending tendency into the
sutures. Microsculpture of this shell has been lost due to unfor-
tunate treatment with acid in cleaning it.
NAUTILUS 73 (1)
PLATE 1
1, Beri72giiis crehricostatus (Ball) . 2, 3, B. kennicotti (Ball) . 4-6, B. imdatus
Ball.
NAUTILUS 73 (1)
PLATE 2
1, 2. Beritigius luidatus Dall. 3, 4, B. eyerdami A. G. Smith, type.
NAUTILUS 73 (1)
PLATE 3
1-4, Beringins eyerdami A. G. Smith.
NAUTILUS 73 (1)
PLATE 4
Decipifiis gracilis McLean 1. (left) Holotvpe, Stanford Univ. Paleo. Type
Coll., no. 8081, X 5. 2. Faraiype, Stanford Univ. Paleo. Type Coll.. no. 8082,
X 5. Off Guavmas, west Mexico.
July, 1959 NAUTILUS 9
List of figures
Plate 1.
1. Beringius crebricostatus (Dall) , Simeonof Id., Alaska. Stan-
ford Univ. Coll. No. 7213. Length, 102 mm.
2. B. kennicotti (Dall) , Petersburg, Alaska. Stanford Univ.
Coll. No. 939-1. Length, 99.1 mm.
3. B. kennicotti (Dall), Kukak Bay, Kodiak, Id., Alaska. Stan-
ford Univ. Coll. No. 7215 (old No. 929-2) . Length, 94.6 mm.
4. 5. B. undatus Dall. Type, from 160 fms., mud bottom, off
Cygnet Inlet, Boca de Quadra, Alaska (U.S.F.C. Sta. 4224).
U. S. National Museum Coll. No. 223031. Length, 79 mm.
6. B. undatus Dall, dredged off Flat Pt., Lopez Id., Puget
Sound, Wash. Calif. Acad. Sci. Coll. No. 34789. Length,
92.3 mm.
Plate 2.
1, 2. Beringius undatus Dall, dredged off Masset, Br. Colum-
bia. Stanford Univ. Coll. No. 7214. Length, 142.8 mm.
3, 4. B. eyerdami, A. G. Smith, new species. Holotype, from
approx. 100 fms., La Perouse Bank, 40 mi. off Cape Flat-
tery, Wash. Calif. Acad. Sci. Paleo Type Coll. (C.A.S. No.
36318). Length, 113.9 mm.
Plate 3.
1. Beringius eyerdami A. G. Smith, new species. Paratype in
W. J. Eyerdam Coll. Length, 120.6 mm.
2. Same. Paratype in Calif. Acad. Sci. Coll. No. 36318. Aber-
rant specimen with obsolete spiral sculpture. Length, 99.9
mm.
3. Same. Paratype in Calif. Acad. Sci. Coll. No. 36318. Aber-
rant specimen with spiral sculpture stronger. Length, 117.2
mm.
4. Same. Raspberry Id., Alaska (Eyerdam) . Calif. Acad. Sci.
Coll. No. 36319. Length, 132.8 mm.
Note: Front views taken with axis of shell horizontal. Back
views with shells resting on their apertures, with axes slightly
tipped upward so as to show sutural characters more clearly.
(To be continued)
A NEW MARINE GASTROPOD FROM WEST MEXICO
By JAMES H. McLEAN
Stanford University
A new gastropod, which fits into the genus Decipifus, recently
proposed by Olsson and McGinty, was collected by the author
in December, 1958, at Guaymas, Mexico. Decipifus belongs to the
family Columbellidae.
"x^
\ ^ '"I f
vx
■v^y.
10 NAUTILUS Vol. 73 (1)
Decipifus Olsson and McGinty, 1958.
Bulls, of Amer. Paleont., vol. 39, no. 177, p. 36. Type species
(monotypy), Decipifus sixaolus Olsson and McGinty, 1958. East
Panama.
Original description: "Shell quite small, Phos-Vike in shape
and sculpture. Protoconch is relatively large, subcylindrical,
formed of 1 to li^ smooth whorls, the final section high and the
apical tip inrolled. Sculpture of the mature whorls is formed by
low, narrow riblets finely beaded by spirals. Aperture semi-
elliptical with a small, indistinct canal at the suture; lip simple;
columella straight, the tip of the pillar slightly twisted; no ex-
ternal fasciole."
Decipifus gracilis, new species. Plate 4
Shell small, fusiform, aperture subovate, 3/g of length. Sculp-
ture consisting of 7 low, flat-surfaced spiral cords between su-
tures, 13 to 15 cords on base, crossed by 13 to 16 somewhat
sinuous axial ribs, (13 on early whorls) , the axial ribs becoming
obsolete on base; entire surface with minute axial striae. Siphonal
notch deep, columella smooth, outer lip simple but reflecting
spiral cords. Ground color buff, with variegated dark brown and
blue-green mottling, brown on the 4 upper cords of each whorl
and on the pillar, many of the bead surfaces white.
Dimensions: Holotype, length 8.2 mm., diameter, 3.7 mm. Para-
type, length 8.0 mm., diameter, 3.5 mm.
Type locality: Bocochibampo Bay, Guaymas, Sonora, Mexico,
about 50 feet offshore from the north end of the bay, in six feet
of water, under rocks. Holotype and paratype collected by the
author, December, 1958.
Repositories: Holotype, Stanford University Paleontological
Type Collection, No. 8081; paratype, No. 8082.
This species appears to be distinct from Amphissa lyrta Baker,
Hanna and Strong (1938) which probably belongs to this genus
also, in that Decipifus gracilis is more slender, has deeper sutures,
7 cords rather than 6 between sutures, 13 to 15 cords on the base
rather than 10, and shows the blue-green mottling. Decipifus
lyrta (Baker, Hanna and Strong) is known from Isla Partida and
other islands near southern Baja California. The two west Ameri-
can species of Decipifus appear to differ significantly from the
four species of Amphissa examined in the Stanford collection.
Although the overall shape and sculpture is similar, the beading
of Decipifus is more pronounced, there is less inner lip callus, and
the two species are much smaller than the minimum size of
Amphissa (9 mm. compared to 13 mm.). The southernmost range
July, 1959 NAUTILUS 11
of Amphissa is Cedros Island, {A, versicolor Dall) , according to
Grant and Gale, 1931. Thus Decipifus and Amphissa are also
geographically distinct.
The types of D. gracilis examined may be immature speci-
mens, for the outer lips are thin and sharp. This may also ac-
count for the absence of the slight posterior canal characteristic
of D. sixaolus. Although the specimens were taken alive, the
operculum was not saved. Conceivably populations of D. gracilis
may be of more general occurrence than is suspected, since col-
lecting has not been as extensive in the subtidal as in the inter-
tidal and dredged areas. Diving in the same spot also yielded
specimens of Engina tabogaensis Bartsch, 1931, and Anachis gilva
(Menke, 1847), both unknown north of Mazatlan until recent
months.
Acknowledgmejits: I wish to express my thanks to Dr. A. Myra
Keen, of Stanford University, for her help in the preparation of
this paper and to Mr. Robert Robertson, of Harvard University,
who first noticed the similarity of "Amphissa" lyrta to the type
species of Decipifus. He also has confirmed the placement of
Decipifus in the Columbellidae from a study of the radula of an
east coast species. (Letter to A. M. Keen dated December 30,
1958.)
Literature Cited
Baker, Fred, G. D. Hanna, and A. M. Strong. 1938. Columbel-
lidae from western Mexico. Proc. Calif. Acad. Sci., vol. 23,
no. 16, pp. 245-254, pi. 24.
Strong, A. M. 1938. New species of west American shells. Ibid.,
vol. 23, no. 14, pp. 203-216, pis. 15-16.
Olsson, A. A. and T. L. McGinty. 1958. Recent marine molluscs
from the Caribbean coast of Panama with the description of
some new genera and species. Bulls, of Amer. Paleont., vol.
39, no. 177, pp. 1-58, pis. 1-5.
NOTES ON THE FEEDING OF MELONGENA CORONA
By ruth D. turner
Museum of Comparative Zoology
At the time that W. J. Clench and I were working on the
Melongenidae for Johnsonia, David and Nevada Schmidt sent
us 6 live specimens of Melongena corona (Gmelin) from Punta
Rassa, Florida. They arrived in early March, 1952, and were
12 NAUTILUS Vol. 73 (1)
immediately placed in a salt water aquarium in the laboratory
where 5 of them are still doing very well (January, 1959) . An
account of the care of these mollusks was published in Turtox
News (Turner, 1956) and notes on egg laying and development
in Johnsonia (1956, p. 162). Continued observations on the feed-
ing of these animals have shown that there must be some means
of communication among them. They invariably feed together,
"ganging up" on one clam even though there are plenty of others
to feed upon. Though these snails are usually rather quiet and
crawl only on the bottom of the aquarium, just before feeding
one specimen becomes very active and crawls up on the blocks
of peat or the sides of the aquarium with its siphon fully ex-
tended and actively waving from side to side. Finding a clam,
usually a small Mercenaria mercenaria (Linne) purchased at the
local grocery store, this first specimen will envelop it with its
foot and within a half-hour all the others will be on the victim
aiding the one that began the process. Just how the proboscis is
introduced between the valves, ^ve have not been able to observe.
However, when feeding is well along, one can pick up the snails
and also, hanging from their extended proboscises, the tightly
closed clam. When feeding is finished, the clam shells are gaping
and completely cleaned out. Melongena corona, unlike Thais
lapillus (Linne), which we have also had in our tanks, never kill
more clams than they can eat and consequently never foul the
tank with rotting, uneaten clam meats. Specimens of Thais lapil-
lus do not feed together but each one attacks a different clam.
They were unable to eat Mercenaria but fed readily on Mytilus.
Clench (1947, p. 66) recorded a similar "ganging up" of speci-
mens of Purpura patula (Linne) when feeding on chitons.
Recently a number of badly broken specimens of Mercenaria
mercenaria were placed in the aquarium and though the meats
of these clams were readily available and each Melongena could
have had a "private dining table" they still all fed on a single
specimen. A clam was placed very close to one specimen of Mel-
ongena which immediately became active and within 15 minutes
all the other Melongena were headed toward this specimen,
ignoring broken clams which were much closer to them. Conse-
quently it does not seem possible that the "signal for feeding"'
could have come from the "odor" of clam meat in the water but
rather from the first Melongena which had begun to feed. That
July, 1959 NAUTILUS 13
this is not true of all members of this family is evidenced by
observations made in the field on Melongena corona johnstonei
Clench and Turner. In Little Lagoon, Alabama, we observed
individuals of this subspecies feeding singly on Tagelus divisus
(Spengler) . In the clear water of the lagoon, the Melo?igena
could be detected at some distance and, if they were not moving
about, we usually found, on picking one up, that the probocsis
was extended down into the burrow of a Tagelus. The soft parts
were complete cleaned out, leaving the shells in place. These
clams with their gaping valves, however, have no protection
against the Melongena and a cooperative attack is unnecessary.
Comfort (1957) reviewed all the known data on the life span
of mollusks, listing 133 species for which some data were avail-
able; of these 35 were prosobranchs. This is an appallingly small
number, and consequently we deem worthy of record the fact
that we have had Melongena corona living in our aquarium for
nearly 7 years. The specimens were at least a year old, and con-
sidering their size probably 2 or 3 years old at the time we
received them. None of the specimens has grown more than a
quarter of a whorl since we have had them. One died in August,
1958, but whether this was from old age or some other cause
we were unable to determine.
Although our specimens have produced egg capsules quite
regularly and the young snails have emerged and crawled to the
water line, conditions in our tank were not satisfactory for their
survival. At least 3 females have been observed depositing egg
capsules and they invariably do this at the end of the aquarium
away from the window and in a common area. There is still
much to be learned and it is hoped that this note will encourage
others who are favorably situated to record similar obser^^ations
on these and other mollusks.
References
Clench, W. J. 1947. The genera Purpura and Thais in the western
Atlantic. Johnsonia 2, pp. 61-91, pis. 32-40.
Clench, W. J. and R. D. Turner. 1956. The family Melongenidae
in the western Atlantic. Johnsonia 3, pp. 161-188, pis. 94-109.
Comfort, A. 1957. The duration of life in molluscs. Proc. Malac.
Soc. London 32, pp. 219-241.
Turner, R. D. 1956. Melongena corona Gmelin, an excellent ma-
rine laboratory mollusk. Turtox News 34 (6), pp. 106-108,
pis. 1-2.
14 NAUTILUS Vol. 73 (1)
SNAILS FROM UPPER PENINSULA OF ALASKA
WITH FEEDING HABITS OF BROOKS LAKE FISHES*
By BRANLEY A. BRANSON and WILLIAM R. HEARD
Department of Zoology, Oklahoma State University
During the summer of 1958 (May-September) the junior
author, working as a fishery aide at the U. S. Fish and Wildlife
Service Research Station at Brooks Lake, collected a few land
snails and some rather important data on the malacophagous
habits of several species of fishes. Brooks Lake is situated in the
Katmae National Monument on the Upper Alaskan Peninsula.
The lake is a relatively deep one (maximum 252 feet) but gastro-
pod habitats are mostly restricted to the shallow north end and
to the peripheral ring of its basin lying less than 10 to 25 feet
below the surface. This is the region of effective plant growth,
consisting mostly of a species in the algal genus Chara.
Several specimens of Succinea strigata Pfeiffer were collected
in a peat bog near banks of the Nakntk River in the region of
King Salmon, July 21, 1958. This form is apparently rather
ubiquitously distributed in Alaska (Hanna, 1956 and others) .
Deroceras laeve (Miiller) is a small grayish or blackish slug
with a soot-colored foot. Our specimens are typical of the one
called Agriolimax (= Limax) hyperboreus (Westerlund) by
Dall (1903), but proved to be conspecific with D. laeve by
Walden (1956) . These specimens were "captured" near Brooks
Lake, July 13, 1958, and days following, on the trips of dead-
fall, small-mammal traps baited with peanut butter. (We have
since used peanut butter in bottle tops to attract Oklahoma slugs
with a great deal of success. In addition, several Polygyra tex-
asiana [Moricand] were collected in this manner, as well as
numerous insects.)
The periphyton referred to above in the brief characterization
of Brooks Lake fosters a population of Lymnaea emarginata Say
that is incredibly large in numbers. Associated with this species,
and only slightly less abundant, are Menetus cooperi planospirus
F. C. Baker and Valvata helicoidea Dall. This massive popula-
tion was observed and a few specimens of each species collected
(Aug. 6 and 19, 1958) by using SCUBA, gear. The following
* Contribution #289 from the Zoology Department and the Research Foun-
dation, Oklahoma State University, Stillwater, Oklahoma.
July, 1959 NAUTILUS 15
notes will show that these gastropods obviously are very impor-
tant in the economy of Brooks Lake piscine inhabitants.
During the course of the summer's work experimental gill
nets were utilized to capture 75 to 80 Dolly Varden charrs,
Salvelinus mahna (Walbaum), one lake trout, 5. namaycush
(Walbaum) , several round whitefish, Prosopium cylindraceum
(Pallas), several blackfish, Dallia pectoralis Bean and thousands
of the ninespine stickleback, Pungitius pungitius (Linnaeus) .
Observations were made on the stomach contents of all the
charrs, but only on samples of the other species of fishes. These
data will be analyzed in considerable detail elsewhere and con-
clusions drawn therefrom. Here, we wish, for the purpose of
records, to report the species involved as food for the fishes men-
tioned above.
Menetiis cooperi planospirus was found in the stomachs of the
round whitefish and blackfish in relatively large numbers and in
the Dolly Varden charr less frequently. In large adults of the last
species, this small snail is probably taken secondarily while the
fish is feeding on the following, larger gastropod species.
Many specimens of Lyrnnaea emarginata, strongly resembling
the form reported as randolphi by Dall, were taken from the
Dolly Varden. One medium-sized fish had 82 snails in its stomach
and an 8 lb. charr was found to contain 253 gastropods, some of
them measuring up to 25 mm. in length. Little else wrs found
in the fish's stomach. Prosopiurn, a known mollusk-eater, was also
found to take many individuals of this species. Dallia apparently
takes only an occasional L. emarginata, which, because of the
small size of the fish, is usually tiny.
The round whitefish and Pungitius were the only fish species
observed to have been feeding upon Valvata helicoidea. The
whitefish devours many Valvata, the stickleback relatively few.
The lake trout is not notorious as a mollusk-eating fish. How-
ever, one specimen of L. emarginata and one empty shell (both
valves) of Pisidium idahoense Roper were found in a large fish.
Since this species eats whitefish in some numbers, it may have
ingested the mollusks along with the fish prey.
These data may turn out to be rather important in helping
to explain why there is little predation upon the valuable sockeye
salmon at Brooks Lake by the Dolly Varden charr. Possibly the
16 NAUTILUS Vol. 73 (1)
charr, in the face of such an abundant supply of readily avail-
able food in the form of gastropods, may take fish species only
when such a menu is lacking in the habitat. This, of course, is
an assumption that will have to be verified.
References
Dall, W. H. 1905. Land and freshwater mollusks. Harriman
Alaska Expd. 13:171 pp.
Hanna, G. D. 1956. Land and freshwater mollusks of the Arctic
slope, Alaska. Naut. 7^ (1) :4-10.
Walden, H. W. 1956. Notes on some Siberian slugs, especially on
the type of Limax hyperboreus Westerlund. Arkiv. Zool.
i(? (4/5): 347-365.
A METHOD FOR FULL RELAXATION OF MURICIDS^
By MELBOURNE ROMAINE CARRIKER and JOHN W. BLAKE
Department of Zoology, Chapel Hill, and Institute of Fisheries Research,
Morehead City, University of North Carolina
Abstract. A method is recommended for full relaxation and
killing in an expanded condition of Urosalpinx cinerea, Eupleura
caudata etterae, Thais haemastoma floridana, Ocenebra erinacea,
Nucella lapillus and Polinices duplicatus. Gastropods are made
partially insensible in a solution of 10 ppm. of 1-naphthyl N-
methylcarbamate ("Sevin") in one atmosphere of COg and then
frozen quickly on dry ice.
Current study of the comparative functional morphology of
boring mechanisms in muricid gastropods by the senior author
(Carriker, 1958a, 1958b) disclosed lack of a method for thorough
relaxation of these marine snails. Of the many chemicals em-
ployed in narcotization of gastropods, cocaine is reported (Lo
Bianco 1899) to provide maximal relaxation, but in the muricid
Urosalpinx cinerea it effects only partial expansion of the soft
parts (Carriker 1943) .
In this investigation, we tested a number of narcotics in search
of a method to produce fully expanded prosobranchs (mainly
Muricidae) for detailed anatomical dissection and experimental
surgery with subsequent recovery of the animal. Active, adult
Urosalpinx cinerea (Say) (collected in Beaufort Inlet, North
iThe bulk of this research was performed at the Institute of Fisheries
Research, and was supported by a U. S. Fish & Wildlife Service grant. Fed-
eral Contract No. 14-19-008-2376.
July, 1959 NAUTILUS 17
Carolina) and Eupleura caudata etterae B. B. Baker (the large
ecologic form of Eupleura, kindly provided by T. Carver, M.
Castagna, and G. Griffith, U. S. Fish & Wildlife Service, from
Chincoteague Bay, Virginia) were the principal experimental
animals employed. In addition small numbers of local Thais
haemastorna fioridana Conrad (from Cape Lookout, N. C; iden-
tification kindly verified by Ruth D. Turner) and the naticid
Polinices duplicatus Say (from Beaufort Inlet, N. C.) , and British
muricids Nucella (=Thais) lapillus (Linne), Urosalpinx cinerea
(Say) , and Ocenebra erinacea (Linne), were tested. The British
gastropods were obtained by the senior author through the gen-
erous cooperation of F. S. Russell, P. R. Walne, and G. D. Waugh
during the summer of 1958 while on a trip to the British Isles.
All experimental snails were maintained in laboratory running
sea-water except the British snails which, not tolerating the high
temperature of the laboratory running sea water, had to be
housed in refrigerated containers under aeration. Thus there
was no possibility of introducing the British snails in native
American waters.
Sets of five or more snails per container were subjected to
(a) increasing concentrations of narcotic, each concentration in
a separate container, (b) gradual addition of narcotic to one
container, and (c) combinations of concentrations of narcotics
in (a) showing promise as relaxing agents. Extent of expansion
of pedal and cephalic regions out of the shell, sensitivity of tenta-
cles to touch, and reaction of the snail to dilute formalin were
utilized as indices of depth of narcosis and degree of relaxation.
Mechanical shocks were avoided which might cause snails to
retract before full narcosis was achieved. Solutions of narcotics
were prepared in sand-filtered sea water ranging in salinity from
20 to 37 o/oo, and temperature of the water in experimental
dishes ranged from 22 to 28° C during the course of experi-
mentation.
Tricaine methanesulfonate (M.S. 222) , metycaine hydrochlo-
ride, dibucaine hydrochloride, and a combination of phenobar-
bital and belladonna, caused snails to retract tightly. So did
Collar's (1938) formalin treatment recommended for killing the
refractory opisthobranch Aeolis in an expanded condition. Slow
cooling (Gohar 1938) brought about slow withdrawal, a response
18 NAUTILUS Vol. 73 (1)
related to behavior in temperate regions where Urosalpinx
migrates bottomward and remains closely affixed to hard surfaces
as temperatures drop in the fall (Carriker 1955) .
Cocaine hydrochloride, phenoxetol (CgH^OCH.CH^OH, sim-
ilar to propylene phenoxetol used by Owen, 1955) , chloretone
and menthol (added as crystals) , ethyl carbamate (urethane) ,
chloral hydrate, and MgSO^ effected partial to full extension
and slight to moderate insensibility. However, even the most
effective of these narcotics produced variable unpredictable de-
grees of relaxation, and percentage of well relaxed insensitive
mollusks was relatively small. All narcotized specimens retracted
fully when cut or immersed in dilute formalin. Combination of
the more effective narcotics in their most effective concentrations
(e.g., phenoxetol 0.1%, ethyl carbamate 0.5%, and cocaine
0.05%) did not compound desirable effects of separate narcotics.
Neither aeration of the narcotizing solution (as attempted with
cocaine and phenoxetol) , nor addition of narcotic slowly (phe-
noxetol, formalin, MgSO^, cocaine, and a mixture of MgSO^ and
cocaine) , nor use of snails carefully removed from the shell
enhanced narcosis.
Full expansion of muricids was obtained by immersion in
(a) sea water under one atmosphere of CO^, (b) 10 ppm. (parts
per million) of tetraethyl monothionopyrophosphate in sea water,
and (c) 10 ppm. of 1-naphthyl N-methylcarbamate ("Sevin") in
sea water.2
Passage of CO^ (evolved from dry ice) through an air-tight
chamber with an outlet causes snails immersed in sea water in
the chamber to extend fully in about seven hours. However,
snails relaxed by this method remain slightly sensitive to touch
and retract vigorously when placed in dilute formalin.
Tetraethyl monothionopyrophosphate^, a liquid soluble in sea
2Loosanoff and associates are developing methods for chemical control of
shellfish enemies (Bull. No. 12, 1956, U, S. Fish & Wildlife Service, xMar.
Biol. Lab., Milford, Conn.) and discovered that Sevin and the pyrophosphate
relax certain gastropods, and that Sevin shows promise in controlling gastro-
pod and other enemies of oysters. Loosanoif kindly recommended the use of
these two chemicals, provided the Sevin, and suggested a concentration for
relaxing muricids which our study confirmed.
3This chemical, CP 847, was kindly contributed by the Development De-
partment, Monsanto Chemical Company, St. Louis, Missouri. It is no longer
manufactured.
July, 1959 NAUTILUS 19
water (and extremely toxic to mammals) relaxes muricids fully
in an hour, and they remain extended in it for at least 24 hours.
However considerable sensitivity is retained by snails during the
first 4 hours; this decreases toward the 10th hour when snails are
less sensitive than when in CO^. Use of pyrophosphate in sea
water of salinity of 20 o/oo produces slightly fuller extension of
snails than in a higher salinity, probably as a result of osmotic
effects. Immersion of narcotized snails in dilute formalin pro-
vokes full retraction.
Freshly prepared solutions of 1-naphthyl N-methylcarbamate
("Sevin") * (relatively safe for humans) produce partial relaxa-
tion of muricids in an hour, and full extension in four hours.
Sevin is less than 0.1 percent soluble in water; solutions were
prepared by adding 0.1 g of crystals to 15 ml of acetone and
mixing this stock solution in sea water. Narcotization with Sevin
is more complete than with either CO, or the pyrophosphate.
Immersion of snails relaxed in Sevin in dilute formalin stimu-
lates slow but complete retraction. Combination of the CO,-
Sevin treatments produces more fully extended snails than either
treatment alone. Solutions of Sevin prepared in sea water of
28 o/oo salinity produce better relaxation than in 35 o/oo. De-
nuded snails do not expand as fully as intact snails in the shell.
Freezing, either slowly or rapidly, causes normal muricids to
retract deeply, whereas rapid freezing of snails relaxed in COg,
pyrophosphate, or Sevin (on dry ice; temperature on this drops
at least to — 56°C) for 24 hours or more kills the snails in a
fully expanded state, and they do not retract into the shell when
dropped into preservative or when they are dissected after thaw-
ing. After shorter periods of freezing many snails remain irritable
and do retract when strongly stimulated.
The results of these experiments led to development of the
following method for killing fully relaxed snails:
1. Narcotize snails in a freshly prepared solution of 10 ppm.
of Sevin (1-naphthyl N-methylcarbamate) in filtered sea water
(25 to 30 o/oo), in a depth of fluid at least thrice the height of
the snails, at room temperature, for 1 hour; keep snails out of
reach of each other and on their backs with their feet extended
4This product is sold under the registered trademark "Sevin" and is
available from Union Carbide Corporation, New York City.
20 NAUTILUS Vol. 73 (1)
freely in the water;
2. Transfer snails to fresh Sevin solution in one atmosphere
of CO, at room temperature and leave there for three hours;
3. Remove snails from the narcotic one at a time and place
the ventral or lateral surface of the foot of each quickly against
the surface of a block of dry ice held in a deep freezer or in an
insulated box; then cover the freezing gastropods with chipped
dry ice, insulate the preparation, and leave for 24 hours.
Freshly thawed snails prepared in this way are ideal for de-
tailed anatomical study since organs retain color, texture, and
pliability characteristic of living relaxed tissues, and take aqueous
stains readily. Relaxed snails may be retained in a frozen state
indefinitely, provided they are not allowed to desiccate, or may
be preserved in a solution of 35 percent ethyl alcohol, 0.5 per-
cent formalin, and 5 percent glycerine in distilled water, which
hardens the tissues slightly. The following gastropods have been
relaxed and killed successfully in an expanded condition by the
Sevin-CO^-quick-freeze technic: American Muricidae: Urosalpinx
cinerea, Eupleura caudata etterae, Thais haemastoma floridana;
British Muricidae: Urosalpinx cinerea, Ocenebra erinacea, Nu-
cella lapillus. American Naticidae: Polinices duplicatus.
Living specimens of these species recover from treatment with
either CO^ or Sevin, and although insensibility is not complete,
it it sufficient to permit rapid experimental surgery on superficial
portions of the body.
Literature Cited
Carriker, M. R. 1943. On the structure and function of the
proboscis in the common oyster drill, Urosalpinx cinerea Say.
Jour. Morph. 75:441-506.
Carriker, M. R. 1955. Seasonal vertical movements of oyster drills
(Urosalpinx cinerea) . Proc. Natl. Shellfish. Assoc. ^5:190-198.
Carriker, M. R. 1958a. Additional information on the mechanical-
chemical nature of drilling by the gastropods Urosalpinx and
Eupleura. Bull. Assoc. Southeastern Biol. 5:5 (abstract) .
Carriker, M. R. 1958b. Comparative functional morphology of
the drilling mechanism in Urosalpinx and Eupleura (muri-
cid gastropods) . XVth Interntl. Congr. Zool. (London) , Sect.
IV, Paper 27: 3 pp.
Gohar, H. A. F. 1938. The preservation of contractile marine
animals in an expanded condition. Jour. Mar. Biol. Assoc.
U. K. 22:295-299.
July, 1959 NAUTILUS 21
Lo Bianco, S. 1899. The methods employed at the Naples Zoo-
logical Station for the preservation of marine animals. Bull.
U. S. Natl. Mus. No. 39: 42 pp.
Owen, G. 1955. Use of propylene phenoxetol as a relaxing agent.
Nature 175'AU.
LAND SNAILS FROM MARYLAND AND VIRGINIA
By WAYNE GRIMM
During the years 1957-1958, additional localities for 5 species
of land snails were found in Maryland and Virginia. The collect-
ing stations of these snails are recorded below.
Mesodon appressus (Say) . On July 1, 1957, twelve specimens
of this species were found under debris in a lumberyard at Cam-
bridge, Dorchester Co., Maryland. This is the second record for
M. appressus on the Delmarva, the first being a dead specimen
from the Chester River across from Chestertown, Queen Anne
Co., Maryland (Vanatta) . Doubtless the presence of this southern
Appalachian species in Maryland is due to accidental introduc-
tion, probably from somewhere in Virginia.
Retinella raderi (Dall) . A single living specimen was collected
on July 27, 1958, at the foot of Dan's Mountain, south of Raw-
lings, Allegany Co., Maryland, on Highway 220. It was found in
wet soil under a rock at the base of a large limestone outcropping.
All other snails associated with it were in the leaf litter between
loose rocks or at the base of the outcropping. Although the area
was searched diligently, no additional specimens of this rare
snail were uncovered. Several bags of leaf mould were collected,
but R. raderi was not among the many snails found therein.
Other snails taken at this unusually fine locality were: Steno-
trema jraternum, Mesodon thyroidus, Triodopsis alholahris, T.
tridentata, T. fraudulenta, Haplotrema concaviim, Ventridens
ligera, V. virginicus, Zonitoides arhoreus, Mesomphix cupreus,
Retinella indentata, R. hurringtoni, Hawaiia minuscula, Guppya
sterkii, Helicodiscus parallelus, Angispira alternata, Punctum
vitreum, Gastrocopta pentodon, G. contracta, G. armifera,
Pupoides albilabris, Vallonia perspectiva, Cionella morseana,
and Carychium exile.
Previously, typical R. raderi was found only in "subfossil" state
at Cumberland, Maryland (Howard Shriver, type 107758
22 NAUTILUS Vol. 73 (1)
U.S.N.M., topotypes A.N.S.P. 73888).
Limax marginatus Miill. Records for this introduced slug are
scattered sporadically on the Coastal Plain.
Maryland: Saltmarsh at Galesville, Anne Arundel Co.; Lum-
beryard at Cambridge, Dorchester Co.; Saltmarsh near Kent Nar-
rows at Grasonville, Queen Anne Co. Virginia: South portion of
Chincoteague Island, Accomack Co.
Previously it was reported from Newfoundland, California,
Colorado, and Missouri.
Gastrocopta cristata (Pilsbry and Vanatta). Numerous speci-
mens of this southwestern species were taken from three open,
comparatively unsheltered areas on the flat Delmarva Peninsula.
At one place it was with G. procera (Gould) . In the field it was
easily separated from procera by its larger size, lighter color, and
more cylindrical shape. One look into the aperture was sufficient
to erase any doubts concerning its identity. This snail may have
been introduced on hay from the southwest. Two lots were de-
posited in the United States National Museum.
Maryland: Under stones in dry weedy area behind Sinclair
Station on Route 50 near Skipton, Talbot Co. Under boards in
lumberyard at Cambridge, Dorchester Co. With G. procera in a
dry area near Chicamacomico River at Drawbridge, Dorches-
ter Co.
Vallonia perspectiva Sterki. This diminutive snail was found
three times in the Appalachian region of Maryland and Vir-
ginia — once in the Blue Ridge, once in the Great Valley, and
once in the Ridge - and - Valley province. Records for it are
surprisingly few in the East.
Maryland: Under dry limestone, Conococheague Creek and
Route 40, Washington Co. Rawlings, Allegany Co. (see above,
R. raderi) . Virginia: Hill near Gooney's Creek Overlook between
mileposts 6 and 7, Shenandoah National Park, Warren Co.
I wish to thank Dr. Joseph P. E. Morrison of the United States
National Museum for his help in identifying Gastrocopta cristata.
Literature Used
Pilsbry, H. A. 1948. Land Mollusca of North America (north
of Mexico) . Monogr. 3, Acad. Nat. Sci. Philadelphia, Vol. 1,
part 2, and Vol. 2.
1
July, 1959 NAUTILUS 23
THYSANOTEUTHIS RHOMBUS, LARGE CEPHALOPOD
NEW TO THE WESTERN ATLANTIC^
By gilbert L. \ OSS and DONALD S. ERDMAN
The Marine Laboratory, University of Miami, and
Department of Agriculture and Commerce of Puerto Rico
On November 5, 1958, at 1205 noon, Mr. Fred Gurke, angling
from the charterboat Sea Queen III operated by Capt. Victor
Rivera, landed a blue marlin about two miles north of the
Carib-Hilton Hotel, San Juan, Puerto Rico. On return to the
dock, the fish was weighed in at 383 1/3 pounds, and then, at the
request of one of us (Erdman) the fish's stomach was opened in
order to examine its contents. Much to the surprise of all con-
cerned, the stomach was found to contain a large squid about
3i/2 feet long and weighing about 23 pounds. It was lodged in
the stomach tail first and except for the loss of the skin from the
strong action of the digestive juices of the marlin, was nearly
intact, only the ends of the tentacles missing. After a preliminary
examination, photographs were taken and the squid was turned
over to Erdman for preservation.
The squid was later brought by Erdman to The Marine Lab-
oratory where a more detailed study was undertaken. After a
careful examination, it was found to be a large specimen of the
squid, Thysanoteuthis rhombus Troschel, 1857. This squid has
not, heretofore, been recorded from the Western Atlantic. Its
measurements are given in the following table.
Table 1. Measurements (in mm.) of Puerto Rican specimen:
This is a large species of squid, although not attaining the
enormous size of the giant squid, Architeuthis, or the Humboldt
"current" squid, Ommastrephes gigas. The largest specimen of
iContribution No. 222 from The Marine Laboratory, University ot Miami.
24 NAUTILUS Vol. 73 (1)
Thysanoteuthis recorded had a dorsal mantle length of 800.0 mm
or about 30 inches.
The species is unusual among the squids for the very large
rhomboidal fins nearly as long as the mantle and the peculiar
shape of the cartilages which unite the mantle with the sides of
the funnel. The pen also is unique among the cephalopods in
that the sides of the vane become free lobed on either side of
the rhachis anteriorly.
According to the records available to us, this is the 22nd speci-
men recorded in the literature and the 2nd adult from the
North Atlantic. According to Pfeffer (1912) 11 specimens were
then known of which all but 3 very small juveniles or larvae
were from the Mediterranean. They ranged from 4.2 mm. to
740.0 mm. in mantle length. Pfeffer in 1910 named another spe-
cies of the genus, T. nuchalis, based on a single specimen of 800.0
mm. mantle length captured off Ningpo in the Yellow Sea. Ac-
cording to the Japanese cephaloped authority, Madoka Sasaki
(1929), this species is identical with T. rhombus. He based his
conclusions on 7 specimens collected in Japanese waters and
ranging in mantle length from 180.0 to 685.0 mm.
In 1947, Barnard, in a brief note recorded the stranding of a
specimen at the Cape of Good Hope and in 1956 Rees and Maul
reported a specimen with a mantle length of 350.0 mm., from
Funchal, Madeira, the only previous record from the Atlantic
Ocean.
Thysanoteuthis rhombus is a powerful squid, with heavy mus-
culature, and is a cosmopolitan warm water species. Rees and
Maul reported that the Madeiran specimen was from a school
of about 20 that came close inshore, so they probably are accus-
tomed to schooling. In schools they must be voracious predators.
Sasaki quotes the Japanese fishermen as saying that they may
leap high above the sea surface and one of his specimens was
taken after it leaped into the bunt of a sail and fell to the deck.
Nothing is known of their food habits although probably they
feed upon small fish.
Despite the paucity of records of adult specimens, this is almost
certainly a common pelagic species. A preliminary examination
of the cephalopods collected by the Dana expeditions in the North
Atlantic has revealed numbers of thysanoteuthid larvae and the
July, 1959 NAUTILUS 25
lack of capture of adults is probably due to their size and speed.
References
Barnard, K. H., 1947. Rare squid at the Cape {Thysanoteuthis
rhombus Troschel). Jour. Conch., 23 (11) :286.
Pfeffer, Georg, 1910. Synopsis der oegopsiden Cephalopoden.
Mitteil. Naturh. Mus., 77:145-198.
. 1912. Die Cephalopoden der Plankton-Expedition. Ergebn.
Plankton-Exped., 2:1-815, atlas.
Rees, W. J. and G. E. Maul, 1956. The cephalopods of Madeira.
Bull. Brit. Mus. (Nat. Hist.) Zool., 5 (6) :259-281.
Sasaki, Madoka, 1929. A monograph of the dibranchiate cephalo-
pods of Japan and adjacent waters. Jour. College Agri.
Hokkaido Imper. Univer., 2^:1-357.
Troschel, H., 1857. Bemerkungen iiber die Cephalopoden von
Messina. Arch. Naturgesch. Berlin, 23 Jahrg., 1857: 41-76.
XANTHONYCHIDAE (PULMONATA)
By H. BURRINGTON BAKER
Since the extension of priority to familial names has been
ratified again at the 25th (1958) International Congress of Zool-
ogy, the status of the family term for American helicoids needs
reconsideration. As pointed out 16 years ago (1943), the oldest
name for an American "helicid" is Xanthonychidae (1879),
which originally included only the Mexican genus Xanthonyx
(1867) . The following chronologic synopsis outlines the history
of these and the related familial names (new ones in italics) :
1867. Crosse, H. & Paul Fischer. J. de Conch. 15:212. Xanthonyx.
1879. Strebel, Hermann & G. Pfeffer. Beitr. Kennt. Faun, mex.,
etc. '^:25. Xanthonychidae (misspelled until 1956).
1883. Fischer. Man. de Conch. :469. Xanthonyx between Binneya
8c Helix in Helicidae.
1884. Tryon, G. W./Struct. Syst. Conch. 3:57. Xanthonyx = Bin-
neya in Helicidae.
1890. Moellendorff, O. von. Bericht. Senckenberg. Ges. :226.
Cochlostylidae ("type g." = Helicostyla; cf. 1909) .
1892. Pilsbry, H. A. Naut. (5:69, footnote. Cochlostylidae not ac-
ceptable.
1898a. Mlldff. Abhandl. Ges. Gorlitz 22:97. Eulotidae ("type
g." =: Fruticicola; cf. 1927).
1898b. Pilsbry %c E. G. Vanatta. Proc. ANSP. 50: 229. Copied
Tryon (1884) in Arionidae (Binneyinae) .
1900. Pilsbry. Proc. Malac. Soc. London ^:24. Xanthonyx in "Be-
logona Euadenia" (z= Eulotidae of 1928) .
26
NAUTILUS
Vol. 73 (1)
1909. Ihering, H. von. Wien. Verb. Zool. Bot. Ges. 5P:430. Heli-
costylinae.
1927. Lindholm, W. A. Archiv Molluskenk. 59:120. Fruticicolidae,
1 ("type g." = Brady baena; cf. 1934) .
1928. Hoffmann, H. Klass. Ordn. Tier-Reich., Pulmonata: 1240.
Fruticicolinae, 2 ("type g." z= Trichia) in Helicidae. Eulo-
tidae (p. 1239) with Xanthonyx as of "Unsicher Stellung,"
Cepolinae Helicostylinae, Epiphragmophorinae & Lysinoi-
nae).
1929. Ihering. Abhandl. Archiv Molluskenk. 2: (2) :50. Frutici-
colidae (2). Eulotinae (p. 61) and Cochlostylinae (p. 70) .
1931a. Hesse, Paul, Zoologica i/(i/2):105. Fruticicolidae (2).
1931b. Thiele, Johannes. Handb. Syst. Weichtierk., pt. 2:688.
Fruticicolidae (1) with Helicostylinae and (p. 697) Epi-
phragmophorinae [Xanthonyx p. 698, & Lysinoe).
1934. Pilsbry. Proc. ANSP. 86:1. Cepolidae (Bradybaeninae) .
1937. Rensch, Use. Arch. Naturgesch. 6 (i) :576. Eulotidae. (Also
M. Perrot, 1938, & G. Cherbonnier, 1943) .
1939. Pilsbry. Land Moll. N. A., ANSP. Monogr. 3, vol. 7:24.
Helminthoglyptidae, with Xanthonyx; Cepoliinae 8c Sonorel-
linae (p. 25) Sc Humboldtianinae (p. 26) . Bradybaenidae,
p. 15.
1943. Baker, H. B. Naut. 5^:82, footnote. Xanthonychidae prior.
1948. Dalsum, J. van. Basteria 72:4. Fruticicola (2) .
1952. Webb, Glenn. Gastropodia 7:1. Xanthonychidae adopted.
1953. Hosoi Sc Sone. Eulota (from Zool. Rec, Mollusca:20; p. 96
lists 3 uses, but none of Fruticicola) .
1958. Turner, Ruth D. MCZ. Occas. Papers on Mollusks 2(22) :
153. Fruticicolidae (1).
Figure 1. Duration of names (to 1956). 8 to 5 = decades (1880-1950).
C = Cochlostylidae. E, Fl & F2 = Eulotidae & Fruticicolidae, 1 & 2 (to
last citations in Zoo. Rec.) : H = Helicostylinae. X = Xanthonychidae
(dotted part = synonymy) .
From the above (Cf. fig. 1), Xanthonychidae did drop out of
sight for 64 years (1943), and was not adopted until 73 years
(1952) after its proposal. However, so far as I can make out,
even its "type genus" was not included under an applicable but
subsequent, familial name until 1928, which may be taken to
July, 1959 NAUTILUS 27
mean that Xanthonychidae remained only 15 (or 24) years in
the synonymy oi any of these. Although Xanthonyx shifted after
1879 from Helicidae (1883) to the synonymy (1884) of an arionid
(1898b), its generic status or inclusion in the helicoids has not
been questioned since Pilsbry's (1900) dissections.
The only 2 applicable, family names, which had been in exist-
ence for 50 years before 1952, are Cochlostylidae (1890) and
Eulotidae (1898). These were based, without descriptions, on
synonyms of the Old World genera Helicostyla and Fruticicola
(= Bradybaena) , respectively, and did not include Xanthonyx.
Cochlostylidae was repudiated when 2 years old (1892) and was
submerged by Helicostylinae (1909) 50 years ago, although it
bobbed up again once subsequently (1929) . Eulotidae was
adopted quite generally. After it was rejected (1927), its use by
authorities (1928, 1929 & 1937) continued for at least 16 years,
and its "type genus" apparently has remained in general use
(1953).
The 5th name, Fruticicolidae (1) , was substituted (1927) for
Eulotidae, without mention of any American genus. It was (and
still is) especially confusing, because Fruticicola (2) traditionally
had been used instead of Trichia in what now is a distinct family
(Hygromiidae Tryon, 1866) or a subfamily of Helicidae. In fact,
the next year (1928) and afterwards (1929 & 1931a), Fruticicoli-
dae (2) , with the hygromiid sense (still used in 1948) of its
"type genus," was preferred by the best European students of
helicoids.
In Eulotidae, Hoffmann (1928) included Xanthonyx, as of
"uncertain position," adopted Helicostylinae, and initiated for
other American genera Cepolinae, Epiphragmophorinae and
Lysinoinae (spelling corrected in 1929).
Unfortunately, Thiele (1931b) went back as far as the 1st
(1927) confusing Fruticicolidae. He included in it Helicostylinae
(1909) and also Epiphragmophorinae, in which he put Xan-
thonyx. Fihhry (1939) substituted Bradybaenidae (-inae, 1934) ,
partly to get rid of this confusion, but also because he judged
Fruticicola to be congeneric with Bradybaena, as did Lindholm
(1927). On the other hand. Turner (1958) copied Thiele.
Until his death. Dr. Pilsbry applied the old "rules'" for synony-
mous "type genera," but thought priority for family (-idae)
28 NAUTILUS Vol. 73 (1)
names should be kept independent from that for subfamily
(inae) terms. In 1939, when he grouped the American helicoids
in a separate family, he found Cepolidae (1934) was a homonym.
Unaware of Xanthonychidae, he quite logically (from his view-
point) proposed Helminthoglyptidae, and the included Cepo-
liinae (his emendation) , Sonorellinae and Humboldtianinae.
He put Xanthonyx (by definition) in the typical subfamily, and
it does approach Monadenia (cf. 1952) . He did say: "Epiphrag-
mophora Doering is an aberrant genus . . ."
The preceding study has proved that Xanthonychidae (1879)
must be accepted for at least the native American genera of heli-
coids, under the present "rules." Its adoption provides one, quite
considerable advantage. Since the sizes of families are matters of
convenience and/or custom, we Americans, North and South,
can leave to the wisdom of our Old World colleagues the ad-
visability of a separate family for the genera of their home lands.
When they become ready to choose its name, they also might
determine the relative status of the typically European Eulotidae
(1898) and of the primarily Philippine Helicostylinae (1909) ;
both names are prior to either Fruticicolidae (1927 or 1928) .
Incidentally, on the basis of counted citations in the 29 vols,
of the Zoological Record since 1927, Eiilota and/or Eulotidae
remain in much more general use than Fruticicola and/or
Fruticicolidae (both usages combined). The last 17 vols, (since
1939) contain 3 notices of Eiilota fruticum (type species) and
one of Bradyhaena fruticum, by Caesar Boettger, but no usage
in Fruticicola. Why not ask the International Commission to sup-
press completely Fruticicola Held, 1837, which is causing all this
confusion?
STUDIES ON THE GENUS MELAMPUS (PULMONATA)*
By PAUL A. HOLLE and CLARENCE F. DINEEN
State Teachers College, Worcester, Mass., and
Saint Mary's College, Notre Dame, Ind.
The ellobiid genus Melampus was erected in 1810 by Mont-
fort, for Bulimus coniformis Bruguiere from French Guiana,
*This work is a portion of a dissertation by the senior author in partial
fulfillment of the requirements for the degree of doctor of philosophy from
the University of Notre Dame.
■NAUTILUS 73 (1)
PLATE 5
i
.iJilllill i
1, Melampiis coffeus (L.) . 2, M. bidentatus Say. 3, M. floridanus Shuttle-
worth. Scales in mm.
NAUTILUS 73 (1)
PLATE 6
I nil mil
I : : ;i
...
4, Melampus from Captiva Island. Florida (lot 37) illustrating varying no
of stripes (0 to 4) . 5, Snails from Nags Head, North Carolina (lot 21)
dlustratnig varying no. and width of stripes. 6, Snails from Stonington, Conn,
(lot 12) illustrating varietv of shapes, spires and spire angles. All scales
July, 1959 NAUTILUS 29
South America. Since 1810, several species that had been described
earlier have been transferred to the genus Melanipus, and many
new species have been added. However, little agreement can be
found on the relative status of the various species. Many have
been reduced to synonymy.
Johnson (1934) recognized four species and two subspecies of
Melampus along the east coast and Gulf Coast of the United
States:
Melampus coffeus coj] ens (L.) , 1758 (Fig. I). — Cedar Keys (Flor-
ida) to West Indies.
M. c. gundlachi Pfeiffer, 1853 — Florida to West Indies.
M. flavus (Gmelin) , 1789 — Cedar Keys (Florida) to West Indies.
M. floridanus Shuttleworth, 1854-1856 (Fig. 3) — Maryland to
Florida Keys.
M. lineatus SsLj, 1822 (Fig. 2) — Prince Edward Island, Canada, to
the Gulf of Mexico.
Johnson listed these categories, without descriptions or a key,
and we relied upon the original descriptions and redescriptions,
upon such type specimens as are extant, and upon the distribu-
tional data in an eflort to separate mixed collections of salt-
marsh snails.
The essence of taxonomic separations is discontinuity, in struc-
tural features if possible, in habitat or behavior or geographical
range, etc. In Melampus discontinuities are hard to find. This
has led to a great deal of confusion, and to synonymizing in
many ways. In the lengthy list of Pfeiffer (1856), curiously
enough, was M. lineatus Say, and this usage must have been fol-
lowed by Johnson. Since Say described M. lineatus as "Variety
A" of M. hidentatus or as a possibly different species, bidentatus
has priority in event that both names refer to the same species.
Based on fine differences in shell thickness and the degree of
development of the plicae in the shell aperture (personal com-
munication, 1953), Morrison (1950) recognized three subspecies
of M. bidentatus:
M. b. bidentatus Say 1822 — South Carolina to Florida and Texas;
Tampico, Belize; Bahamas; Cuba; N. W. Jamaica; Gonave Id.,
Haiti; Tortola Id.?
M. b. lineatus Say 1822 — New Jersey to North Carolina.
M. b. corneus (Deshayes) 1830 — Prince Edward Island, Canada,
to Staten Island, New York.
30 NAi TILLS \"ol. 7:> {\)
Morrison (^I93n transtened M. fioviddnus to the genus
Detracio.
With such taxonomic disagreenKiu. it is eas\ to understand
whv distributional records are contused. Even records in other
genera have been cited: thus. Hinkley (1907) identified as
Mehvnpus tloriddnus some snails collected in northeastern Mexico
which Dall (Hinkley. 1907) identified as Tralid cin^uhito. Xor
does Johnson's summary (1954) on range provide a solid foun-
dation. He stated that Melampus coffeus was limited in range
trom Cedar Keys, Florida, to the AVest Indies: Pulle\ (1952)
reported M. cofjeus as the onlv species on the coast ot Texas.
Johnson (1934) either overlooked or disbelieved Hinkley 's iden-
tification of M. floridanus from northeastern Mexico, since he
gave the range limits of this species as Maryland to the Florida
Kevs. Johnson (1954) cited M. bide?itatus (as M. lineatus) as
extendino from Prince Edward Island to the Gulf of Mexico,
but did not indicate an exact limit along the Gulf of Mexico.
Among collections studied by the senior author are specimens
identified as M. bidentatus and/or M. lineatus from the coasts
of the Gidf of Mexico and the Caribbean as far as Nicaragua.
These collecting sites are reliably recorded, although the identi-
fications are uncertain and therefore the range of the bidentatus -
lineatus complex cannot be stated Avith finality.
The confusion between M. bidentatus Say, and M. lifieatus
Sav. is particularlv troublesome. Snails fitting Say's original de-
scription are abundant along the northeastern seaboard of Xorth
America. Under his diagnosis of ^[. bidentatus. however, he listed
a 'A'arietv A" as "verv possibly a distinct species . . .' if so. it
mav be called "lineatus. " To this day no worker has demon-
strated clearlv that bidejitatus and lineatus are distinct — vet
both names recin- repeatedlv in the literatiue.
According to Savs 1822 description, the t\pe specimens of
M. bidentatus and M. lifieatus were in the collection^ of the
"Academv and the Philadelphia Museum." All these specimens
were not inherited bv the Academv of Natural Sciences in Phila-
delphia, and a search bv the senior author indicated that they
have been lost.
Under these circumstances, the best appoach aj^peared to be a
thorough studv of topotvpes collected from localities along the
July, 1959
NAUTILUS
31
Table 1, Shells studied for shell characteristics.
X0^^^G.
32 NAUTILUS Vol. 73 (1)
Lot
#
QROUP A 1. 0 0
3. 0 0
5. 0 0
9. 0 0
12. $0 15 1 - U
16. 0 0 .
17. 22 0
20. k9 22 1 - U
21. 21 0
25. 0 0
26. k 8 1-3
27. 0 0
28. 0 0
29. 82 35 1 - 7
31*. 92 3 1 - U
37. 85 11 - 5
39. 96 13 1 - 5
U2. 31 15 1 - U
U6. 33 0
U7. n 52 1 - 6
U8. 60 0
h9. 95 0
50. 71 0
51. 100 33 2 - 8
52. 100 17 1 - 7
53. 100 33 1 - 5
55. 65 0
58. 100 11 1 - 3
63. 67 5 2-5
67. 96 6 3 - U
(moup B 15. 82 0
18. 1(0 0
2U. 90 0
38. 88 0
la. 100 0
liU. 100 0
GROUP C 32. 100 0
5U. 100 0
57. 100 0
60. 100 0
65. 100 0
66. 80 0
68^ 22 2
Table 2. Col. A - ^ of eadx lot havixtg a 3rd denticle.
" B - }S of each lot having parietal ridges,
" 0 - range In the nuaber of parietal ridges.
July, 1959
NAUTILUS
33
Table 3. Stripes - /C of each lot having stripes, and number.
34
NAUTILUS
Vol. 73 (1)
Lot
6
GROUP A
(ROUP B
GROUP C
1.
3.
5.
9.
12.
16.
17.
20.
21.
25.
26.
27.
28.
29.
31.
37.
39.
U2.
1(6.
Ii7.
U8.
19.
50.
51.
52.
53.
55.
58.
63.
67.
15.
18.
2U.
38.
la.
hk.
32.
5U.
57.
60.
65.
66.
68.
68
65
75
71
90
7U
77
80
8U
102
7U
85
90
78
107
98
nii
88
81
78
79
88
77
98
95
100
76
9U
98
90
60
59
72
63
68
67
86
86
85
90
87
86
89
55 - 85
50-85
60-95
60-95
55-115
60-95
60-95
60-100
65 - 105
75 - 115
60-85
75 - 100
75 - 105
60 - 105
90 - 120
80 - 110
95 - 125
80 - 110
70 - 95
70 - 90
60-95
80 - 100
60-100
80 - 120
75 - 125
90 - 125
65 - 105
70 - 115
75 - 125
75 - 110
50 - 70
15-85
60-85
50 - 80
55 - 85
60 - 80
65-
65 -
75 -
75 -
75 -
65-
75-
110
110
no
no
105
105
105
Table U. Angle of spire:
Plicae:
Col. A - mean angle (in o) of all spires.
" B - range (in o).
Col. C - J6 of lot having plicae*
" D - range.
July, 1959 NAUTILUS 35
Atlantic, Gulf of Mexico and Caribbean coasts from Canada to
Central America.
Shell characteristics: Although 5,897 shells collected from 71
localities were carefully studied in an attempt to clarify the taxo-
nomic difficulties, the data of only 43 selected localities are in-
cluded in this report for the sake of brevity (Table 1).
Initially the shells of each lot (from each collecting site) were
measured for length, and separated into 1/2 mm. length groups,
such as 5.0 to 5.4 mm., 5.5 to 5.9 mm. Using terminology pri-
marily that of Abbott (1954) , the following observations were
made on each shell:
1, form and position of denticles;
2, presence and number of parietal ridges;
3, kind of striping, and the number of stripes;
4, angle of the spire;
5, presence and number of plicae on the inside of the lip.
An analysis of the data on these points showed that several
characteristics occurred in regular combinations, and provided a
basis for the separation of shells into three distinct groups. These
characteristics were used to construct the following key:
I. 2nd (parietal) denticle located immediately inside the
shell aperture 1
1. 3rd (parietal) denticle, when present, near the 1st
(columellar) denticle, or plainly separated from (anter-
ior to) the 2nd (parietal) denticle; parietal ridges pres-
ent in some specimens; stripes 0-4, usually 4; angle of
spire 50-125° Group A
2. 3rd (parietal) denticle, when present, located close to
or partly fused with the 2nd (parietal) denticle; parietal
ridges absent; stripes 0-2, usually 2 and near the shoul-
der; angle of spire 45-85° Group B
II. 2nd (parietal) denticle located far inside the shell
aperture Group C
(To be continued)
NOTES AND NEWS
Dates of Nautilus. — Vol. 72, no. 1, pp. 1-36, pis. 1-4, was
mailed July 21, 1958. No. 2, pp. 37-72, pis. 5-8, Oct. 1, 1958. No.
3, pp. 73-108, pis. 9-11, Jan. 15, 1959. No. 4, pp. 109-148, title page
and indexes, pi. 12, April 2, 1959.— H. B. B.
36 NAUTILUS Vol. 73 (1)
Another record of Arion ater. — While on a camping trip in
1937, Mrs. Chace and I stopped in a camp ground in the north-
east section of Seattle. In scouting around for snails, I found an
area of semi-marsh, probably pretty wet in the rainy season. Here
I saw many large, rather chunky and spotted slugs, which I later
found to be Arion ater, the European slug. This colony is known
to several people locally, but apparently never has been reported
in the literature. — E. P. Chace.
Carychium clappi, new name, for Carychium costatum
Hubricht, 1951, Naut. ^5:59, not C. costatum Freyer, 1856, Sitz-
ber. K. K. Akad. Wissensch. (for) 1855, p. 20, pi. 1, fig. 5, =
Zospeum costatum. The new name is proposed in honor of
George Hubbard Clapp, who described it from Alabama, but did
not name it (1906, Naut. 19:\S9) . — Leslie Hubricht.
Unionid introduction in Massachusetts. — On September 8,
1958, the writer began an experiment in hybridization by intro-
ducing three midwestern unionid species into the Ipswich River
drainage in eastern Massachusetts. Such introductions should be
recorded.
The unionids, Lampsilis radiata siliquoidea (Barnes), Ano-
donta grandis Say, and Elliptio dilatatus (Raf.) were collected
at Base Line Lake, Livingston Co., Michigan, by J. P. E. Mor-
rison, P. F. Basch, J. B. Burch, H. J. Walter, and the writer.
Adult specimens, 70 of L. r. siliquoidea, 16 of ^. grandis, and 7 of
E. dilatatus, were placed in a small, recently created lake near
the Valley Road in the Putnamville section of Danvers, Essex
Co. and 66 of L. r. siliquoidea were placed in Silver Lake, Wil-
mington, Middlesex Co. The writer intends to introduce equiva-
lent numbers of L. r. radiata (Gmelin) , A. cataracta Say, and
E. complanatus (Sol.) at the Danvers locality in the spring of
1959 and to observe the lake in succeeding years for possible
further evidence^ of gene flow between related groups. This lake
apparently contained no unionids prior to this introduction.
The Wilmington locality contained only A. cataracta and E.
complanatus before the Lampsilis introduction and I hope that
it will serve as a control to reveal possible phenotypic changes in
iClarke, A. H., Jr., 1958, American Malacological Union Annual Reports
for 1957. pp. 15-16.
July, 1959 NAUTILUS 37
L. r. siliquoidea, resulting from the lower calcium carbonate
content of eastern Massachusetts water. Both of these new locali-
ties contain many species of fish and are similar to Base Line
Lake in physical ecology. — Arthur H. Clarke, Jr.
RuMiNA DECOLLATA IN OKLAHOMA." — On 12 January, 1959, Mr.
R. C. Harrel found several specimens of R. decollata (L.) under
an iron pipe lying on moist, sandy soil near Ada, Pontotoc
County, Oklahoma. Three of these specimens were subsequently
sent to the author as a gift from Dr. C. J. Dennis, of East Central
State College, Ada. To my knowledge this is the first record for
this species in Oklahoma. Pontotoc County is in mid-southern
Oklahoma and lies in the so-called Texan biological province.
This particular part of Oklahoma is characterized by low, rolling
hills, sandy soil, oak-hickory associations along streams and
cross-timbers on the hills.
The specimens represented several stages of growth, from small
and immature to large, fully adult, living snails. The three
specimens sent to Oklahoma State University measured 27.5, 28.0,
and 14.0 mm. in total length. The apical whorls were decollated
from each specimen.
Seemingly in many places this species, where it has been intro-
duced by the activities of man, has become very numerous and
often a pest. Last July (1958) I found it to be one of the more
common species in Zilker Park, Austin, and rather abundant on
the banks of the San Gabriel River, Georgetown, Williamson
County, both in Texas. These snails are very easily transported
in greenhouse materials. In 1948, 15 February, several specimens
of the snail were found around the roots of a shipment of violets
from Fort Worth, Texas. Supposedly, this, or some similar ve-
hicle, is the route via which Rumina entered southern Oklahoma,
after which it escaped. Whether it will be successful is academic. —
Branley a. Branson.
LiGUUs hatched. — I just want to let you know that my first
Ggg from Liguus was hatched April 10. From all I know, this is
the first time, while in captivity. This winter, I collected 2 live
pictus in Florida. — R. Atmus (from letter) .
2Contribution 292 from the Zoology Department and the Research Fotinda
tion, Oklahoma State University, Stillwater,
38 NAUTILUS Vol. 73 (1)
North Dakota. — Is anyone interested in receiving snails or
slugs? If so, I would save specimens for a taxonomist. I do not
know if anyone ever concentrated on North Dakota moUusks. —
R. L. Post, N. D. Agricultural College, Dept. of Entomology,
Fargo, North Dakota.
PUBLICATIONS RECEIVED
The gastropod genus Assiminea in the Philippines. By R.
Tucker Abbott. Proc. Acad. Nat. Sci. Philadelphia 770:213-278,
pis. 15-25. 1958. — This studies the anatomy, ecology, habits and
geographic distribution of 21 species (4 new) and subspecies (2
new) from the Islands and adjacent areas. The very dubious Syn-
cera (literally hearty or sound!) is rejected for the genus, and
the various sections, which have been proposed, are considered
artificial or unusable groups. A "Catalog of names connected
with Assiminea" covers the world. But, does not Assimineidae
date from 1856, instead of 1858 (pp. 215 8c 265)? Incidentally,
"Synceratidae" Bartsch was corrected to Synceridae by Pilsbry &
Bequaert, 1927.— H. B. B.
Distribution and variation of the Hawaiian tree snail Acha-
tinella bulimoides Swainson on the windward slope of the Koolau
Range, Oahu. By d'Alte A. Welch. Proc. Acad. Nat. Sci. Philadel-
phia 77(?: 123-211, figs. 1-2, pis. 10-14. 1958.— In this continuation
of exhaustive studies on variation, especially of color forms, in a
species, which has a range of 72 square miles, 5 subspecies are
proposed as new. These beautiful shells are represented now by
rapidly vanishing, relict colonies or "demes" on the ridges, which
are outlined on maps. — H. B. B.
Studies on the biological control of schistosome-bearing
SNAILS. I. The control of Australorbis glabratus populations by
the snail, Marisa corniiarietis, under laboratory conditions. — II.
Ditto, by the leech, Helobdella fusca. By Eli Chernin, Edward H.
Michelson k Donald L. Augustine. Amer. J. Tropic. Med. &
Hygiene 5 (2) :297-307; 308-314. 1956.— III. The effects of popula-
tion density on growth and fecundity in Australorbis glabratus. —
IV. Further observations, etc. By Chernin & Michelson. Amer. J.
Hygiene <55 (1) :57-70; 71-80. 1957.— V. Ditto I, of Biomphalaria
I
July, 1959 NAUTILUS iii
pfeifferi. By Michelson & Augustine. J. Parasit. 43 (2) :135. 1957.
— The ampullariid ingested eggs and the leech attacked juveniles
especially. Crowding apparently inhibited population growth. —
H. B. B.
Notes sur les limaces. Philomycidae et Limacidae de la re-
publique El Salvador. By C. O. van Regteren Altena. Arch. Mol-
lusk. (97(1/3) :27-31, 2 figs. 1958.— Middle American records are
added for Pallifera costaricensus (variations in color pattern and
penis figured) and Deroceras laeve. — H. B. B.
On the structure of copulative apparatus of Hippeutis com-
planatus (L.) . By Ja. I. Starabogatov. Zool. Zhyr. Ak. Nauk
SSSR. i7 (11): 1743-4, 1 fig. 1958.— This planorbid has a lateral
(not apical) opening in its verge (penial papilla) , and the in-
clusion of it (and maybe Hippeutis also) in the genus Segmen-
tina is suggested. — H. B. B.
Descripcion de una especie nueva de Pomacea de Venezuela.
By T. Pain & Sergio Arias C. Nov. Cien. Ser. Zool. (Caracas)
2^:5-11, pis. 1-2. 1958. — P. falconensis from Chichiriviche, Falcon,
with male anatomy. Apparently its type locality is about 20 miles
from that selected for P. chemnitzii in 1930. — H. B. B.
WILLIAM H. WEEKS SHELL COLLECTION: Now being of-
fered for sale. To receive free lists, send name and address to:
George E. Jacobs, 853 Riverside Drive, N. Y. 32, N. Y.
CASH OFFERED: For a sea shell collection and cone shells.
E. L. Mauseth, Alden, Minnesota
How TO COLLECT SHELLS: Published by the American Malacological Union.
$1.00. Write:
Margajiet C. Teskev, Sect., Route 2, Box 318, Marinette, Wis.
Directory of conchologists. — The 1960 edition will be published the first of
that year. Price will be $2.50, postpaid. For inclusion, write:
John Q. Burch, 4206 Halldale Ave., Los Angeles 62, Calif.
Sea Shells
of Tropical
West America
MARINE MOLLUSKS
FROM LOWER CAUFORNIA TO COLOMBIA
A. Myra Keen
This is the first attempt to list and provide illus-
trations of the sea shells of the entire area of the Pan-
amic marine province — the area between the Gulf of
California and Colombia. Concise descriptions of
1,650 species of sea shells, with about 1,500 illustra-
tions, cover most of the recorded forms larger than
about one-fifth of an inch in length. For the smaller
forms, sample illustrations are given. About 70 type
specimens are here figured for the first time.
The notes on geographic distribution of the species
indicate where the collector might expect to find them,
and an extensive bibliography is included to help the
scientist and serious amateur make use of the scattered
literature. The glossary explains technical terms com-
monly used by collectors, but an effort has been made
to use nontechnical language wherever possible.
Some of the most colorful forms are shown on the
ten pages of four-color plates. $12.50
Stanford University Press
Stanford^ California
THE NAUTILUS
Vol. 73 OCTOBER, 1959 No. 2
AN UNUSUAL OCCURRENCE OF MYA ARENARIA L.
AND NOTES ON OTHER MARINE MOLLUSKS
By ARTHUR S. MERRILL
U. S. Fish and Wildlife Service, Woods Hole, Mass.
Many species of mollusks having a pelagic larval stage settle
upon navigation buoys. These buoys are brought in periodically
for maintenance, at which time the attached species can be
collected. This paper is the result of sampling one such buoy at
the Coast Guard Station, Woods Hole, Massachusetts.
On June 14, 1956, the Coxen's Ledge lighted bell buoy was
placed on station in Buzzards Bay Channel midway between
Gooseberry Neck and Cuttyhunk, Massachusetts (Latitude 41°
27' 00" N. and Longitude 70° 59' 20" W.), and on June 13, 1958,
just two years later, it was brought in for servicing. The buoy is
of the type equipped with a hollow cylindrical stabilizing tube.
The inner surfaces of these tubes provide a sheltered area {see
arrow, figure 1) for many species of animals and plants possibly
unable to withstand the rigors of existence on the ouside of the
buoy. When the buoy was examined, two large specimens of the
softshell clam, Mya arenaria L., were found far up in the stabil-
izer tube nestled in a thick growth of hydroids. Several other
species of mollusks were collected at the same time. These are
listed and discussed briefly.
The normal habitat of the soft-shell clam is within the mud or
sand of the bottom. Our records show that small Mya have been
taken from many buoys. To my knowledge, none approaching
the size I took from this buoy have been reported. Furthermore,
the length of these two specimens averaged 70.0 mm. which
exceeds the normal growth of a two-year-old clam in its usual
habitat. To evaluate the significance of this find, it is desirable
to review briefly some biological aspects of this species.
According to Turner (1953), the preferred habitat of Mya is
generally the upper estuary, although it is found at the heads
of bays and in inlets along the lower estuarine zone where there
is no significant reduction in salinity. Verrill and Smith (1873)
39
40 NAUTILUS Vol. 73 (2)
record softshell clams from the half-tide mark to depths of 40
fathoms, although only smaller clams have been dredged from
the greater depths.
The time at which M. arenaria spawns and the duration of
the pelagic larval stage are functions of temperature (Turner,
1948) . Spawning may begin before May in Wickford Harbor,
Rhode Island (Landers, 1954) , or as late as August in Malpeque,
Prince Edward Island (Stafford, 1912). Metamorphosis may be
complete in 12 days or require as long as three weeks. As far as
can be determined from the evidence at hand, spawning in the
Buzzards Bay area takes place about the time the buoy was placed
on station in June, so probably the Mya spat settled in it soon
after it was put out.
After metamorphosis, the tiny clams usually settle to the bot-
tom and, by means of a byssus, temporarily attach to sand grains,
rocks, seaweed or shell (Turner, 1949). They usually burrow into
the substrate and take up the sedentary phase of their existence
by the time they are about 25 mm. long.
Th large specimens we collected from the buoy were appar-
ently able to survive out of their usual environment because of
the chance protection and support offered by the hydroids and
other growths within the tube. Probably most of the small clams
which attach to buoys fall to the sea bottom, possibly even before
losing the power of byssal attachment. Probably these two speci-
mens were the sole survivors of an initially large set. They not
only survived but grew to a larger size than is usual under
normal conditions.
Newcombe (1935) was able to determine age and the rate of
growth of the soft-shell clam by counting and measuring the
"annual rings" on the shell valves. In the cold Bay of Fundy
waters, he found a two-inch clam to be about five years of age.
Turner (1949) notes that Mya in Massachusetts waters grows
two inches (50 mm.) in two to two and a half years, after which
time growth becomes significantly slower.
I attempted to read the annual rings on the two specimens,
and the measurements of these rings are as follows:
Spawned — June 1956 Specimen #1 Specimen #2
1st winter ring 30.2 mm. 26.5 mm.
2nd winter ring 58.3 mm. 60.4 mm.
Captured— June 1958 68.0 mm. 72.2 mm.
October, 1959
NAUTILUS
41
Figure 1. Sample navigation buoy. Arrow indicates the sheltered inner por-
tion of the stabilizer tube, from which the mollusks w^ere taken.
If my aging is correct, the clams were over two inches (50 mm.) ,
the minimum legal size in Massachusetts, by the time they were
a year and a half old, whereas in this area they do not usually
reach this size until about a year later.
Investigators have studied and compared the growth rate of
Mya and of other commercial bivalves both in their natural
habitat and under conditions similar to those found in buoys.
Belding (1915) suspended soft-shell clams in boxes at various
depths from a raft. He compared these with clams grown in boxes
buried to the level of the substrate in the shallow water near the
shore. Clams suspended from the raft grew faster, and this he
attributed to the faster current. Mossop (1921) secured a number
of blocks bearing mussels (Mytilus edulis L.) to an anchored
buoy and found that the greatest rate of growth occurred near
the surface of the water. In Europe, where mussel farming is
practiced extensively, a faster growth is found in those cultivated
on wooden frames above the sea bottom. Quayle (1956) found
that the raft method of oyster culture greatly enhanced the
42 NAUTILUS Vol. 73 (2)
growtli oi the suspended oysters. He further noted that this
method results in lower mortality because it eliminates silting
and reduces predation.
Belding (1930) considered the most important factor in clam
growth to be a good current which carries food, oxygen and
salts and acts as a sanitary agent. That this condition is met in
the surface waters offshore where buoys are located is evident
not only by the fast growth recorded for Mya in this particular
buoy but also for Mytilus and many other species of mollusks
from this and other buoys.
The mollusks collected in the Coxen's Ledge buoy are listed
below. Included are notes on the frequency of occurrence of the
species to this particular buoy, measurements of the smallest and
largest specimen found for each species, and, where possible,
the number of year classes represented according to our in-
terpretation.
Mya arenaria Linne: very rare. Two specimens of the following
height and length: 41.6 X 68.0 mm. and 42.2 X 72.2 mm.
Anomia simplex d'Orbigny: abundant. Largest 27.6 mm., small-
est 6.4 mm. in height. There seem to be two year classes, one
from about 20-27.6 mm., the other 6.4 to 12.0 mm.
Hiatella arctica (Linne) : rare. Largest 17.0 mm. in length. Rare
in this buoy but usually not at all uncommon when found in
buoys.
Mytilus ediilis Linne: most abundant. Largest 71.5 mm., smallest
2.0 mm. in height. Three year classes as follows:
Spawned 1956, 50.0-71.5 mm.
Spawned 1957, 20.0-40.0 mm.
Spawned 1958, 2.0-15.0 mm.
Mitrella lunata (Say): abundant. Little range in size, average
4.0 mm. in length.
Crepidula fornicata (Linne) : common. 5.6-28.5 mm. in height.
Two year classes, one from about 19.0 to 28.5 mm., the other
5.6 to 13.0 mm.
Crepidula plana Say: fairly abundant. 20.7-27.0 mm. in height.
Apparently only one year class represented.
References
Belding, D. L. 1916. A report upon the clam fishery. Fiftieth
Annual Report of the Commissioners on Fisheries and Game
for the year 1915. Commonwealth of Mass., Boston, Public
Document No. 25.
1930. The soft-shell clam fishery of Massachusetts. Mass.
Dept. Conservation, Div. Fish and Game, Mar. Fish. Series
No. L
October, 1959 nautilus 43
Landers, W. S. 1954. Seasonal abundance of clam larvae in Rhode
Island waters, 1950-1952. U. S. Fish and Wildlife Service,
Spec. Sci. Rept., Fish. No. 117.
Mossop, B. K. E. 1921. A study of the sea mussel {Mytilus edulis,
Linn.) Sect. IV — The rate of growth of the sea mussel
{Mytilus edulis L.) under various intertidal and floating
conditions at St. Andrews, New Brunswick. Contr. Can.
Biol., No. 2.
Newcombe, C. L. 1935. Growth of Mya arenaria L. in the Bay of
Fundy region. Can. Jour. Res., 13.
Quayle, D. B. 1956. The raft culture of the Pacific oyster in
British Columbia. Fish. Res. Bd. Can., Progress Rept. of the
Pacific Coast Stations, No. 107.
Stafford, J. 1912. On the recognition of bivalve larvae in plankton
collections. Contr. Can. Biol., No. 14.
Turner, H. J. 1948. Report on investigations of the propagation
of the soft-shell clam, Mya arenaria. Appendix I. The soft-
shell clam industry of the east coast of the United States.
Div. Mar. Fish., Dept. Conservation, Commonwealth of
Mass., Boston.
1949. Report on methods of improving the shellfish re-
sources of Massachusetts. Factors influencing the abundance
of soft-shell clams in Massachusetts. Div. Mar. Fish., Dept.
Conservation, Commonwealth of Mass., Boston.
1953. A review of the biology of some commercial mollusks
of the east coast of North America. Sixth Report on the
Investigations of the Shellfisheries of Massachusetts. Div.
Mar. Fish., Dept. Nat. Res., Commonwealth of Mass., Boston.
Verrill, A. E. and S. I. Smith. 1873. Report upon the invertebrate
animals of Vineyard Sound and adjacent waters, with an
account of the physical features of the region. Report of the
U. S. Fish Commission for 1871-72.
A NEW BERINGIUS FROM THE PACIFIC NORTHWEST,
WITH COMMENTS ON CERTAIN DESCRIBED FORMS
By ALLYN G. smith
(Concluded from July no.)
The single, well preserved specimen from Chignik, Alaska, in
Mr. Eyerdam's collection is comparable in general aspect and
sculptural characters to the normal form from off the Washing-
ton coast on which the holotype description is based. There are
14 major spiral ribs on the body whorl and 8 each on the penulti-
mate and antepenultimate whorls. Nuclear whorls and operculum
44 NAUTILUS Vol. 73 (2)
are missing. Mr. Norberg's specimen in tlie Tromsoe Museum
is said to be similar, as is the single specimen in the San Diego
Museum from off Vancouver Island.
Mr. Eyerdam's single shell from Raspberry Island, Alaska
(C.A.S. No. 36319), is a large "dead" specimen with sculptural
characters that are intermediate between the first of the two
aberrant shells from the type lot described above (pi. 3, fig. 2)
and the other heavily ribbed specimens. It is illustrated on pi. 3,
fig. 4. This shell is cream-white, covered in places with a blackish-
brown periostracum. Nuclear whorls are missing. The summits
of the postnuclear whorls descend somewhat into the sutures.
Spiral sculpture consists on the penultimate and early post-
nuclear whorls of 3, strong, rounded cords on the lower half and
several other less prominent ones that become progressively
fainter above the periphery and fade out entirely at the tabulate
summits. On the body whorl are about 8 fairly strong spiral
cords below the periphery, which, with their corresponding inter-
spaces, widen toward the outer lip. Above these the spiral cords
become more or less obsolete; below them is the usual series of 8
smaller cords on the outer canal. Overall microsculpture consists
of many fine, closely spaced spiral lines crossed by a series of
equally fine, somewhat less closely spaced and less regular growth
riblets that are weakly nodulose or beaded at the points of cross-
ing. The low, rounded boss inside the upper end of the aperture
is present.
The single shell from Bainbridge Island in Puget Sound, col-
lected by Mr. Eyerdam, is a worn, "dead" specimen that is solid
and heavy in texture but relatively smooth over-all, without any
traces of heavy spiral ribbing whatever. Sculpturally it is much
like the second of Mr. Stiles' aberrant specimens (pi. 3, fig. 3) ,
having two widely spaced, subobsolete cords above the periphery
of the body whorl with indications of several still weaker ones
below. The outer canal, which is somewhat worn, has several
(at least two and possibly more) weakly incised grooves. The
upper portions of the earlier whorls descend somewhat into the
sutures; the summit of the body whorl, however, is not tabulate
and slopes downward in a straight line to the point where the
rounded portion of the whorl begins. The maze of weak, pro-
tractive cording on the upper part of the body whorl, which is
October, 1959 nautilus 45
present on the second of Mr. Stiles' aberrant shells, also occurs
on this specimen. The outer lip is thick and slightly recurved at
its middle part. There is a small siphonal fasciole.
Measurements of the series of shells at hand yield the follow-
ing comparative data:
Max. No. of
Specimens Length^ Diam. L/D whorls
10 adult shells from the type
lot (extremes)
Chignik, Alaska; sculpture
normal
Vancouver Id., British Colum-
bia; sculpture normal
Aberrant shell from type lot;
spiral sculpture obsolete
Aberrant shell from type lot;
spiral sculpture stronger
Bainbridge Id., Puget Sound;
sculpture like preceding
Raspberry Id., Alaska; spiral
sculpture intermediate
In view of the fact that considerable variation may exist be-
tween shells of the same species in this family of marine, carniv-
orous mollusks, probably all the specimens from the general
vicinity of La Perouse Bank, for the present at least, should be
placed in the same species of Beringius. There is little doubt
that earlier workers in the group, based on single shells coming
in separately, would have described at least two and perhaps
three species among the material now at hand. Because of the
known variation in shells from the type lot it seems wiser to
take the conservative step of naming only one species. As more
shells become available, however, a change in this approach may
be indicated. The existence of two specimens, one from the type
lot with relatively smooth sculpture and another quite similar
shell from Puget Sound present a puzzle in speciation that cannot
be solved without more material. As should be pointed out, the
animals in two of the heavily ribbed normal shells prove to be
females. Whether specimens with male animals will show sculp-
2No attempt has been made to estimate the length or number of lost nu-
clear whorls. Dimensions are in mm.
46 NAUTILUS Vol. 73 (2)
tural and other differences in their shells that are constant is also
a question for the future to answer.
Beringius eyerdami is most closely related to B. undatus Dall
but differs in possessing a more globose shell with a shorter spire
and particularly in the far more strongly developed spiral ribs
and channels. It is named for Walter J. Eyerdam in recognition
of his many contributions to the knowledge of conchology and
for the fact that he furnished the first specimen of this new spe-
cies several years ago.
Thanks are due to my associates, Drs. G. Dallas Hanna and
Leo G. Hertlein for advice and assistance with the manuscript.
The excellent photographs used for illustrations are the work
of Mr. E. C. Crompton, the California Academy's photographer.
STUDIES ON THE GENUS MELAMPUS (PULMONATA)
By PAUL A. HOLLE and CLARENCE F. DINEEN
(Concluded from July no.)
Denticles in the aperture. All shells possess at least two con-
spicuous denticles in the aperture; a columellar (1st) denticle
and a parietal (2nd) denticle. An additional parietal denticle
(3rd) is present in some shells (Table 2A) .
The 1st denticle varies little, but the variation in the shape
and position of the 2nd denticle proved significant. In groups A
and B it is located immediately inside the shell aperture, whereas
in group C it appears much farther inside and greatly facilitates
identification of the latter group.
The 3rd denticle is subject to wide variation, its position ap-
pears meaningful, and it may be present or absent. Thus when
present in specimens of group A, it consists of a rounded tubercle
adjacent to or partially fused to the 2nd denticle. In group A, the
percentage of specimens possessing the 3rd denticle was greater
in the southern localities (Table 2A). When present in specimens
of groups B and C, it is rounded and plainly separated from
(anterior to) the 2nd denticle. The incidence of this 3rd denticle
remained consistently high in all habitats south of east-central
Florida. A large percentage of specimens in lots of group B pos-
sessed a 3rd denticle, although the incidence was somewhat lower
in lots from northern localities. The incidence of the 3rd denticle
October, 1959 nautilus 47
in group C is consistently high. All shells of group C came from
southern localities. Consequently, the presence of the 3rd denticle
is correlated with latitude.
Parietal ridges. Low, limy elevations toward the posterior end
on the columellar side of the aperture were present only in some
specimens of group A and varied in the number per shell (Table
2B, 2C) . In general parietal ridges were more prevalent in south-
ern populations.
Stripes. In this study, only the pale stripes between the shoulder
and columellar denticle on the apertural side were counted.
The percentages of specimens in each lot having stripes, and the
variation in number of stripes, are shown in Table 3. In group A,
the incidence of striped specimens varied erratically from lot to
lot, but most stripe-bearing specimens had 4 stripes. The number
of stripes in some lots of group A varied from 0-4, (pi. 6, fig. 4) .
Both the number and width of the stripes varied (fig. 5) . Nearly
all specimens of group B had 2 stripes. Group C with few excep-
tions had 2 or 3 stripes; a majority had 3 stripes.
Angle of spire. The average angle of all specimens in each lot
is given in Table 4. The range in spire angles is wide even among
specimens from a single locality (fig. 6) . For group A, the aver-
age angle of the spire increases gradually from north to south.
For groups B and C, the average angle varies little for shells from
all localities.
Plicae on inside of lip. Some of the shells possessed ridges or
plicae on the inside of the lip. The percentages of shells in each
lot having plicae, and the numerical range, are given in Table 4.
In group A, all specimens (except those from extreme northern
localities) possessed plicae. All specimens of group B and almost
all group C possessed plicae, but no geographical nor ecological
pattern could be found among those of group C lacking this
structure.
If Ball's (1894) explanation for plicae is correct, it is easy to
understand why there would be considerable variation in this
characteristic. According to Dall, the columellar attachment of
snail shells without spirals is as long as the mantle edge, so that
withdrawal into the aperture results in little to no wrinkling or
folding. In forms with strong spirals, such as Melampus, by con-
trast, the attachment area is narrow and separate from the aper-
48 NAUTILUS Vol. 73 (2)
ture. As as result, the distal edge of the mantle is folded during
withdrawal — comparable to the wrinkling of a flattened napkin
when being pulled through a ring. Minerals deposited by the
wrinkled mantle edge would follow its contours and form plicae.
Discussion and summary
Confusion concerning the genus Melampiis appears due to
difficulties in identifying these snails along traditional lines. Shell
structure is variable. The distribution of each species overlaps
others. Life history information has been scarce or lacking alto-
gether (Morrison, 1950) .
As a result of this study of shell structure, two of the five cate-
gories listed by Johnson (1934) are readily identified:
Melampus floridanus (pi. 5, fig. 3) , discussed previously as
"Group B," is segregated on the basis of three features: (1) the
anterior position of the 3rd denticle; (2) its small size (less than
8.5 mm.) ; and (3) two chestnut stripes near the shoulder. On
the basis of habitat study, it would seem that this species can
tolerate low salinity, hence can be found greater distances from
sea water, especially in the upper portions of streams draining
into salt marshes.
Melampus flavus, discussed previously as "Group C," is segre-
gated on the basis of three features: (1) the anterior position
of the 3rd denticle; (2) the internal position of the parietal
denticle; and (3) the presence of three stripes in a majority of
striped specimens. Habitat studies indicate that this species fre-
quents areas which remain submerged by sea water for long
periods. They are often found in rocky areas. In this vigorous
environment they have developed a harder, thicker shell.
Group A has been more resistant to analysis. A study of shell
characteristics and interpretation of habitat and distribution data
suggest three possibilities: (1) that these snails all belong to a
single species of wide variability, this variation showing some
local features (perhaps as clones) and some correlation with
latitude; (2) that these snails belong to two species, a northern
and a southern, the ranges of which overlap along the middle
eastern coast of Florida; or (3) there are two species, a northern
and a southern, with no outliers of the southern species north
of Florida but with outliers of the northern species in southern
localities.
October, 1959 nautilus 49
If further facts support the first possibility (i.e., all in a single
species) , then all members of Group A should be called
Melampus coffeus. If the second and third possibilities should
prove valid, then the southern form would be called M. coffeus
(pi. 5, fig. 1) , the northern form M. hidentatus (fig. 2)
The first possibility is supported by the following:
1. All members have both a columellar and parietal denticle,
with the third denticle becoming more prevalent in southern
portions of the range (Table 2A) .
2. The presence of parietal ridges increases in the more southern
areas (Table 2B) .
3. The number of stripes does not exceed 4, with 4 being the
most prevalent number in the northern populations, but grad-
ually becoming variable in southern populations (Table 3).
4. The angle of the spire changes gradually from acute in the
north to obtuse in the south (Table 4A) .
5. The number of plicae is highly variable but increasing in
number from north to south (Table 4C) .
The second possibility (two species, a northern and southern,
the ranges of which overlap along the middle eastern coast of
Florida), is supported by the following:
1. The percentage of specimens having a third denticle (Table
2A) and parietal ridges (Table 2B) remains nil in most pop-
ulations until the populations of northeastern Florida are
considered.
2. The number of stripes in specimens north of Florida is pri-
marily four, while the southern forms tend to be erratic
(Table 3).
3. The spire angle of specimens north of Florida remains essen-
tially very small, tending to be greater in the more southern
forms (Table 4A) .
The third possibility (two species, a northern and southern,
with no outliers of the southern species north of Florida but
with outliers of the northern species in southern localities) is
suggested by a number of lots studied. They are:
1. Lots 27 and 28 lack a third denticle (Table 2A).
2. Lots 27-28, 48-50 lack parietal ridges (Table 2B) .
3. Lots 28, 46, 48-50 contain specimens most of which have four
stripes (Table 3) , and have very small spire angles (Table
4A).
4. Collecting data, when available, suggest that the northern
and southern forms prefer different habitats — the former pre-
ferring a grassy habitat as opposed to mangrove areas for the
southern form.
50 NAUTILUS Vol. 73 (2)
Resolution of the three possibilities will require additional
studies, especially habitat and life history studies (as was sug-
gested by Morrison, 1950).
As a result of the present study, however, several points have
been clarified:
1. Three different species of Melampus can be distinguished
among the salt-marsh snails of the Atlantic coast, the Gulf of
Mexico, the Caribbean, and Bermuda:
M. coffeus (L) : From the east central coast of Florida south-
ward, including the Gulf of Mexico, the Caribbean and
islands of the West Indies, and Bermuda: M. bidentatus may
be a synonym, or a subspecies, or a separate species, with
range extending from Georgia northward to the Maritime
Provinces of Canada.
M. flavus (Gmelin) : From the east central coast of Florida
southward, including the Gulf of Mexico, the Caribbean
and islands of the West Indies, and Bermuda.
M. floridanus Pfeiffer: From Chesapeake Bay southward, in-
cluding the Gulf of Mexico.
2. In the event that future study demonstrates the duality of
"Group A" and hence the need for a second name, M. bidentatus
is available and should have priority over M. Uneatus. As was sug-
gested by Pilsbry (1927), no reason is apparent for considering
it preoccupied.
3. Variability in the characteristics used by Morrison (1950)
in separating subspecies of M. bidentatus appears continuous
enough that his subdivision seems inadvisable; geographical dis-
tribution of the varieties he recognized shows no discontinuities
that w^ould urge retention of the subspecific names.
4. The specific name M. floridanus should be credited to Pfeif-
fer (1856), not Shuttleworth. The 1854 monograph by Adams and
Adams lists this specific name but gives no description. Accord-
ing to Dr. William Clench (personal communication) , a Mr.
Rugel collected the original Florida specimens for Shuttleworth,
and they are now part of the Cuming Collection in the British
Museum (Natural History) . Neither Rugel nor Shuttleworth
described the specimens; both merely gave them museum num-
bers. Hence floridanus remained a nomen nudum until Pfeiffer
validated it in 1856. As a matter of courtesy, and an index to its
frequent appearance in literature, it could also be M. floridanus
"Shuttlew^orth" Pfr.
October, 1959 nautilus 51
5. Until specimens from Cayo Blanco, Cuba, have been studied
and found to match Pfeiffer's (1853) description of M. gund-
lachi, this name well may be left as incertae sedis.
References
Abbott, R. Tucker 1954 — American seashells. D. Van Nostrand
Co., Inc., New York, xiv— 541, 100 figs.
Adams, H. and Adams, A. 1854 — Monographs of Ellobium and
Melampus, two genera of pulmoniferous Mollusca. Proc.
Zool. Soc. London, 22:7-13.
Dall, W. H. 1894 — The mechanical cause of folds in the aperture
of the shell of Gastropoda. Amer. Nat. 2<^: 909-9 14.
Hinkley, A. A. 1907 — Shells collected in northeastern Mexico.
Naut. 27:68-72.
Johnson, C. W. 1934 — List of marine Mollusca of the Atlantic
Coast from Labrador to Texas. Proc. Boston Soc. Nat. Hist.
^^: 1-204.
Montfort, Denys de 1810 — Conchyliologie Systematique. VoL 2
(Univalves). 1—676, 161 figs.
Morrison, J. P. E. 1950 — American Ellobiidae — an annotated list.
Amer. Malacol. Union Ann. Rpt. (1950) :8-10.
. 1951 — Two new western Atlantic species of pulmonate
mollusks of the genus Detracia and two old ones (family
Ellobiidae) . Jour. Washington Acad. Sci. ^7:17-20.
Pfeiffer, L. 1853 — Neue Auriculaceen. Zeitschr. f. Malakozoologie.
70:124-127.
. 1856 — Monographia Auriculaceorum viventium. Sumptibus
Theodorei Fischer, Casselis. xiii — 209.
Pilsbry, H. A. 1927 — Nomenclature of Leuconia, Melampus and
Truncatella. Naut. 40:125-126.
Pulley, T. 1952 — An illustrated checklist of the marine mollusks
of Texas. Texas Jour. Sci. ^ (2) : 167-199.
Say, Thomas, 1822 — An account of some of the marine shells of
the United States. Jour. Acad. Nat. Sci. (Phila.) . 2:221-276.
PLEISTOCENE MOLLUSCAN NOTES, II
FAUNULE FROM HUNTINGTON BEACH MESA, CALIF.
By JAMES W. VALENTINE
University of Missouri, Columbia
Huntington Beach Mesa is one of a series of low mesas and
plains developed along the coast between Palos Verdes Hills and
San Joaquin Hills, California. To the south, it is separated from
Newport Mesa by the Santa Ana Gap, and to the north, from
52 NAUTILUS Vol. 73 (2)
the small Bolsa Chica Mesa by Bolsa Gap (Poland, Piper et al.,
1956, pi. 1). These mesas lie athwart the Newport-Inglewood
structural zone, and their surfaces are commonly deformed.
Poland, Piper et al. (1956) suggest that they represent remnants
of a formerly widespread surface correlative with the surface of
the Upper Pleistocene Palos Verdes sand.
Marine sediments exposed in stream-cut bluffs and in gravel
pits below the surface of Huntington Beach Mesa have been
assigned to the Lower Pleistocene San Pedro sand (Poland,
Piper et al, 1956, pp. 62-62, pi. 3). However, Mr. Robert G.
Thomas has recently discovered marine fossils in these sediments,
and they indicate an Upper Pleistocene age.
Marine sands and gravels are exposed in a sand pit about
2 miles north of the city of Huntington Beach, Orange County,
California (see Description of fossil localities, to follow). Mol-
lusk shells are locally abundant in several lenses, but are in poor
condition, being chiefly broken and also leached by ground
water. Collections made from three localities in the sand pit
(UCLA, localities 3655, 3656, and 3657) contain a total of 47
species that can be identified definitely and 9 more that may
only be compared or referred to described species or genera,
because of their fragmentary condition (table 1) .
The identified mollusks all represent species that are living
today. Recent populations of most of them live on sandy sub-
strates and along exposed sandy beaches. Only a few species, as
Tegula ligulata and Megatehennus himaculatus, probably re-
quire a solid substratum, and these species are rare in the col-
lections. The most abundant species (for which ecologic data
are available) are exposed sandy beach forms, as Tivela and
Donax, and slightly deeper-water forms that are common along
exposed coasts on relatively coarse, current swept or wave-agitated
bottoms, as Glycymeris suhohsoleta, Tellina salmonea, and
Spisula planulata. These assemblages evidently represent depths
of less than about 5 fathoms along an exposed coast. The sedi-
ments are lenses of medium to coarse cross-bedded sands and
pebble and cobble gravels and also suggest deposition in agitated
shallow water. The collection from UCLA. Locality 3655, com-
posed of fewer species and of relatively more exposed sandy
shore species than the other two, probably represents a beach
October, 1959 nautilus 53
deposit; perhaps the other fossils are in beach deposits also.
Two species are present that live today only well to the south,
where they form part of a northward extension of the Pana-
manian fauna that inhabits embayments along southwestern
Baja California, Mexico. These are Crassinella branneri and
Petricola parallela. These species are both present in the Palos
Verdes sand at San Pedro, California, but are not known in
Lower Pleistocene deposits.
C. branneri is known to live from Laguna Scammon, Baja
California (Woodring et al., 1946, p. 88) to Corinto, Nicaragua
(Eyerdam in Burch et al., 1944, no. 39, p. 9) . Possibly, C. bran-
neri is conspecific with the Recent C. pacifica, which has essen-
tially the same recorded geographic range. C. branneri is par-
ticularly characteristic of warm-water facies of the Upper Pleisto-
cene in southern California, wherein it is recorded from Pacific
Palisades, California (Valentine, 1956, p. 195) to Bahia San
Quintin, Baja California (Orcutt, 1921), chiefly in moderately
protected shallow inner sublittoral facies but also in exposed-
shore facies. A single Recent record of branneri from San Diego
(Dall, 1921, p. 31) is probably based on fossil material (as was
the case with Dosinia ponderosa; see Woodring et al., 1946, p. 84,
footnote 81a) .
Petricola parallela is known to live from Laguna Scammon,
Baja California (Hertlein and Strong, 1948, p. 195) to Corinto,
Nicaragua (Pilsbry and Lowe, 1932, p. 99), from the littoral
zone to 8 fathoms. In the Upper Pleistocene, it is recorded from
San Pedro (U.C.L.A. collections) to near Newport Beach, Cali-
fornia (Bruff, 1946, p. 232) chiefly in protected shallow-water
facies.
A single specimen in the collections represents Tegula
brunnea, which lives only to northward in the Oregonian prov-
ince today. This species is characteristic of exposed rocky-shore
facies of the Upper Pleistocene in southern California, but is
recorded in other Upper Pleistocene facies and also in the
Lower Pleistocene. In the Upper Pleistocene, it usually is associ-
ated with other Oregonian rocky-shore species to form a distinc-
tive cool-water element. T. brunnea commonly lives somewhat
offshore near the surface on kelp (Smith and Gordon, 1948,
p. 201) as well as inter- and sub-tidally on rocks. Shells of brun-
54
NAUTILUS
Vol. 73 (2)
nea in the essentially sandy-bottom, warm water association at
Huntington Beach Mesa well may have been transported from
kelp beds offshore in cooler water, perhaps on broken kelp fronds.
No species that are characteristic of the Lower Pleistocene are
found in the collections; however, most such species belong to
other facies. Chiefly because of the occurrence of the southern
forms, then, and in the absence of conflicting evidence, the as-
semblages at hand are considered to be Upper Pleistocene.
Table 1. Mollusca from the Upper Pleistocene at Huntington
Beach Mesa. Abundance symbols are: R, less than 9; C, 9 to 32;
and A, more than 32 specimens in the collection per 1,000
specimens.
Pelecypoda
Localities: 3655 3656
Glycymeris subobsoleta (Carpenter) C A
Modiolus cf. M. capax Conrad R
Modiolus sp. R
Leptopecten latiauratus (Conrad) R R
Hinnites giganteus (Gray)
Anomia peruviana Orbigny R
Ostrea lurida Carpenter R
Crassinella branneri (Arnold) R C
Lucinisca nuttalli (Conrad) R
Trachycardium quadrigenarium (Conrad) R
"Cardium" sp.
Protothaca staminea (Conrad) R
Tivela stultorum (Mawe) C C
Petricola parallela Pilsbry & Lowe
Tellina bodegensis Hinds R
T. salmonea Carpenter R C
Macoma secta (Conrad) R
Donax gouldi Dall A C
Siliqua lucida (Conrad) R
Solen sicarius Gould
Spisula planulata (Conrad) C
Schizothaerus nuttalli (Conrad) R
Cryptomya californica (Conrad) R R
Corbula luteola Carpenter C C
Zirfaea pilsbryi Lowe
SCAPHOPODA
Dentalium neohexagonum Sharp & Pilsbry R
D. pretiosum Sowerby R
3657
A
R
R
R
R
R
R
R
R
A
R
R
R
R
C
R
C
C
R
C
R
October, 1959 nautilus 55
Gastropoda
Megatebennus bimaculatus (Dall)
Pupillaria parcipicta (Carpenter)
Calliostoma dolarium (Holten)
C. gemmulatum Carpenter
C. ligatum (Gould)
C. tricolor Gabb
Tegula brunnea (Forbes)
T. ligulata (Menke)
Halistylus pupoideus (Carpenter)
Tricolia? sp.
Aletes? cf. A. squamigerus Carpenter
Fartulum occidentale Bartsch
Bittium sp.
Epitonium cf. E. indianorum (Carpenter)
Crepidula coei Berry
Neverita reclusiana alta Arnold
N. reclusiana imperforata (Dall)
Acanthina spirata (Blainville)
Ocinebra interfossa Carpenter
Aesopus chrysalloideus (Carpenter)
Mitrella carinata (Hinds)
"Nassa" delosi Woodring
"Nassa" fossata (Gould)
"Nassa" mendica cooperi Forbes
Olivella biplicata (Sowerby)
O. pedroana (Conrad)
Narona cf. N. cooperi (Gabb)
Conus californicus Hinds
Acteocina culcitella (Gould)
Thus in summary the sediments exposed on Huntington Beach
Mesa are essentially contemporaneous with terrace deposits on
Newport Mesa and probably with the Palos Verdes sands in the
Long Beach and San Pedro regions. The sea was then capable
of supporting mollusks that are today restricted to the Pana-
manian province, presumably because it was warmer, at least at
times, than at present. The shallow, sandy, current and wave
agitated sea bottom where Huntington Beach Mesa now stands
was inhabited by a shallow-water molluscan community similar
to recent southern California communities in such habitats but
including Panamanian species. The upper surface of Huntington
Beach and nearby Mesas is probably a depositional surface, rep-
resenting the last epicontinental deposits of the sea during the
series of events — the "cycle" — that formed the Palos Verdes ter-
56 NAUTILUS Vol. 73 (2)
race platform and its marine cover. The surface is commonly
overlain by thin non-marine sediments that thicken considerably
where the former shoreline was bold, as along its landward mar-
gin at Palos Verdes Hills. A somewhat similar surface seems to
be preserved at an altitude of about 20 feet in the Mission Bay —
San Diego Bay district, California (Valentine, in press).
It is a pleasure to acknowledge the aid of Professor W. P.
Popenoe and Mrs. Lou Ella Saul, University of California, Los
Angeles, in comparing certain specimens, and of Peter U. Rodda,
Bureau of Economic Geology, University of Texas, in collecting
the fossils. Robert G. Thomas, Engineering Geologist, California
State Division of Water Resources, kindly called these deposits
to my attention.
Description of fossil localities
UCLA. Locality 3655. Essentially horizontal sands exposed on
eastern margin of sand and gravel pit, on west face of ridge north
of oil sumps, in N 14 of NE i/4. Sec. 34, T. 5 S, R 11 W, Hunting-
ton Beach Mesa, Seal Beach quadrangle. Orange County, Cali-
fornia. Southwest nose of ridge is now cut away. Uppermost
fossiliferous beds; altitude approximately 25 feet. Rodda and
Valentine, collectors, spring 1957.
UCLA. Locality 3656. Approximately 12 feet stratigraphically
and topographically below UCLA, locality 3655, in a 2-3 foot
pebble and cobble conglomerate. Rodda and Valentine, col-
lectors, spring 1957.
UCLA, locality 3657. A 2-foot pebble conglomerate exposed
in the northeast corner of the sand and gravel pit described in
UCLA. 3655. Evidently represents the same horizon as UCLA.
loc. 3656 (exposure is discontinuous) . Rodda and Valentine,
collectors, spring 1957.
References
Bruff, S. C. 1946. The paleontology of the Pleistocene molluscan
fauna of the Newport Bay area, California. Univ. Calif.
Pubs., Bull. Dept. Geol. Sci., 27 (6): 213-240, 12 text-figs.
Burch, J. Q. (ed.) et al. 1944. Distributional list of the west
American marine mollusks from San Diego, California, to
the Polar Sea, Pt. I, Pelecypoda. Conch, Club So. California,
Minutes, nos. 33-45, pagination by issue.
Dall, W. H. 1921. Summary of the marine shellbearing mollusks
of the northwest coast of America, from San Diego, Cali-
fornia to the Polar Sea, etc. U. S. Nat. Mus., Bull. 112,
217 pp., 22 pis.
Hertlein, L. G.. and Strong, A. M. 1948. Eastern Pacific expedi-
tions of the New York Zoological Society. Mollusks from
October, 1959 nautilus 57
the west coast of Mexico and Central America. Pt. 6
Zoologica 33: 163-198, 2 pis.
Orcutt, C. R. 1921. Pleistocene beds of San Quentin Bay, Lower
California. West Am. Scientist 19: 23-24.
Pilsbry, H. A., and Lowe, H. N. 1932. West Mexican and Cen-
tral American mollusks collected by H. N. Lowe, 1929-31.
Proc. Acad. Nat. Sci. Philadelphia 84: 33-144, pis. 1-17, 6
text-figs.
Poland, J. F., Piper, A. M., et al. 1956. Ground-water geology
of the coastal zone, Long Beach-Santa Ana area, California.
U. S. Geol. Surv., Water-Supply Paper 1109, 162 pp., 8 pis.,
2 text-figs.
Smith, A. C, and Gordon, Mackenzie, Jr., 1948. The marine
mollusks and brachipods of Monterey Bay, California, and
vicinity. Proc. Calif. Acad. Sci., ser. 4, 26 (8) : 147-245, pis.
3-4, 4 text-figs.
Valentine, J. W. 1956. Upper Pleistocene Mollusca from Potrero
Canyon, Pacific Palisades, California. Trans. San Diego
Soc. Nat. Hist. 12 (10) : 181-205, pi. 13.
(in press) Pleistocene molluscan notes. L The Bay Point
formation at its type locality. Jour. Paleontology 34.
Woodring, W. P., Bramlette, M. N., and Kew, W. S. W. 1946.
Geology and paleontology of Palos Verdes Hills, California.
U. S. Geol. Surv., Prof. Paper 207, 145 pp., 37 pis., 16 text-
figs.
MOLLUSKS OF THE SALT RIVER, KENTUCKY
By JOSEPH ROSEWATER
Museum of Comparative Zoology, Harvard University
An expedition was made by Dr. W. J. Clench and myself dur-
ing September, 1958, to collect mollusks in the Kentucky and
Salt River systems, Kentucky. We left Ann Arbor, Michigan,
September 6, 1958, at the close of a most enjoyable meeting of
the American Malacological Union. Traveling south through
southern Michigan and Ohio, we collected at several stations
in the Great Lakes drainage, and upon arriving in Kentucky
on September 7, at two localities on the Licking River. Since
the water level was somewhat high, we proceeded to the Kentucky
River where several stations were made on the main river and
its three forks. This was followed by a week-end trip to Cleve-
land, Tennessee, to visit Herbert Athearn, an ardent collector
of fresh-water mollusks. During this side trip, fine collections
were made in the Powell and Sequatchie Rivers, Tennessee, in
58 NAUTILUS Vol. 73 (2)
the Conasauga River of northwest Georgia and in the Green
River, Kentucky.
The Sak River drainage system occupies a roughly oval area
in north central Kentucky. Along most of its eastern boundary,
it is separated from portions of the Kentucky River system by
only a few miles. On the south and west, it is bordered by the
Green River system. On the north, tributaries of the Salt River
extend north of Louisville, Kentucky. It flo-^vs into the Ohio
River about 20 miles southwest of Louisville almost precisely
on the line of 38° N. Latitude. Two major forks, the Rolling
Fork and Beech Fork, and the Salt River propei- originate fairly
close together in Boyle and Casey Counties, Kentucky. The
Rolling Fork, the most southern branch, flo^ss in a northwesterly
direction to its confluence with the Salt River proper about nine
miles southeast of the Ohio River. The Beech Fork flows north,
then west to join the Rolling Fork about twelve miles southeast
of the confluence of the latter with the Salt River. The Salt River
proper parallels the Beech Fork during its initial northerly flow
but continues farther north than the latter, then turns west and
flows in this direction to its confluence with the Rolling Fork,
then northwesterly to the Ohio River. \'en' approximately, the
drainage area of the Salt River system covers 3000 to 4000 square
miles. The system occupies a portion of the "Blue Grass" area
of Kentucky, a region of limestone eroded to form a generally
rolling surface, and is given over in large part to farming. Also
this area supports one of Kentucky's outstanding industries, that
of the distillation of fine whiskey. One asks himself on picking
up a series of especially large Quadrula quadrula (Rafinesque),
"Could there be some correlation between the large size of these
bivalves and the dumping of distillery wastes into streams of
this area?"
We based Salt River collecting operations at Bardstown, Nel-
son Co., Kentucky, and spent four days making ten stations in
this river system. \Ve intended to make several additional sta-
tions in the Salt and also the lowei* Kentucky River working our
way east (and home) . Ho^vever, a heavv t^vo-inch rain ended the
ideal collecting conditions which had existed for nearly two
weeks and our last stations could not be made. Consequently,
a thorough report on the mollusks of the Kentucky River must
be delayed until further collections can be made there. Our
October, 1959 nautilus 59
collections from the Salt River were substantial, however, and
are therefore reported upon at this time.
Three stations have been made on the Salt River previously
that we know of, although other collectors have undoubtedly
visited the area. The known visited localities are those of Clench
and Okkelberg in 1927, corresponding to the Clench-Rosewater
stations 1934 and 1935 plus one which we did not make: Salt
River, 4 miles west of Lawrenceburg, Kentucky. An account of
the fresh-water mussels collected by Clench and Okkelberg was
published by Clench and van der Schalie (1944). No account
was given of the gastropods collected.
Ortmann (1926, p. 187) stated that the Salt River system
should be expected to have an Ohioan fauna rather than a
Cumberlandian one. Clench and van der Schalie showed this to
be true and our 1958 collections only reinforce this view. On the
basis of the 1958 collections, four species of mussels may be
added to the list of Clench and van der Schalie for the Salt River:
Villosa lienosa (Conrad) ; Carunculina parva (Barnes) ; Trun-
cilla donaciformis (Lea) ; Anodonta grandis Say. On the other
hand, three species mentioned in 1944 were missing from those
collected in 1958: Cy progenia irrorata (Lea) ; Ohovaria subro-
tunda (Lea) ; Anodonts. imbecillis Say.
Goodrich (1939, p. 2 and 1940, p. 13) stated that forms not
separable from Pleurocera acutum Rafinesque^ occur in tribu-
taries of the Cumberland and Duck Rivers, Tennessee. This is,
apparently, also true of the Beech Fork and the Salt River proper
stations (1934,-36, -40, -41) as forms representing and, at present,
indistinguishable from P. acutum were also found there.
To my knowledge, there is no previous published record of
Pleurocera canaliculatum (Say) from the Salt River system. We
found it, along with Lithasia obovata (Say) at station 1939, a
locality with a distinctly large river ecology. The Lithasia had
remarkably complete, pointed spires for this species. The other
locality recorded for P. canaliculatum, station 1934, yielded
only a few dead specimens. For this reason, it is questionable
whether the species lives there or was washed down from an
area above a partially demolished rock dam where the Beech
2 Morrison (1954, pp. 359-364) discusses the taxonomy of the subfamily
Pleurocerinae in North America and gives reasons for use of different
nomenclature than is employed here; also see Hemming (1951, pp. 6-17) .
60 NAUTILUS Vol. 73 (2)
Fork runs wide and deep. Attempts to collect above the dam
failed because of hazardous conditions due to steep and slippery
banks at this locality.
I wish to acknowledge the aid of my associates, Richard I.
Johnson and Arthur H. Clarke, Jr., in the department of mol-
lusks. Museum of Comparative Zoology, in checking identifica-
tions of certain mussels. H. B. Herrington identified Sphaerium
striaiinum (Lamarck) . Thanks are given to Dr. W. J. Clench
for checking the identification of Physa and Campeloma, for
sharing with me his broad knowledge of fresh-water moUusk
collecting and for the many valuable experiences we shared in
the field. Financial aid for this trip was made possible by the
Friends of The Department Of Mollusks Fund, Museum of
Comparative Zoology.
The following is a list of the collecting stations visited in the
Salt River system, Kentucky, in September, 1958. Preceding each
station and its description is the station number. This number
is repeated in the list of fresh-water mollusks denoting where
each species was collected.
1932. Rolling Fork, Salt River, 7 miles southwest of Lebanon,
Marion Co., Ky. (Ky. Route 55) ; September 16, 1958. Ledge
rock with gravel-rock overlay; water clear, quite warm; gastro-
pods and bivalves rare, the latter often lying out on substrate, the
former crawling on rocks and gravel.
1933. Cartwright Creek, 13 miles southeast of Bardstown,
Washington Co., Ky. (U.S. Rt. 150) ; September 16, 1958. Ledge
rock; water very shallow; no bivalves found; Goniobasis abundant
in pools and on bridge pillars.
1934. Beech Fork, Salt River, 1 mile southwest of Bardstown,
Nelson Co., Ky. (U.S. Rt. 31E) ; September 17, 1958. Gravel,
some rock; water low, dammed above bridge; a large sewer pipe
entering just below dam; dead bivalves and snails abundant;
Physa abundant on sewer sludge.
1935. Rolling Fork, Salt River, 1 mile south of New Haven,
Nelson Co., Ky. (U.S. Rt. 31E) ; September 17, 1958.. Rocky
gravel; water clear in spite of heavy rain, flowing over rapids
and shoals; bivalves abundant, pleurocerids fairly so.
1936. Beech Fork, Salt River, about 3 miles south of Bardstown,
Nelson Co., Ky. (Ky. Rt. 49) ; September 18, 1958. Ledge, gravel,
stones, sand and some silt; dead bivalves abundant, apparently
washed out of substrate by recent freshets; pleurocerids in very
great abundance on mud and rocks; viviparids in sand; dead
sphaeriids in great abundance on banks.
October, 1959 nautilus 61
1937. Rolling Fork, Salt River, Raywick, Marion Co., Ky.
(Ky. Rt. 527); September 18, 1958. Gravel bars with some rock;
bivalves abundant; Goniobasis abundant in drying puddles.
1938. Wilson Creek, about 2 miles northwest of Boston, Nel-
son Co., Ky. (Ky. Rt. 61) ; September 19, 1958. Ledge rock;
water clear, fairly warm, silt on ledges; no bivalves collected;
Physa and Lymnaea rare; Goniobasis very common.
1939. Rolling Fork, Salt River, 1 mile southwest of Lebanon
Junction, Bullitt Co., Ky. (Ky. Rt. 434); September 19, 1958.
Steep, muddy banks, channel narrow and deep; water roily,
rather fast flow; bivalves probably present but impossible to
collect; Lithasia obovata and Pleurocera canaliculatum not un-
common on banks.
1940. Salt River, 3 miles southeast of Mt. Washington, Bullitt
Co., Ky. (U.S. Rts. 150 and 31E) ; September 19, 1958. Gravel-
rock bottom, sand and silt; water fairly clear; bivalves very
abundant, especially in backwater areas; pleurocerids rather rare.
1941. Salt River, 5 miles west of Taylorsville, Spencer Co., Ky.
(gravel road) ; September 19, 1958. Gravel-rock bar, shoals ex-
tending for several hundred feet on either side of bridge; water
somewhat murky, a few rapids; large bivalves very abundant
above rapids; pleurocerids common on rocks and mud at mar-
gins, Campeloma very abundant in sandy mud.
MOLLUSKS OF THE SaLT RiVER SYSTEM
Viviparidae
Campeloma ponderosa (Say)
Lioplax sulculosa (Menke)
Pleuroceridae
Pleurocera acutum Raf.
P. canaliculatum (Say)
Goniobasis semicarinata (Say)
Lithasia obovata (Say)
Physidae
Physa integra Haldeman
Lymnaeidae
Lymnaea humilis Say
L. columella Say
Unionidae
Fusconaia flava (Raf.)
Crenodonta gigantea (Barnes)
C. costata (Raf.)
X X X X X
X X X
X X X X
X X
X X X X X X X X X
X
: X X X
X
X
X X X
X X X
XXX X X
62 NAUTILUS
Quadrula pustulosa (Lea)
Q. quadrula (Raf.) x x
Tritogonia verrucosa (Raf.) x
Elliptio dilatatus (Raf.) x
Lasmigona costata (Raf.) x
L. complanata (Barnes) x x
Anodonta grandis Say x x
Alasmidonta calceolus (Lea) x
Strophitus rugosus (Swainson)
Ptychohranchus fasciolaris x
(Raf.)
Ohliquaria reflexa Raf.
Actinonaias carinata (Barnes) x x
Leptodea fragilis (Raf.) x x x
Proptera alata (Say) x
Carunculina parva (Barnes)
Villosa lienosa (Conrad)
Lampsilis anodontoides (Lea) x x
L. radiata siliquoidea (Barnes) ^ x
L. ovata ventricosa (Barnes) x x x
Truncilla donaciformis (Lea)
T. truncata Raf.
Dysnomia triquetra (Raf.)
Sphaeriidae
Sphaerium striatinum
(Lamarck) x x
Literature cited
Clarke, A. H., Jr. 1958. Distribution and apparent introgression
of the Unionidae in central New York. The American Malac.
Un., Annual Rpts. for 1957, Bull. 24: 15-16.
Clench, W. J., and Henry van der Schalie. 1944. Notes on Naiades
from the Green, Salt, and Tradewater Rivers in Kentucky.
Papers Mich. Acad. Sci., Arts, and Letters, 29: 223-229.
Goodrich, Calvin. 1939. Pleuroceridae of the St. Lawrence River
basin. Occ. Papers Mus. ZooL, Univ. of Mich., No. 404: 1-4.
1940. The Pleuroceridae of the Ohio River drainage system.
Occ. Papers Mus. ZooL, Univ. of Mich., No. 417: 1-21.
Heming, Francis. 1951. Report on the question whether, in order
to avoid confusion in nomenclature, it is desirable that the
plenary powers should be used by the International Com-
mission On Zoological Nomenclature to vary the type species
of the genus "Pleurocera" Rafinesque, 1818 (Class Gastro-
poda, Order Mesogastropoda) . Bull. Zool. Nomenclature,
2: 6-17.
3 Clarke (1958, pp. 15, 16) has shown that reproductive isolation is lack-
ing between L. radiata (Gmelin) and L. siliquoidea (Barnes) and that the
latter may be considered a subspecies of the former.
October, 1959 nautilus 63
Morrison, J. P. E. 1954. The relationships of Old and New World
melanians. Proc. U. S. National Mus., 103: 357-394.
Ortmann, A. E. 1926. The Naiades of the Green River drainage
in Kentucky. Ann. Carnegie Mus., 17: 167-188.
ON THE GREEN LAND MOLLUSK FROM NEW GUINEA
By J. B. HENRARD
Oegstgeest, The Netherlands
Some time ago, I received a letter from a fellow conchologist,
who called to my attention an article by Dr. Clench, in Breviora,
1957, no. 76. In this paper, a curious green land-shell was de-
scribed; it was said to be from New Guinea, but without exact
locality or collector. The species was received from Mr. Poling,
who bought it from a dealer, and presented it to the Museum of
Comparative Zoology, at Harvard University. My American
friend, w^ho knew that I had brought together a very large col-
lection of land-snails during the years I stayed in Dutch New
Guinea, suggested that this green snail probably would be repre-
sented in my cabinet. Indeed, since I had received a copy of the
publication from him and had read Dr. Clench's paper carefully,
I recognized the shell at once, and had it from several, widely
separated localities in northwest New Guinea. The shell was
considered by Dr. Clench as a member of the subfamily Chlori-
tinae, and a new subgenus Verdichloritis was created for it. The
photo in the article, enlarged 4 times, is rather bad. The name of
the species finally was given as Eustomopsis (Verdichloritis)
polingi Clench.
All the specimens in my collection agree so completely, in
dimensions and in all other characters, with the description of
Dr. Clench, that I do not hesitate to accept them as belonging
to his species. However, I have a different opinion in regard to
the correct genus to accept for this species, because all my
examples, which are adult or were taken alive, have a very
striking character. Just behind the reflexed peristome, the body
whorl has a very curious gibbous crest, which is separated from
the peristome by a shallow sulcus, that runs downwards to the
open umbilicus. This character does not occur in the genus
Chloritis, but is present in all species of the subgenus Cristigibba
Tapparone-Canefri, which was established for those species of
the genus Planispira that have such a gibbous crest. This sub-
64 NAUTILUS Vol. 73 (2)
genus was accepted as a section by Thiele, Handb., p. 681. This
character is absent from Chloritis, as may be seen by comparison
of Thiele's descriptions of the two genera. Shells with hairy
periostraca occur in both.
I wish to give here all the data on the 1949 discovery of this
species.
Mollusk collecting in the tropical regions is much more dif-
ficult than elsewhere, because the collector is hampered by the
very dense vegetation of the primeval forest. The presence of
snails often is indicated by dead and worn shells among rotten,
wet leaves at the bases of trees and shrubs. In somewhat more
open places, two species of Planospira are rather common. Both,
although variable in banding and in color of peristome, are
easily recognizable. These species are: Planispira tortilabia
(Lesson) and P. corniculum (Hombr. Sc Jacq.) . Both belong to
the subgenus Cristigibba also. I collected plenty of them; their
beautiful varieties are still insufficiently described. During my
holidays, I was a guest of some friends, who had a bivouac near
Klamesien, a locality situated on the western part of the large
peninsula called "De Vogelkop" (i.e.. Bird's Head) . At Klame-
sien, I could make many exploring trips in all directions; this
locality proved to be very rich; and a fine collection was brought
together. One day, under very dense vegetation, I found a dead
but perfect shell of a dull grayish color, but with the same gib-
bous crest as in other members of the Cristigibba group. It was
thus a species of the genus Planispira, but was quite unknown
to me, being smaller than the two species mentioned above.
Moreover this specimen was peculiar because, within the mouth
on the parietal area, there was a green spot. Further search
yielded more, similar specimens and also a much damaged one,
which was, however, provided with a green periostracum and
was still somewhat hairy. Fortunately this damaged shell had
the last part of the body whorl with a quite intact peristome,
and proved to be conspecific with the first examples, which were
totally denuded shells. This green species is arboreal, and de-
tection of living ones among the dense vegetation of nearly the
same color was very difficult. The locality was combed as thor-
oughly as possible but only two living specimens were obtained.
The greenest Planispira would be a suitable name for this unique
October, 1959 nautilus 65
species. Understandably, when other localities were visited, my
friends and I tried to locate this species, and we did find them,
but never in sufficient quantities. My friend J. Kanter handed
me two fine specimens, found by him near Klamaloe, and Mr.
H. C. Kavelaars, a very good shell-hunter, gave me a very fine
one found between the Klaga- and Klasafet River. All these are
from localities north of the Gulf of McClure. We were, however,
fortunate enough to detect this species also south of the Gulf,
when we had a beautiful and very profitable trip to the large
island of Salawati. Here we went further on the Waiboe River,
a long way, to our bivouac near Waileh in the northern part of
the island. There, on the so-called Waiboe Ridge, a calcareous
area, very fine mollusks could be procured, and once more we
obtained a few living specimens. Our conclusion was that this
species, which occurred in so widely separated localities, is not
actually very rare, but appears to be, because it is so difficult to
obtain. As a member of the genus Planispira, its name now be-
comes: P. (Cristigibha) polingi (Clench) .
In regard to the other green shells, mentioned in Dr. Clench's
paper, there are fundamental differences between this Planispira
and such genera as Liguus, Amphidromus, and all the other
shells mentioned by him. In all those shells, the coloring matter
is situated in the calcareous layer of the whorls, while in this
green Planispira only the periostracum and its hairs are green.
To the species, in which totally green whorls occur, I may add
the very beautiful Amphidromus ventrosulus Moellendorff,
which, according to Fischer and Dautzenberg, is a subspecies of
A. smithi Fulton. In this shell, the aperture also is green-colored.
STUDIES ON MOLLUSK POPULATIONS: 4
By R. STOHLER
Department of Zoology, University of California, Berkeley 4.
T. S. Oldroyd (1918) described a new variety of Olivella hi-
plicata (Sowerby) and named it angelena. It was distinguished
from the typical species as "being more delicate and slender."
He also states: "Sowerby 's type came from Monterey and does
not occur near San Pedro living, but is found fossil there in the
Pliocene and lower Pleistocene. Variety angelena is found fossil
in both the upper and lower San Pedro beds of the Pleistocene."
66 NAUTILUS Vol. 73 (2)
From the text of the note the reader is led to assume that
O. b. angelena is based on a fossil shell, but the holotype in the
Oldroyd collection at Stanford University is a fresh shell from
Los Angeles.
In 1921 the same author described two additional varieties
of Olivella bipUcata. The first of these two varieties he named
fucana and stated that it was broader across the middle and
lower part of the aperture than var. angelena. In addition to the
type locality "Straits of Fuca, near Cape Flattery" where the
variety was collected alive he also lists the Pliocene at San
Pedro. The second variety, named parva, had been collected at
Point Abreojos in Lower California by Henry Hemphill and was
described as being "nearest angelena, but much smaller, a little
broader in proportion; . . .; found in the upper Pleistocene at
San Pedro." With the descriptions of these two new varieties,
Oldroyd figured presumably the type specimens of all three
varieties as well as of a shell he called "typical" (obviously
meaning a typical O. biplicata).
Mrs. Oldroyd (1927) reprinted the descriptions almost ver-
batim, except that the first variety was then called angelina, that
in the description the type locality was given as Los Angeles
County (not included in the original description) and that
nothing was said about the fact that T. S. Oldroyd (I.e.) made
no reference whatever to a living specimen. Furthermore, the
range of the variety was given as from San Pedro to San Diego.
On her plate 26, Mrs. Oldroyd illustrated the varieties; how-
ever, the figures evoke the impression that they have been badly
retouched and close inspection indicates the possibility that the
photographs of the shells were cut out and pasted up for
reproduction.
Through the generous cooperation of Dr. A. Myra Keen, for
which I wish to express my gratitude, I was permitted to measure
the holotype of each variety in the Oldroyd collection at Stan-
ford University; at the same time all other designated specimens
in the type lots of the same collection were measured. To insure
greater accuracy, all specimens were measured again after ap-
proximately one year had elapsed. The results are given in
Table 1.
D. S. and E. W. Gifford (1944) gave a number of measure-
1
I
I
October, 1959 nautilus 67
merits, made on a large series of shells collected in various
localities. They used the length-width index as an indicator of
the relative obesity of shells. This index is obtained by dividing
the width of the shell by its length and multiplying the result
with 100. The Giffords showed that the populations they studied
fully overlapped, as far as the indices were concerned, all Old-
royd's varieties, although for the indices of the latter they limited
their calculations to the measurements of the type figures in
the Nautilus (1921) and the measurements of the shells as
recorded by Mr. Oldroyd.
The problem of relative obesity and absolute shell sizes at-
tracted my attention. Olivella biplicata, being a rather common
shell and easily collected throughout the year, at least in certain
localities, seemed to be an especially favorable object for a more
generalized study. A number of observations concerning the
natural history of this species were made in the course of this
particular inquiry and some of them, which may have a bearing
on some of the aspects involved in the questions revolving
around the validity of Oldroyd's varieties, will be included in
this report.
To insure accuracy as well as speed in measuring the shells
(a total of over 6200 was involved) a modified sliding caliper
was used to construct an imaginary rectangle around the shell.
Figure 1 illustrates the manner in which the width of the shell
was measured and Figure 2 shows the measuring of the length
of the shell. The dial allows direct reading of tenths of milli-
meters, while the bar shows the centimeters. By adding the cross
bar, accurately ground to form a snug gliding fit for the legs of
the caliper, truly comparable measurements on all shells could
be made with ease. The two ends of the canal of the shell were
used to form a firm base, as shown in the figures; the width was
obtained in all cases by gently rotating the shell between the
legs of the caliper until the greatest width was found. That the
method was reliable is borne out by the fact that no differences
were obtained in the measurements of the same shells in the
Stanford collection made over a year apart.
Table 2 gives the results obtained from 7 lots, collected in
California from Drakes Estero, Marine County (appr. 38° N)
to La Jolla, San Diego County (appr. 33° N). The accompanying
68
NAUTILUS
Vol. 73 (2)
i"4l"'i"T'
lOiO IliO I2|0
Figue 1: Sliding calipers, modified. The base of the canal of the shell is
gently rotated until the largest diameter is ascertained; the dial reads in
tenths of millimeters.
Figure 2: Sliding calipers, modified. To ascertain the greatest height of the
shell, the base of the canal is set firmly upon the moving leg of the instrument
and moved until the apex just touches the stationary leg.
October, 1959
NAUTILUS
69
PP ^^Y'T-^-''" ^
Figure 3: Outline map of California showing the six localities in which
Olivella biplicata (Sowerby) were collected for this study. DE = Drakes
Estero; DR = Duxbury Reef; PP = Pillar Point; MH z:^ Monterey Harbor;
GP — Government Point; LJ = La Jolla.
map (Fig. 3) shows the 6 localities at which the collections were
made. In Table 3, the figures obtained from 11 separate groups
collected in one locality at more or less regular intervals are
reproduced. Additional lots have been collected, measured and
their indices computed for the Duxbury Reef area (near Bolinas,
Marin County, California) but the figures are not included here
as it was considered unnecessary to add more of the same results.
In actuality, the study at Duxbury Reef covered collections over
a period of 18 months with semi-monthly visits, meteorological
conditions permitting; the total included 4258 individuals, and
70
NAUTILUS
Vol. 73 (2)
Table 1: "Varieties" of Olivella hiplicata (Sowerby)
in the Oldroyd Collection at Stanford University.
among them 49 mating pairs (M. P.) .
In all three tables the same terms are used. Measurements are
in units of one-tenth millimeters; thus the holotype of O. bipli-
cata angelena Oldroyd measures 278 units or 27.8 mm. in length
and 137^ units or 13.75 mm. in greatest width. From each lot,
after all measurements were completed, the figures for the largest
(i.e., longest) and smallest (i.e., shortest) specimen were se-
lected; in the same manner the slenderest or least wide and the
most obese or widest specimen was singled out. Finally the
purely arithmetical averages were computed. The information in
October, 1959
NAUTILUS
71
II
CO 00 <30
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CO CO
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00
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^ t^ <30
O Tt^ Co
ut) GO ^
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O
E
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O
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I
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00 CO
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<:o
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o ^ "^o
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O TtH o
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00 lO^
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<:o
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(M '^ Oi
05 r-H lis
'^ CO ^^
t^ CO •
(M 1-H o:)
CO(N(^
(M CO Oi
(N i-H Q)
2^00
<:o
t^ CO <so
t- CO 00
CO t^ Oi
■^ 00 iJ^
o ^ o
o t-
CO 1-H ?^
lO Ir^ 00
o o •
O Tt^ ^
CO CO •
C5 00 >-i
CO o •
(M ^ QO
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O 05 00
CO 'TiH Oi
(N CO ^
iCjnqxRQ
00
(N lO 00
1—1 CO
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00 CO Oi
CO <M •
(M ^ ir:5
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(M T^ "-^
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05 CO <to
1-1 (M i^
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59
-^ o 00
^ rt^ •
(M 1-1 2^
coio "^
-^ (N ^
T:t^ CO Oi
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a
the tables is considered sufficient for the present study.
Of the three Oldroyd varieties, as will be recalled, angelena
was supposed to be slenderer and fucana broader or more obese
than "typical" Olivella biplicata. For reasons to be discussed
below, these two varieties will not be compared here with "typi-
cal" shells. Table 1 shows, by the obesity index, that O. h. fucana
is, indeed, more obese than O. b. angelena, at least if only the
holotype is considered. As may be noted, however, the "slender"
variety includes at least one specimen that is more obese than
the most obese of the so-called obese variety; furthermore, as
appears from the averages, the "slender" variety is actually more
72 NAUTILUS Vol. 73 (2)
obese than the "obese" variety. This alone should be sufficient
to show the invalidity of the criterion used by Oldroyd to dis-
tinguish the two varieties.
In regard to O. biplicata parva Oldroyd, as may be pointed out,
we have, in a fairly good sample, a shell of a small average; as I
have been unable to obtain large, unselected population samples
from Punta Abreojos I am not prepared to make, at this time,
any positive statements. However, I believe that the table gives
some clues, when taken together with the information contained
in the other two tables, as to the nature of this particular lot
(see the discussion further below) .
A careful comparison of the various figures in Table 2 shows
that, from sample to sample, there is but little variation in the
data, a fact which should not be surprising, although it might
be expected that in the extremes of the distributional range of
a species having such a large geographical range, differences
might be observed. This still might hold true for O. biplicata
if significant population samples from the true extremes of the
range were available; Table 2, however, includes only a somewhat
southern quarter of the entire range, it being the area from 33°
to 38° north latitude, while, according to Keen (1937) the
species may be found from 25° to 49° N.
(To be continued)
THE CHARLES M. WHEATLEY COLLECTIONS
By RICHARD I. JOHNSON
Charles Moore Wheatley was born in 1822, and in 1842, at
the age of 20, he published privately a "Catalogue of the shells
of the United States, with their localities." A second edition
appeared in 1845. His only other paper on conchology was pub-
lished in 1865 in the American Journal of Conchology 1, pp.
65-67: "Revision of M. Petit's Catalogue of the Genus Mono-
condylaea." He was an engineer by profession, also interested in
geology, and made some important fossil discoveries in Penn-
sylvania. Wheatley died on May 6, 1882.
In his article of 1862 on "The history of conchology in the
United States," G. W. Tryon mentions that Wheatley 's collec-
tion had become the property of Union College, Schenectady,
New York. By this date, Isaac Lea had described 19 species which
he mentioned as being "In the cabinet of C. M. Wheatley." This
October, 1959 nautilus 73
collection after suffering from neglect was transferred from
Union College to the Schenectady Museum Association during
the 1930's where it remained until 1959, essentially, in the con-
dition in which it was packed for transfer. The author was in-
vited by the director, Mr. Donald S. Smith, to remove to the
Museum of Comparative Zoology, Cambridge, Massachusetts,
any specimens that were, in his opinion, of scientific interest.
In this effort, Mr. Smith and Mr. Russell Carter, science curator,
offered every assistance and courtesy.
Apparently about 1860, Wheatley decided to specialize in
freshwater shells. This is attested to by his advertisements for
exchanges in the American Journal of Conchology. He there-
fore left only a token series of freshwater lots in this collection.
Of the species specifically mentioned by Lea as being in the
collection, only Anodonta schroteriana and Unio micans were
found.
In a published, though undated, sales catalogue issued, pre-
sumably, after Wheatley's death, a John A. Tydner of Philadel-
phia offered the freshwater collection for sale. He divided the
collection into two similar series which were offered for sale at
$2000 and $1750. From the information supplied by Dr. R. T.
Abbott, apparently both of these collections are now in the
Academy of Natural Sciences of Philadelphia. One is on "perma-
nent deposit" from the University of Pennsylvania and the other
was formerly the property of E. D. Cope.
Now in the collection of the Museum of Comparative Zoology
are some of the lots mentioned in the "Catalogue" of 1842,
though not many. There are 30 lots of Jamaican land snails from
C. B. Adams. (The main C. B. Adams collection is in the
Museum of Comparative Zoology) . Included are 62 type lots
from the collection of Hugh Cuming of species presumably sel-
ected by him and described by Pfeiffer, Broderip, Sowerby, Jonas,
Beck, and Reeve. Most of these species are Philippine land snails.
(For a discussion of the Cuming types see: Clench, W. J., 1945,
"Some Notes on the Life and Explorations of Hugh Cuming,"
Occ. Pap. Moll., Harvard Univ. 1, no. 3.)
From Bishop S. Elliott are 3 species of Unionidae described
by Isaac Lea and 2 species of Cuban land shells from Elliott
described by Poey. There are 6 more unionid types described by
Lea, 3 from W. A. Haines and 3 from other sources. Already
mentioned are the 2 type lots of the species from Wheatley
74 NAUTILUS Vol. 73 (2)
which Lea described. Finally, there are 3 type lots of land shells
described by A. A. Gould.
In all, the first Wheatley collection contained 108 type lots
in addition to which are almost 300 other lots with sufficient
data or historic interest to be worthy of retention.
TWENTY-FIFTH ANNUAL MEETING OF THE
AMERICAN MALACOLOGICAL UNION
The AMU. returned to its birthplace for this silver anniver-
sary meeting, for the Academy of Natural Sciences was where
the organization meeting was held in 1931. (Annual meetings
were suspended during World War II) . Of those present at the
first meeting, the following were on hand when, on Tuesday,
June 30, 1959, the twenty-fifth annual meeting was called to
order: H. B. Baker, Joshua L. Baily, Jr., William J. Clench,
Harald A. Rehder and Fred Tobleman.
The scene was Haverford College on the western outskirts of
Philadelphia, and there the members of the Philadelphia Shell
Club vied with one another to make comfortable the largest
assemblage in AMU. history. When the meeting was over, 133
had signed the register, and agreement was unanimous that it
had been an enjoyable and instructive four days.
Split sessions were held for the first time in AMU. meetings,
and in two halls President R. Tucker Abbott and Vice-president
Katherine Van Winkle Palmer presided as dual audiences heard
the following papers and talks:
"In search of Neopilina." Arthur H. Clarke, Jr.; "Sanibel
Island shell fair," Lulu B. Siekman; "Origin of the Land and
Fresh-water mollusks of the Bahamas," William J. Clench;
"Small beginnings," Adlai B. Wheel, Sr.; "Classification in the
Olividae," John Q. Burch; "Early Philadelphia conchologists,"
John D. Parker; "The strange phenomenon of autotomy in
Tremoctopus/' Richard W. Foster; "Mechanics of a shell club,"
Robert J. Wagner; "Some conchological miscellanies," Joshua
L. Baily, Jr.; "Expedition to the Philippines," John du Pont;
"Some techniques for anatomical work," Ruth D. Turner; "Shell
collecting around the world," Joseph J. Kuchar; "Marine
zoogeography, with special reference to the South Pacific area,"
Harald A. Rehder; "Our experiences photographing mollusks,"
Mrs. Theophil Kuczynski; "The egg mass and gross embryology
October, 1959 nautilus 75
of Pleurocera canalicidatum," Joseph Rosewater; "Mollusks, a
sound color movie"; "Unearthing Gould's types," Richard I.
Johnson; "Observations on Murex stainforthi and M. monodon,"
Virginia Orr; "Chromosomes of Pomatiopsis and Oncomelania,"
John B. Burch; "Quantitative sampling of aquatic mollusks,"
Charles B. Wurtz; "The Coosa revisited," Paul F. Basch; "The
Mollusca of Bernard Palissy, 1580," Aurele LaRocque; "Xan-
thonychidae (Pulmonata)," H. Burrington Baker; "Subspeciation
in Triodopsis," Joseph Vagvolgyi; "The mollusk fauna of Ba-
hamian mangroves," Robert Robertson; "Collecting at Broome,
Western Australia," Virginia Orr.
At a brief business meeting on Thursday morning, the fol-
lowing were elected officers for 1959-60:
President, Katherine Van Winkle Palmer; Vice-president,
Thomas E. Pulley; 2nd Vice-president, Chairman-incumbent,
AMU.-Pacific Division; Secretary-treasurer, Margaret C. Teskey;
Publications Editor, George M. Moore; Councillors-at-Large,
Alger P. Blaine, John E. Fitch, Richard Foster, Alan Solem.
It was announced that the council had accepted the invitation of
McGill University Museum to hold the 1960 meeting in Mon-
treal, Canada, in mid-August, and that the names of Dr. Julia
Gardner and Dr. Ralph Arnold had been added to the list of
Honorary Life Members, and that Dr. Paul Bartsch had been
elevated to Honorary Life President.
Other features of the meeting were the informal amateur sym-
posium of Tuesday evening, a showing of slides and movies of
former meetings following dinner on Wednesday, and an espe-
cially enjoyable social session which occupied all of Thursday
afternoon and evening.
Starting with a bus ride to the Academy of Natural Sciences
where exceptional exhibits of shells, shell products and books
arranged by members of the Philadelphia Shell Club awaited
inspection, guests were given a guided tour of the Department
of Mollusks, treated to a delightful tea by Mrs. Ruth Ostheimer
and Miss Anne Harbison, later were entertained by General
Frank and Dr. Jeanne Schwengel at the cocktail hour which
brought memories of many other such occasions to those who
have enjoyed the hospitality of the Schwengels in the past.
The annual dinner at 8:00 was held in the ANSP. auditorium
and made memorable by excellent steak, an inspirational talk
by Dr. Kenneth W. Prescott, Managing Director of the Museum,
76 NAUTILUS Vol. 73 (2)
and by individual place favors of pearl-bearing oysters (real
oysters, real pearls!) each tidily preserved in its own take-
home jar.
Following breakfast on Friday morning two chartered buses
transported those making the field trip to Cape May, New Jersey,
where a tour of the Snow clam canning factory, picnic lunch on
the Point, an afternoon devoted to collecting land and marine
shells and finally the long ride back to Haverford rang down
the curtain on the twenty-fifth annual meeting. — Margaret C.
Teskey.
NOTES AND NEWS
Zachrysia auricoma (Ferussac) in Miami, Florida. — Recently
Dr. H. A. Denmark, entomologist of the State Plant Board of
Florida, sent a series of land and freshwater mollusks from
Florida for routine determination. In this sending, there were
two species of Zachrysia: Z. provisoria (Pfr.) and Z. auricoma
(Ferussac). The record of Z. provisoria from Miami was reported
upon by Clarke (Naut. 70: 142, 1957) . So far as I can determine,
this is the first record of Z. auricoma outside of Cuba. A sub-
species, Z. a. havanensis Pilsbry, has been reported from Quinta,
Merida, Yucatan and the city of Panama, Panama, by Bequaert
and Clench (Publ. no. 457, Carnegie Inst, of Washington, p. 64,
1936).— W. J. Clench.
Green shells. — On an earlier page, J. B. Henrard states that
the calcareous layer in Liguus shells is green. In those before me,
it actually is whitish, but is so thin and translucent that the
green in the periostracum shows through and often is visible
from inside the aperture. — H. B. B.
iNTERTmAL STRANDING of CUoTie Umaciua in Massachusetts. —
On May 5, 1959, 29 living individuals of the gymnosomatous
pteropod, Clione limacina (Phipps, 1774), were collected from
pools in sand and mud and from Zostera beds where they were
stranded by the receding tide on the Lynn Harbor side of Little
Nahant, Essex Co., Massachusetts. To my knowledge, a published
record of this species occurring inshore on this coast has been
made only once in the past 91 years. Wood (1869, Proc. Port-
land Soc. Nat. Hist., 1, Pt. 2: 185-188) noted its presence in
October, 1959 nautilus 77
numbers in Portland Harbor from April 6 to May 7, 1868.
Previous to this DeKay (1843, Zool. of N. Y., Pt. 5, Mollusca,
p. 6) reported it from New York bays in April, 1883. Dr. J. H.
Welsh, Biological Laboratories, Harvard University, stated (per-
sonal communication) that he has collected this species at
Nahant on at least two occasions between 1938 and 1945, but
has not seen it again until this year. Possibly the species does
occur inshore more often than is generally supposed. However,
undoubtedly notice would have been made of these animals if
this occurrence were frequent because of the easily visible bright
red coloration of the organs which are outstanding in their deli-
cately opaque bodies. Probably onshore winds, ocean currents,
and spring tides during April and May have much to do with the
sporadic appearance of this pelagic species.
The usual habitat of Clione limacina is in surface waters of
the open ocean in boreal portions of the Atlantic and subarctic
Atlantic and Pacific oceans. In the western Atlantic, it is found
as far south as the latitude of Cape Hatteras (see J. J. Tesch,
1950, Dana-Report No. 36; also, references have been made to
Clione limacina on the English coast by J. E. Morton, 1958,
J. Mar. Biol. Ass. U.K., 37: 287-297 and M. V. Lebour, 1931,
ibid., 17: 785-795) . The only other report of the inshore occur-
rence of a gymnosomatous pteropod species in New England is
that of Danforth (1907, Proc. Bost. Soc. Nat. Hist., 34: (1): 1-19)
who described Paedoclione doliiformis which appeared in Casco
Bay, Maine, during August and September, 1902. — ^Joseph
RosEWATER, Museum of Comparative Zoology, Harvard Uni-
versity, Cambridge, Massachusetts.
Melongena egg cases. — Captive animals of Melongena corona
(Cf. Naut. 7i: 11-13) produced egg cases again, beginning on
June 23, 1959, when 9 cases, one with 112 eggs, were placed on
the side of the aquarium. — Ruth D. Turner.
PUBLICATIONS RECEIVED
New land Mollusca from the Admiralty and Bismarck Isl-
ands. By William J. Clench. Amer. Mus. Novit, no. 1863: 6 pp.,
7 figs. 1957. — Pseudocyclotus coultasi, P. incendium, Dendro-
trochus coultasi, Nesonanina unidentata and Sulcobasis (Goldie-
lix) fasciata (from Manus I.) are proposed as new. — H. B. B.
78 NAUTILUS Vol. 73 (2)
The land and freshwater Mollusca. By William J. Clench.
From "The Nat. Hist, of Rennell I., British Solomon Is." vol.
2:155-202, 3 figs., pis. 16-19. l9bS.—Palaeohelicina (s.s.) mayri,
Taheitia whitneyi, Setaepoma mayri, Nesopoma, genus of Assim-
ineidae with N. eyerdami (type) & N. galathea, Omphalotropis
nebulosa guppyi, O. quirosi, Charopa hoeyeri, C. insularis,
Quirosella, genus of "Ariophantidae," with Q. coultasi (type) ,
Q. wolffi %z Q. knudensi, Trochomorpha mcleani, Crystallopsis
{s.s.) crystallina, subgenus Cristovala (type Helix tricolor Pfr.) ,
with C. (C.) rennellensis, (vague genitalia) Eustomopsis hellon-
ensis, and E. renschi are new. The radular figs, of Quirosella
(like Nesonanina) apparently confuse the cusps on the laterals
with their basal thickenings. — H. B. B.
SoBRE "Turbonilla (Pyrgiscus) dispar" Pilsbry, 1897 (Gas-
tropoda, Pyramidellidae) . By H. de Souza Lopes. Sobre un novo
gastropodo brasileiro do genero "Solariella" Wood, 1842 (Tro-
chidae) . By Lopes & Paulo de Sa Cardoso. Rev. Brasil. Biol.
18(1) :17-21, 11 figs., and 59-64, 3 figs. \9b%.—Solariella carvalhoi
is new, from 57 meters depth, S. Paulo. — H. B. B.
Note sur Drepanotrema anatinum et Taphius peregrinus
(Pulmonata, Planorbidae) . By W. Lobato Paraense 8c Newton
Deslandes. J. de Conch. 775:152-162, 22 figs. 1958.— As usual with
the authors, the genitalia, radulae and shells are figured ex-
quisitely.— H. B. B.
System and phylogeny of Planorbidae (Gastropoda Pul-
monata) . By Ja. L Starabogatov. Bull. Moskov. O-va. Isp. Prirod.
(Soc. Investigators of Nature), Biol. 63 (6): 37-53, 4 figs. ^ phylo-
genetic scheme. 1958. — In the family Planorbidae are included
the typical subfamily (tribes sens. str. & Segmentinini) , Planor-
bulinae and Biomphalariinae (tribes s. s. ^ Acrorbini, new) . In
Camptoceratidae (^ Bulinidae) are recognized Camptocera-
tinae (= Coretinae -\- Helisomatinae) , Plesiophysinae, Bulin-
inae and Physastrinae, new (^ Miratestinae) . Figured ovotestes,
prostates and jaws are used for the separation of the major
divisions. — H. B. B.
About anatomy and systematics of subgenus Carinogyraulus
(Gastropoda, Planorbidae) . By Ja. I. Starabogatov. Nauchn.
Dokl. Byssh. Shkol. (Sci. Rept. of Superior School), Biol. no.
4:16-20, 4 figs. 1958. — In this subgenus of Anisus, genitalia,
radulae, jaws and pallial complexes of "Gyraulus" (C.) trape-
October, 1959 nautilus iii
zoidesrohustonudus, G. (C.) paradoxus 2ind G. (C.) lychnidicus
are described and figured. — H. B. B.
Materialen zur Kenntnis der Zonitiden (Gastropoda) des
Kaukasus und der Krim. By Adolf Riedel. Ann. Zool. Polska
Akad. Nauk 77(11) :383-427, pis. 29-30, 1 map, 14 textfigs. 1958.
— The morphology and distribution of one Crimean and 4 Cau-
casus species of OxycJiilus are studied carefully. These are dis-
tributed in subgenera Schistophallus, Cellariopsis, Morlina and
(new) Longiphalhis, type Helix filicum Krynicki. — H. B. B.
On the molluscan adhesive epithelium. By Bengt Huben-
dick. Arkiv for Zool. 0 (7118) :001-006, 1 fig., pis. 1-3. 1957.— In
Ancylus lacustris, the cells at the retractor attachments have
microvilli, which apparently fit into minute shell depressions. —
H. B. B.
The development of the penial stylet in Gyraulus (Moll.
Pulm.). By Bengt Hubendick. Arkiv for Zool. 11 (24):427-429, 4
figs. 1958. — All the stylet is secreted by surrounding epithelia,
and later protrudes. — H. B. B.
A NOTE on Protancylus p. & F. Sarasin. By Bengt Hubendick.
Beaufortia d (78) : 243-250, 9 figs, 1958.— Additional anatomic
data indicate that this genus belongs in the Planorbidae. —
H. B. B.
A NOTE ON THE TAXONOMY OF THE BRAZILIAN VECTOR SNAILS of
"Schistosoma mansoni." By Bengt Hubendick. Rev. Brasil. Biol.
7(9(1) :37-40. 1958.— Prefers Biomphalaria to Taphius. Again,
why not Planorbina? Or Australorbis, which is in most general
use?— H. B.B.
Studies of the genus Gundlachia (Pulmonata, Ancylidae) .
By Paul F. Basch. Occas. Papers Mus. Zool. Univ. Mich. no. 602,
9 pp., 2 figs. 1959. — The status of the genus (sensu lato) remains
uncertain, but the radula of G. (Kincaidilla) meekiana is unlike
that of the typical group. It is like that of Laevapex, 1903, which
unfortunately was made prior to Ferrissia, 1903, by Hannibal,
1912.— H. B. B.
Preliminary study of newly hatched oyster drills, Urosal-
pinx cinerea (Say) . By Melbourne Romaine Carriker. J. Elisha
Mitchell Sci. Soc. 75 (2):328-351, 10 figs. 1957.— Reactions of
baby drills to water currents, light, gravity, substrata, evapora-
tion, and emanations from young hard clams were tested in the
laboratory. They were voracious and very active. — H. B. B.
x^^^C'^rx
WILLIAM H. WEEKS SHELL COLLECTION: Now being of-
fered for sale. To receive free lists, send name and address to:
George E. Jacobs, 853 Riverside Drive, N. Y. 32, N. Y.
A CHECK LIST OF THE MARINE SHELLS OF ST. CROIX,
U. S. Virgin Islands, with random annotations. Up to date.
Lists 650 species plus 60 subspecies, including 20 new. With
70 illustrations. The "random notes" contain much new infor-
mation. Price: $4.10, post free.
G. UsTiCKE, 1 North Street, Christiansted, St. Croix, Virgin Is.
How TO COLLECT SHELLS: Published by the American Malacological Union.
$1.00. Write:
Margaret C. Teskey, Sect., Route 2, Box 318, Marinette, Wis.
Directory of conchologists. — The 1960 edition will be published the first of
that year. Price will be $2.50, postpaid. For inclusion, write:
John Q. Burch, 4206 Halldale Ave., Los Angeles 62, Calif.
THE NAUTILUS
Vol. 73 JANUARY, 1960 No. 3
CARIBBEAN SPECIES OF TRUNCATELLA
Bv ALFREDO DE LA TORRE
Guggenheim Fellow, Division of Mollusks
U.S. National Museum, Washington, D.C.
Pfeiffer (1839:356) in the report about his trip to Cuba,
describes the species Truncatella pulchella as follows: "Testa
imperforata ovato-cylindrica, gracili, pellucide fulva; anfract. 4
minutissime striatis, ad suturam subcrenulatis, ultimo semis-
triato; peristomate crasso, albido; apertura subelliptica. Long.
2, diam. 14'". Operculum tenue corneum."
In studying this description, I conclude that this is a species of
about 4.36 mm. length (= 2 German lines) , with rather obso-
lete axial sculpture, and a thick peristome.
One year later (1840:253), in the continuation of the report,
the same author describes T. bilabiata as follows: "Testa cylin-
dracea solidula, niti ce carnea, longitudinaliter confertim cos-
tulata; anfract. 4i/2 convexis; sutura profunda; apertura obliqua
ovali peristomate incrassato, duplicato. Long. 2i/2 diam., 34 lin."
This species is, according to the description, a little larger
than pulchella, being of 2i/2 German lines length (= 5.45 mm.) ,
and having closely spaced longitudinal ribs.
Clench and Turner (1948:151) write the following about
this species: "Our thanks are due to Dr. C. G. Aguayo for the
gift of several lots of Truncatella from Cuba and for the loan
of a type series of T. pulchella Pfeiffer, a series collected origi-
nally by Gundlach." In the same publication (pp. 156-158) these
authors describe what they consider to be pulchella Pfr., based
on a lectotype (Museo Poey 181, from Cardenas, Cuba) selected
by them, and mention also other specimens considered by them
to be paratypes, from the same locality, saying that: "Additional
types are probably in the Museum of Stettin, Germany." (Loc.
cit. p. 157).
In the same publication (p. 157) they stated that "T. pul-
chella is readily separated from T. bilabiata by being larger and
having a simple lip," giving for their T. pulchella a length of
79
80 NAUTILUS Vol. 73 (3)
5.5 mm. (small specimens) to 7.3 mm. (large specimens) . These
dimensions do not agree with the original description by
Pfeiffer of pulchella. This species is, according to Pfeiffer some-
what smaller than bilabiata and much smaller than the species
supposed to be pulchella by Clench and Turner. On the other
hand, the T. pulchella of Clench and Turner (not of Pfeiffer)
is in perfect agreement in the characters, dimensions and figures,
with T. caribaeensis "Sowerby" Reeve (1842:94, pi. 182, f. 2).
The last name is, as far as I know, the earliest one available
for this larger species of Truncatella of the Caribbean region.
At the same time, I have proved to my satisfaction that T.
succinea C. B. Adams (1845:12), described from Jamaica, is
absolutely identical with T. caribaeensis "Sowerby" Reeve, which
as the earlier name has priority. Dr. Morrison, who helped me
in the preparation of this publication, agrees with me in these
opinions.
As stated above, I have arrived at the following conclusions:
(1) T. bilabiata Pfr. is only the strongly ribbed form of the
previously published species T. pulchella Pfr. (2) The species
called T. pulchella, by Clench and Turner in 1948 is not pul-
chella Pfr., and must be known as T. caribaeensis "Sowerby"
Reeve, which is apparently the earliest name available for that
species. It is larger than pulchella Pfr., with the outer lip of the
peristome simple, and the lip usually thin.
In agreement with the foregoing, I have been able to examine
numerous lots of Truncatella from Cuba and other Caribbean
regions which prove that T. pulchella Pfr. (non Clench and
Turner) is very variable, going from a smoothish form with
the duplex condition of the outer lip and peristome in a very
reduced or obsolete condition, to a form presenting strong
axial ribs and with the duplex condition of the outer lip very
prominent. This last form is what properly may be called
T. pulchella, form bilabiata Pfr.
The typical T. caribaeensis "Sowerby" Reeve has well marked
axial riblets and so is identical with T. succinea C. B. Adams; the
smoothish form of this species was improperly called T. sub-
cylindrica Gray, by Pfeiffer, in his writings. The true species
T. subcylindrica (Linne) (= laevigata Risso) is a European
species.
January, 1960 nautilus 81
Going back to the problem of T. pulchella Pfr., I must point
out that Clench and Turner (loc. cit. p. 151) are in error in
considering a series of T. pulchella, from Cardenas, Matanzas
Province, Cuba, collected by Gundlach (from which these
authors have selected a lectotype) as typical. As I have stated
in previous paragraphs, I do not agree with them in consider-
ing this a type series of the species. At the same time I also
suspect that this locality can not be considered the type locality
(see A. Torre, 1952:19, also pp. 9 and 10) . As I stated in 1952,
Pfeiffer, during his trip to Cuba in 1839, visited only a few
localities in the neighborhood of Havana and Matanzas and
remained in Cuba only three months. So, the "neighborhood
of the city of Matanzas" was probably the locality from which
he collected the type specimens of T. pulchella; not from
Cardenas. It was only in 1841, two years after the publication
of the original description of pulchella Pfr., that "Gundlach
moved from Tl Fundador de Canimar,' Matanzas, to the Tinea
San Juan,' in Cardenas, extending in this way his area of explora-
tions" (A. Torre 1952:10).
The year 1856 (loc. cit. p. 193) is when Pfeiffer gave the
localities in Cuba, for T. pulchella as "prope Matanzas et
Cardenas;" evidently collected by Gundlach at the last locality
on a date posterior to the original publication of the species.
Again, in 1857 (pp. 118-119) Pfeiffer lists the species of Trunca-
tella of Cuba, giving under pulchella Pfr., the same localities
as above; "Cardenas and Matanzas, in the same living conditions
as subcylindrica."
My only conclusion is that the selection of a lectotype by
Clench and Turner for this species is inappropriate because the
lot from which it was selected is not a series of pulchella com-
ing from the original material collected by Pfeiffer and Gundlach
in 1839. What is more important, it is not even a specimen of
the species pulchella Pfr., but represents caribaeensis "Sowerby"
Reeve, which is a different and larger species.
At the same time and in accordance with the foregoing state-
ments, I am here restricting the type locality of T. pulchella Pfr.
to the "vicinity of the city of Matanzas" and am considering
the selection of a lectotype by Clench and Turner as invalid.
It would be very interesting to know if there are typical speci-
82' NAUTILUS Vol. 73 (3)
mens of this species in the Museum of Stettin, Germany, as
Clench and Turner suspect, and then to study them and get
more detailed information about the type locality.
In 1856 (pp. 175-196) and again in 1857 (pp. 118-119)
Pfeiffer listed the species of Truncatella of the Island of Cuba;
the data given in both publications are in agreement with my
conclusions. In the publication of 1857, the species listed for
Cuba are the following: T. carihaeensis (with succinea as a
synonym) ; T. subcylindrica (Gray) Pfr. [this is not, as I state
in another part of this paper, the species subcylindrica (Linne) ,
which is a European species, but only the smooth form of
carihaeensis Reeve] ; T. bilabiata ("collected under fallen leaves
of Coccoloba, at the mouth of Canimar River, Matanzas, Cuba,")
which is the type locality of this form (see A. Torre 1952:19) ;
T. scalar is Michaud (Pfeiffer himself gives his T. costata as a
synonym) , collected at Matanzas, Cuba. The other two species
listed as Truncatella (elongata and lirata Poey) , are now known
to be Geomelania and not Truncatella.
Pfeiffer, who was named "The Prince of Malacologists," was
evidently a brilliant student of the genus Truncatella and his
writings on this genus merit the most careful consideration.
I must cite also, in support of the conclusions herein adopted,
the work by Kiister (1855). In this publication, the figures and
references given by the author absolutely support what I have
stated in this paper. Also, W. G. Binney (1859:184-190) cor-
rectly described and figured T. pulchella Pfr., declaring on page
185 that "he had sent to Pfeiffer and Poey the Florida specimens
of Truncatella and these were identified by them."
Before concluding these notes, I wish to make reference to
the species T. barhadensis Pfr., listed by Clench and Turner
(loc. cit. p. 153) in the synonymy of T. bilabiata Pfr. This
species. Dr. Morrison has proved to me, is a good species, geo-
graphically limited to Barbados, W.I., and perhaps other lesser
Antillean Islands.
Also I wish to refer to T. scalaris scalaris Michaud, and T.
scalaris clathrus Lowe, which are distinguished from one another
by the number of axial ribs on the body whorl, there being
8 to 11 in the typical form and 12 to 16 in clathrus. Since this
character is variable and the two forms overlap on some of the
January, 1960 nautilus 83
West Indian Islands, probably they can be considered only as
simple forms of the same species, instead of as subspecies.
Genus Truncatella Risso.
Truncatella Risso, 1826, Hist. Nat. de I'Europe Merid. 4, p.
124.
Type species: T. laevigata Risso (^ Helix subcylindrica
Linne) SD., Gude 1921 (According to Clench and Turner,
1948) .
Subgenus Truncatella s. s.
Truncatella pulchella Pfeiffer.
T. pulchella Pfr., 1839, p. 356 (Cuba) . Kiister, 1855, pp. 10-12,
pi. 2, figs. 11-15. Pfr., 1856a, p. 192-193 (Cuba, near Matanzas
and Cardenas; Puerto Rico, Jamaica and St. Thomas [Gives
as dimensions 41/9 mm. length X 1^9 to 2 mm. width]. Binney,
1859, p. 189, pi. 75, fig. 10 (W.I. and Fla.) . Binney, 1865, p. 99.
T. bairdiana C. B. Ads., 1852, Ann. Lyceum Nat. Hist. N. York
5, p. 437 (p. 213 in separate) ([West] Panama). Clench and
Turner, 1948a, p. 153, pi. 66, fig. 7 [Occurrence in West Panama
due to accidental transport from east Coast?].
T. capillacea "Gundlach" Pfr., 1859, p. 77 (Caimanera,
Guantanamo, Cuba). Clench and Turner 1948a, pi. 66, fig. 6
(Same locality) .
T. bilabiata Pilsbry, 1948 p. 1069, fig. 571b (in part) . Clench
and Turner, 1948a, p. 153 (in part).
Type locality: "Vicinity of Matanzas City, Matanzas Prov.,
Cuba" (herein restricted) . Other localities: In Cuba: Mantan-
zas and Cardenas (Ph.) ; Playa Bellamar, Matanzas (A. Torre!) ;
"El Embarcadero," Finca Bacunagua, Los Palacios, P. del Rio
(A. Torre!) ; also, Jamaica, Haiti, Florida (East and West
Coast) , Bahamas and St. Croix (USNM) .
Truncatella pulchella Pfr., form bilabiata Pfr.
T. bilabiata Pfr., 1840, p. 253 (Cuba. 1856, p. 192 (Cuba:
"Right bank of the mouth of the Canimar River, Matanzas
(Gundlach) , but not in the Island of Carmen, Gulf of Mexico
(Kiister)." Plsbry 1948, p. 1069, fig. 571a (in part). Clench and
Turner 1948a, p. 153-155 (in part) . A. Torre, 1952, p. 19. [The
type locality is fixed, according to previous restriction by Pfeiffer
in 1856].
Type locality: Right bank of the mouth of the Canimar
River, Matanzas, Cuba. Other localities: In Cuba: Baracoa and
Gibara, Oriente (Gundlach) (Ace. to Pfr.) ; "El Embarcadero,"
Finca Bacunagua, Los Palacios, P. del Rio (A. Torre) ; Buey-
vaquita, Matanzas (1 specimen, dead, in sand dredged) (A.
84 NAUTILUS Vol. 73 (3)
Torre!) ; Playa de Bellamar, Matanzas (A. Torre!) (Very com-
mon in this last locality in a cave in the seashore, etc.).
In the lots of T. pulchella Pfr. from Jamaica, Haiti, Florida,
Cuba, Bahamas and St. Croix, that I have studied at the USNM.,
there are usually mixed specimens of the typical smoother form
(pulchella), and of the costate form (bilabiata), but generally
the smoother form is scarce and the costate form very abundant.
From the Bermuda Islands there are several lots, but, in these,
all the specimens observed by me belong to the costate form or
bilahiata Pfr. The records given by Clench and Turner as
bilabiata (loc. cit.) from Barbados, W.I., really belong to the
species barbadensis Pfr. The species pulchella and the form
bilabiata apparently do not live at that locality.
Truncatella caribaeensis "Sowerby" Reeve.
T. caribaeensis "Sowerby" Reeve, 1842, p. 94, pi. 182, fig. 7
(not fig. 2, as inadvertently given by Clench and Turner in 1948)
(No locality given.) Kiister, 1855, p. 9, pi. 1 figs. 35-37 and pi.
2 f. 1-4 [these last four figures correspond to the smoother va-
riety, improperly called subcylindrica Gray, by Pfeiffer] (West
Indies, Mexico and Alabama). Pfeiffer, 1856, p. 185 (Cuba and
Jamaica). Pfeiffer, 1857b, p. 119 (Vicinity of Matanzas, Cuba
and Island of Jamaica) .
T. succinea C.B. Ads., 1845, Proc. Boston Soc. Nat. Hist. 2,
p. 12 (Jamaica). Pfr., 1846a, p. 118.
T. goiildii Pfr., 1846a, p. 118, Pfr., 1846b, p. 183 |in synonymy
of caribaeensis Reeve].
T. caribaeorum Pfr., 1846a, p. 118.
T. variabilis Pfr., 1846b, p. 183 [nude name in the synonymy
of T. caribaeensis} .
T. guerinii "Parreyss" Pfr., 1856 p. 185 [nude name in the
synonymy of T. caribaeensis].
T. subcylindrica (Gray) Pfr., 1856, p. 186-187 (in part)
(Cuba: Matanzas and Cardenas) ; (Puerto Rico, St. Thomas
Bermuda, etc.). Pfeiffer 1857b, p. 119 (^= caribaeensis Kiister,
1855, T. 2, f. 1-4, sec. Pfr.) Cardenas, Matanzas Prov. Cuba).
Non T. subcylindrica Linne, 1767, Syst. Nat. 12th. Ed. p. 1248
(Europe) . [This is the smooth form of caribaeensis; typical
caribaeensis being regularly axially striate].
T. pulchella Pilsbry, 1948, p. 1070, fig. 572b, c. Clench and
Turner 1948a, p.l56, pi. 68, f. 1-6 [Non pulchella Pfr. 1839].
T. pulchella form caribaeensis Pilsbry 1948, p. 1071, fig. 572 a.
Type locality: Cuba (?) (As I have not seen the reference by
Pfeiffer in 1846 (p. 183) in which this author probably gives
Cuba as locality, I am considering this as type locality with
January, 1960 nautilus 85
doubt, until I can check that reference. Other localities: North
Carolina, Fla., Texas, Bermuda, Bahamas, Cuba (Playa de
Manimani, P. Rio) ; Cardenas (Varadero) , Matanzas; punta
Alegre, Camagiiey; Punta de Piedra, Banes, Cabo Cruz, Oriente;
Isla de Pinos) ; Porto Rico, Virgin Islands (St. Thomas and St.
Croix); Jamaica; Hispaniola (Haiti) ; Lesser Antilles (St. Bar-
tholomew, Martinique, Guadeloupe, Barbados, Trinidad, Cura-
cao); all given by Clench and Turner, 1948: 157-158), (under
the name pulchella). Bahamas, Cuba (Cabo Cruz, Matanzas);
Hispaniola (Haiti) ; Jamaica; Cayman Ids. (Given by the same
authors under the name succinea C. B. Ads., loc. cit. p. 159).
Playa de Bellamar, Matanzas, Cuba (A. Torre!) (Common) ;
Matanzas (Cuba) and Jamaica (Pfr.) ; Cardenas, Matanzas
Prov. (Pfr.) ; W. Indies, Mexico and Alabama (Kiister).
In the USNM, there are records from Cuba of several locali-
ties: Isla de Pinos; Little Cayman; Jamaica; Hispaniola (Haiti);
Porto Rico; Virgin Is.; Guadeloupe; Trinidad; Bermuda; Flor-
ida (East and west coasts) ; Texas; Bahamas and St. Croix.
Subgenus Tomlinitella Clench and Turner.
Tomlinella Clench and Turner, 1948a, p. 159 [non Tomli-
nella Viader 1938, Bull. Mauritius Inst. 1: 6].
Tomlinitella Clench and Turner, 1948 b, p. 169.
Type species: Truncatella scalaris Michaud, (OD.)
Truncatella scalaris (Michaud) .
Rissoa scalaris Michaud, 1830, Descr. Genre Rissoa p. 18
(Locality unknown) .
Truncatella costata Pfr., 1839, p. 356 (Cuba).
Truncatella cumingii C.B.Ads., 1845, Proc. Boston Soc. Nat.
Hist. 2, p. 12 (Jamaica). Pfeiffer, 1846a, p. 119 [declared iden-
tical to T. costata Pfr.]
Truncatella scalariformis C.B.Ads., 1845, loc. cit. p. 12 (Ja-
maica) (Non Reeve, 1842) (Ace. to Clench and Turner, 1948a:
160).
Truncatella adamsi Pfr., 1846a & 1846b, pp. 119 k 189 [New
name for scalariformis C.B.Ads., non Reeve 1842].
Truncatella scalaris Pfeiffer, 1856, p. 194-195 (Cuba and
Jamaica) .
Truncatella (Tomlinella) scalaris Clench and Turner, 1948a,
p. 160-161, pi. 71, fig. 1-4 [figures 5 and 6 of the same plate
probably belong to the subspecies clathrus Lowe.]
Truncatella (Tomlinitella) scalaris Clench and Turner 1948b,
p. 169.
86 NAUTILUS Vol. 73 (3)
Type locality: Port Antonio (Jamaica) (Neoholotype desig-
nated by Clench and Turner from that locality 1948a; 161).
Other localities: Cuba; Hispaniola (Haiti) ; Jamaica (Clench
and Turner); Matanzas, Cuba (Pfr.); Jamaica (Pfr.) ; Playa
de Bellamar, Matanzas, Cuba (scarce) (A. Torre!) . Haiti,
Jamaica and St. Croix (USNM.) (Teste Morrison) . (See also
under T. scalaris clathrus).
Truncatella scalaris clathrus Lowe.
T. clathrus Lowe 1832, Zoological Journ. 5, p. 303 (Locality
unknown). Reeve, 1842 p. 94, pi. 182, fig. 3 (no locality given)
[This is the first figure of this species]. Shuttleworth, 1852, p.
155 (Porto Rico and St. Thomas) . Pilsbry 1948, p. 1069, f. 571c.
T. clathra Pilsbry 1900, p. 506, pi. 62, fig. 13 (Bermuda Ids.,
Porto Rico and St. Thomas.)
T. scalaris piratica Clench and Turner 1948a, p. 161, pi. 72,
fig. 1-4 (Bermudas) . (Teste Morrison).
T. (Tomlinitella) scalaris piratica Clench and Turner, 1948b,
p. 169.
Type locality: (?) according to the literature (Shuttleworth,
1852: 155) the restriction of the type locality to Puerto Rico
seems possible, as this author cites this form from Puerto Rico
and St. Thomas (collected in the last locality by Blauner) . But
we have not seen a single specimen of T. scalaris (Michaud)
or of T. scalaris clathrus Lowe from either one of those two
localities, so, we are in doubt if this species or the form clathrus
really live on Porto Rico and St. Thomas or not. Other locali-
ties: Bermuda Islands (Clench and Turner); Cuba, Haiti, Flor-
ida, Bahamas. The localities in Cuba being Varadero, Matanzas;
Bahia de Cabanas (P. Rio) (USNM. collection) . Also: Playa
de Bellamar, Matanzas (in sand, mixed with T. scalaris scalaris)
(Michaud) both of them being rare (A. Torre!) .
This form is distinguished from the typical one because it has
12-16 axial ribs in the body whorl, instead of 8-11 as in the
typical scalaris (Michaud) . According to Dr. Morrison, these
two forms seem to have a different geographic distribution,
although they overlap in Haiti, and, according to specimens
collected by the writer they overlap in Cuba also. Dr. Morrison
thinks that the specimens coming from Florida and the Bahamas
in the collection of the USNM. all correspond to the form
clathrus and none to the typical scalaris Michaud. I keep them
separate, as distinct subspecies, although they possibly may
prove to be only different extreme forms of the same species.
January, 1960 nautilus 87
Truncatella bahamensis Clench and Turner.
T. bilabiata bahamensis Clench and Turner 1948, Johnsonia,
vol. 2 no. 25, p. 155, pi. 67, f. 1-3.
Type locality: Northwest Point, Little Inagua, Id., Bahama
Ids. Other localities: Limited to the Bahama Islands.
Truncatella barbadensis Pfeiffer
T. barbadensis P£r., 1856, p. 192 (Barbados). Pfeiffer, 1857a,
p. 337 (Barbados, W.I.) .
T. bilabiata Clench and Turner 1948a, p. 153-155 (in part).
Type locality: Barbados, Lesser Antilles. Other localities: in
the collection of the USNM. There are records of this species
from several localities in Barbados and also, records of a small
form of the species from Guadeloupe and Antigua (teste Mor-
rison) . The records given by Clench and Turner in 1948 (p. 155)
for bilabiata, from Barbados, probably correspond to this species,
these authors giving the name barbadensis as a synonym of
bilabiata Pfr. I think that they are really different species,
barbadensis being always strongly axially ribbed as well as
different in shape and other characters. The previous study by
Dr. Morrison on specimens of the collection of the USNM. is in
perfect agreement with my conclusions here.
I wish to acknowledge here my gratitude to Dr. Joseph P. E.
Morrison, for the valuable help given to me in preparing these
notes and list.
Key to the western Atlantic and Caribbean species of Truncatella.
Shell small (3-6 mm. total length) . Outer lip generally duplex
Costae 17 or more on the body whorl; no microscopic spiral
sculpture between the ribs.
Shell smoothish. Duplex condition of the outer lip some-
what obsolete T. pulchella.
Shell not smoothish. Duplex condition of the outer lip
well marked
Shell narrower and more cylindrical and elongated than
the following group (4 to 5 remaining whorls in the
adult shell) .
Axial ribs numerous and close together (about 40 on
the body whorl) T. barbadensis.
Axial ribs generally less numerous and more widely
spaced than in the preceding species (about 20 on the
body whorl) T. bahamensis.
Shell shorter and less cylindrical (3 to 4 remaining whorls
in the adult shell) T. pulchella, form bilabiata.
Costae 16 or less on the body whorl; microscopic spiral sculp-
ture between the ribs.
88 NAUTILUS Vol. 73 (3)
Costae 8 to 1 1 on the body whorl T. scalaris.
Costae 12 to 16 on the body whorl T. scalaris clathrus.
Shell larger (6 to 8 or 9 mm. in length) . Outer lip simple. Axial
costae well developed or more or less obsolete T. carihaeensis.
Bibliography
Binney, W. G. 1859. The terrestrial mollusks of the United
States and the adjacent territories of N.A., vol. 4, pp. 1-207,
pis. 75-80.
1865. Land and fresh-water shells of N.A., Pt. 3, pp. 1-220.
Clench, W. J. k R. D. Turner, 1948a. Johnsonia, vol. 2 (25),
pp. 149-164, pi. 63-73
1948b. Occ. Papers Moll., Harvard, vol. 1 (13), pp. 157-212.
Kiister, H. C. 1855. Syst. Conchyl. Cab., Truncatella & Paludin-
ella, pp. 1-20 pis. 1-2.
Pfeiffer, Ludwig. 1839. Archiv. f. Naturg. 5th yr., vol. 1, pp.
346-358.
1840. Archiv. Naturg., 6th. yr., vol. 1, p. 250-261.
1846a. Zeitschr. Malak., vol. 3, pp. 1 J 3-120.
1846b. Ibid., pp. 177-190.
1856. Monogr. Auricul. Viv., app. 2, pp. 175-196.
1857a. May 8. Proc. Zool. Soc. London, pt. 24, no. 322, pp.
336-339.
1857b. Malak. Blatt., vol. 3, pp. 118-119.
1859. ALilak. Blatt., vol. 6, pp. 66-102.
Pilsbry, H. A. 1900. Trans. Conn. Acad., vol. 10, p. 491-509, pi.
LXIL
1948. Land Moll, of N. America, vol. 2, pt. 2, pp. 1065-1074,
figs. 569-573.
Reeve, Lovell. 1842. Conchologia Systematica, Vol. 2, London,
337 pp., pis. 130-300.
Shuttleworth. R. J. 1852. Diagnosen neuer Mollusken, no. 1
(May 1852), Bern, 163 pp.^
Torre, Alfredo de la. 1952. Rev. Soc. Malac. "Carlos de la Torre",
vol. 9 (2), pp. 9-25, pis. 1-4.
ANATOMY OF RHODACMEA CAHAWBENSIS WALKER,
1917, A RIVER LIMPET FROM ALABAMA
By PAUL F. BASCH
Department of Biology, Kansas State Teachers College, Emporia
The genus Rhodacmea was established by Bryant Walker in
1917 to include a number of limpet-like freshwater mollusks,
found mainly in the southeastern United States. Heretofore,
nothing has been known of the anatomy of these snails besides
the shell and radula, both of which present distinctive features.
NAUTILUS 73 (3)
PLATE 7
I mm.
B
I mm.
250 micro
\- Shell of juvenile specimen of Rhodacmea cahaxcbensis, dorsal view;
note depression^ near lefl side of apex. B Shell of adu U ^pec^n n of ^.
cahaivbensis, rieht side view; note erosion of apical region^ C. \ entiai view
of head region. D: Jaw, dissected from an adtdt specimen, flattened. E. Cen-
tral and fi?st 7 lateral teeth of radula; marginals not shown.
NAUTILUS 73 (3)
PLATE 8
mantle
salivary gland (cut)
penial complex
brain
esophagus -
vagina
spermattiecal
duct -^
vas deferens ^ ^
prostate
buccal
mass
intestine
ovotestis
mantle^ ^--""'cr'-^
male genital opening^ ^5*''*''***^'Ci^
female genital opening
rectum
pseudobranch
A: Central nervous svstem of RJwdacmea cahcnvbensis. B: Superficial dis-
section of animal to show relative positions of organs in life. C: Deeper
dissection; most of the digestive and reproductive systems removed. Note
positions of shell muscles. D: View of left side of adult specimen, shell and
portion of mantle containing kidney removed. Note position and character
of pseudobranch.
January, 1960
NAUTILUS 73 (3)
NAUTILUS
89
PLATE 9
female genital opening
penial complex
-male genital opening
A: Penial complex of Rhodacmea cahawbensis, semidiagrammatic B- Sec-
tion of vas deferens, through plane b-b' on A; note ciliated lumen. C- Sec-
tion through penis and penis sheath at plane c-c' on A. D: Section through
penis sheath at plane d-d' on A. E: Reproductive system of R. cahawbensis
dissected out and separated to show relationships of organs
90 NAUTILUS Vol. 73 (3)
The apex of the shell in all species is tinged with a reddish
coloration, and a thick callus of shelly material usually is
found within the apical region. The enormous, unusually
shaped lateral teeth of the radula (pi. 7, fig. E) , the small mar-
ginals and the small number of teeth per row have cast doubt
upon the validity of associating Rhodacmea with the family
Ancylidae. The present investigation was undertaken to de-
termine the proper systematic position of this genus through
anatomical study.
The specimens used were collected by the author and Dr. John
B. Burch on June 6 and June 8, 1959, from stones and dead
mussel shells in rapids of the Cahaba River west of Helena,
Section 19, T 19S, R 4 W, Shelby County, Alabama. Identifica-
tion as R. cahawbensis w^as made by comparison with the holo-
type specimen, No. 102635, in the collection of the University
of Michigan Museum of Zoology. The type locality for the
species is at Gurnee, Shelby County, Alabama, about 8 miles
south of the Helena location. Specimens from the present col-
lection have been deposited in the University of Michigan
Museum of Zoology.
I am greatly indebted to Dr. Henry van der Schalie for making
the collecting trip possible and for the generous use of his facil-
ities, and to Dr. Harold W. Harry, of the University of Alabama,
Birmingham Center, for invaluable assistance in the field.
In spite of unusually high waters which made collecting diffi-
cult, sufficient specimens were secured to make a thorough ana-
tomical study of R. cahaivhensis possible. The animals were
placed in a 1% solution of Nembutal until relaxed, fixed in
the iield with formalin-acetic acid-alcohol fixative, and stored
in 70% alcohol. A few specimens were brought back to Ann
Arbor alive, but they did not survive more than four days
even when packed in ice. Gross dissections were made of 8 spec-
imens, and one was embedded in paraffin and sectioned serially
at 10 microns, for histological study. Radulae of adult and
juvenile animals were removed, cleaned in dilute sodium hypo-
chlorite solution, stained with Orange G, and mounted in
Canada Balsam or Euparal.
Shell: The shells of the juvenile specimens are distinctly
radially striate, and show a characteristic pit-like depression on
January, 1960 nautilus 91
the left side of the apex (pi. 7, fig. A.) The precise form of this
depression varies somewhat among the specimens collected, but
it is present in all. Both the depression and striae usually are
lost in the adult when the apex becomes eroded, and are not
apparent on the holotype. Both of the characters, however, are
present in adult UMMZ. specimens of Rhodacmea filosa (Con-
rad) from the Coosa River, Alabama, and in certain lots identi-
fied as R. elatior (Anthony) from Eagle Creek, Owen County,
and Green River, Hart County, Kentucky. The utility of these
apical characters as taxonomic markers is unfortunately dimin-
ished by the erosion so frequently encountered in certain species.
Juvenile shells are a rusty brownish color when collected, and
shells of the adults (pi. 7, fig. B) are black, often with tiny
clumps of algae growing on them. When cleaned with dilute
oxalic acid, the periostracum is revealed as a thin, light corneous
layer over the shell, and in those specimens in which the apex
is at least partially retained, the pink coloration becomes
evident.
External morphology: The animal of R. cahawbensis is super-
ficially bilaterally symmetrical, resembling other ancylid genera
in this respect. The head and foot are a light gray color, and the
translucency of the tissues permits the color of the internal
organs to show through. The relatively large head bears promi-
nent eyes and much reduced tentacles, through the thin integu-
ment of the front of the head the pink buccal mass is clearly
visible. Beneath the shell, the color of the dorsal surface of the
mantle is quite variable. In some specimens, it is uniformly gray
with a concentration of pigment near the periphery giving the
effect of a black ring, in others pigment is almost entirely absent
or may appear only as irregular black blotches. The pseudo-
branch is small and simple, without folds or ridges, and bears
the rectum and anus. The foot is broad, flat, and oval in out-
line, as in other ancylids.
Integument and muscles: The integument of R. cahawbensis
presents no peculiarities, and histologically is similar to that
described by Basch (1959) for Laevapex fusciis. The foot and
velar muscles are also similar to the latter species, but the right
anterior shell muscle in all specimens examined extends a much
greater distance caudally than does the homologous structure
92 NAUTILUS Vol. 73 (3)
in Laevapex or Ferrissia. The placement of shell muscles in
Ancylus fluviatilis is different from that of any of the American
forms yet examined, and in the more distantly related Acroloxus
lacustris the pattern is again different. When more genera
of the family Ancylidae become known anatomically, this may
show that the pattern of shell muscle distribution is significant
in tracing relationships.
Respiratory system: The pseudobranch (pi. 8, fig. D) des-
cribed above, resembles that of Ferrissia or Ancylus, and not the
far more complex structure found in Laevapex. The pulmonary
cavity is almost obsolete, its opening represented only as a nar-
row horizontal slit just dorsal to the anterior part of the pseudo-
branch. Undoubtedly, the habitat of this species, in rapids of a
swiftly-flowing stream, permits it to obtain sufficent oxygen
through the general body covering and renders a highly devel-
oped respiratory structure unnecessary.
Digestive system: The general plan of the digestive tract is
much the same as that of Ferrissia, Laevapex, or Ancylus. The
salivary glands, fused posteriorly, are longer than those presently
known from any other American form, but the most striking
feature of the digestive system is the extraordinarily long radular
sac. This structure (pi. 8, fig. C) extends well back into the
body, as in Ancylus fluviatilis, the common river limpet of
Europe. Whether the conformation of the radular sac is de-
pendent upon the diet of these animals, or whether it reflects
phylogenetic relationships, is at present unknown.
The peculiar radula was used as a major criterion for estab-
lishing the subfamily Rhodacmeinae by Walker (1917), and is
difficult to interpret in view of the strong similarities in other
features between Rhodacmea and other ancylid genera. The jaw
(pi. 7, fig. D) with its many small lateral elements, resembles
that of Ancylus far more closely than it does any of the known
American species (cf. Hubendick, 1947: fig. 15) .
Excretory system: The kidney of Rhodacmea shows the char-
acteristic pattern of flexures found in other ancylid snails (with
the exception of Acroloxus), and histologically presents the same
picture. The shape of the kidney appears to be a far more con-
servative character than radular or reproductive structures and
may be useful in the discrimination of phylogenetic groups
January, 1960 nautilus 93
within the family.
Nervous system: The brain (pi. 8, fig. A) is much the same as
that of Laevapex, and of Basommatophora in general. The size,
shape, and number of otoliths in the statocysts agree with those
of Laevapex. Eyes and osphradium show no peculiarities.
Circulatory system: The structure, position, and relations of
the heart (pi. 8, fig. B) are as in Laevapex, Ferrissia, and other
sinistral ancylids.
Reproductive system: In this system, Rhodacmea differs from
the other genera of ancylids whose anatomy is known. The
ovotestis has relatively few follicles (8 to 10), resembling that
of Ferrissia and Ancylus. The seminal vesicle is a variable struc-
ture, probably changing shape in all species according to the
progress of the breeding season. In the specimens of Rhodacmea
examined, the seminal vesicle consists of a number of papillose
side branches of the hermaphroditic duct (pi. 9, fig. E) appear-
ing identical in shape to that of Ancylus (Lacaze-Duthiers,
1899), and similar to Ferrissia parallela (Baker, 1928). In
Ferrissia tarda, the seminal vesicle is a simple sac (Hoff, 1940) ,
and in Laevapex fuscus it is a greatly expanded structure as
large as the ovotestis (Basch, 1959) .
The albumen gland in the living Rhodacmea consists of a
number of glistening, bright yellow, elongated pear-shaped folli-
cles which collapse quickly when placed in fixative. In shape,
this structure resembles the homologous gland in Ancylus.
The organ here called the uterus is actually a compound
gland composed of at least 3 clearly differentiated histological
regions, one or more of which may represent the separate
nidamental gland as found in other ancylids and many other
Basommatophora. In the family Planorbidae, there is a tendency
for this gland to become incorporated into the wall of the uterus
(Baker, 1945), and, in such forms as Laevapex fuscus, it is diffi-
cult to distinguish uterine from nidamental tissue. The dorsal
wall of the hollow uterus is greatly thickened in the head region,
where it extends forward over the penial complex and part of
the buccal mass. The cytoplasm in this area stains a smooth
light gray color with haematoxylin and eosin, and nuclei are
very scarce. This appears to be a storage area. The thickened
area tapers off gradually posteriad, and disappears at about the
94 NAUTILUS Vol. 73 (3)
level of the left anterior shell muscle. Here the lateral wall of
the uterus consists of an extremely active secretory epithelium
whose many cells are filled with cords of brilliant red-staining
droplets, present in enormous numbers. The third kind of
tissue, folded in a complex pattern, is found in the medial wall
of the uterus and continues posteriad. Here the cytoplasm shows
a deep blue velvety appearance, with few nuclei. This area is
that which connects proximally with the other organs of the
reproductive system.
The prostate is an irregularly shaped extension of the vas
deferens, and in some specimens seems almost to be connected
with the portion of the uterus just described. The characteristic
pattern of folded lobes which is found in Laevapex and to some
extent in Ferrissia parallel a is entirely absent in Rhodacmea
cahawhensis. The distal portion of the male system (the penial
complex) is unique among the known ancylids. The penis is
simple (pi. 9, fig. A) showing neither the ultra-penis arrange-
ment found in Laevapex nor the flagellar structure found in
various modifications in Ferrissia species and in Ancylus.
Relationships: Only a relatively small number of the dozens
of described species of ancylids have been dissected, so that
meaningful evaluations of the significance of anatomical char-
acters are extremely difficult. Still, something can be said about
the relationships of Rhodacmea. The general body morphology,
kidney, brain, pseudobranch, and digestive tract (except for the
radula) all show that Rhodacmea is indeed an ancylid snail and
related to the other genera mentioned in this paper. Several
features, such as the shape of the jaw, the radular sac, and the
albumen gland seem to indicate an affinity with the genus
Ancylus, although these resemblances may be superficial and
coincidental. The shell, radula, and penial structure, which all
are unique among ancylid snails, support the separation of
Rhodacmea as a genus, and these characters probably have suffic-
ient weight to uphold Walker's separation of the Rhodacmeinae
as a distinct subfamily.
Literature Cited
Baker, Frank Collins. 1928. The fresh water Mollusca of Wis-
consin. Part 1. Gastropoda. Wise. Geol. and Nat. Hist. Surv.
Bull. No. 70, 507 pp.
January, 1960 nautilus 95
1945. The molluscan familv Planorbidae. Urbana. Univ.
111. Press. 530 pp.
Basch, Paul F. 1959. The anatomy of Laevapex fuscus, a fresh-
water limpet (Gastiopoda: Pulmonata) . Misc. Publ. Mus.
Zool. Univ. Mich., 108: 1-56.
Hoff, C. Clayton. 1940. Anatomy of the ancylid snail, Ferrissia
tarda (Say) . Trans. Amer. Micros. Soc, 59 (2) : 224-42.
Hubendick, Bengt. 1947. Phylogenetic relations between the
higher limnic Basommatophora. Zool. Bidrag Fr. Uppsala,
23: 141-64.
Lacaze-Duthiers, H. de. 1899. Des organes de la reproduction de
VAncylus fluviatilis. Arch. Zool. Exp. et Gen., 3rd Series, 7:
33-120.
Walker, Bryant. 1917. Revision of the classification of the North
American patelliform Ancylidae. Naut. 31 (1) : 1-10.
Editor's note. For the benefit of those accustomed to the
Pilsbryan anatomic terms, which long have been established in
geophile Pulmonata, the "penis" in the preceding paper is a
verge or penial papilla; the "penial complex" corresponds to
an introverted penis; and the "flagellar structure" is a penial
appendix, on the vergic sheath or capsule. Dr. Pilsbry, since
1895, employed flagellum for a free appendix or caecum on the
epiphallus, which is not developed in the limnophiles. — H. B. B.
STUDIES ON MOLLUSK POPULATIONS: 4
Bv R. STOHLER
(Concluded from October no.)
As will be noted from Table 3, concerning specimens collected
at Duxbury Reef only, throughout the second half of the year
1950, there was a variation in the indices for this locality almost
as large as that in the entire area represented by Table 2. At
Duxbury Reef, O. biplicata varied from a slender 45.90 to an
obese 64.81. Oldroyd's varieties vary, similarly, from a slender
47.06 to an obese 61.74 (not differentiating between the several
"varieties," but all taken together) . Thus, the population at
Duxbury Reef well exceeds in its variations the varieties singled
out on what seem to have been purely subjective impressions by
T. S. Oldroyd. As may be stated, then, this study reinforces the
opinion of several authors as implicitly expressed (the Giffords,
1944, state: ". . . seems to indicate that Mr. Oldroyd dignified
mere individual variations by names, rather than . . .", and
Keen, 1937, omitted listing the varietal names in the check list) .
96
NAUTILUS
Vol. 73 (3)
o
CD
f— '
CO
Ox
O
S5 S ^
January, 1960 nautilus 97
The three varietal names given by Oldroyd have no taxonomic
significance and should be relegated to the synonymy of O. bipli-
cata (Sowerby) .
As indicated before, several other observations were made in
connection with this particular investigation which are con-
sidered worthy of record. In Table 2 are recorded measurements,
etc., made on what appears as two separate lots from La Jolla,
designated as I and II. Mr. Richard Shaw (see Stohler, 1952)
collected both lots at one time in one spot. The present writer
separated out from this single lot (and designated as II) all
shells that showed peculiar brownish markings on the early
whorls, reminiscent of O. pycna Berry. I thought that the mark-
ings might be an indication of hybridization, an idea which had
to be abandoned very quickly, because no O. pycna were con-
tained in that particular collection nor have any specimens of
this latter species been collected at that locality since then.
While lot II did not include as large a specimen as lot I, nor as
obese a one, nevertheless the arithmetical averages are identical
for all practical purposes. From observations made since that
time, seemingly about one fourth of all O. hiplicata shells show
such brown lines on the early whorls and that this is merely
another of the variables.
The smallest specimen in our collections obtained so far was
gathered by Dr. Cadet Hand (Department of Zoology, Univer-
sity of California, Berkeley) in the harbor at Monterey (see
Stohler, 1952) . As the averages of this lot indicate, there was a
gieater proportion of small shells picked up than elsewhere.
The sand in this harbor seems very fine and lacks coarse grains,
pebbles and rocks. In other areas to be discussed presently, the
sandy areas were not as uniform but included even fairly large
rocks and the fine sand was limited to relatively small pockets.
In attempts to section histologically fixed specimens, the stomach
was found to be full of sand in every case, a fact which seems to
indicate the olivellas ingest the sand in their feeding process.
This in turn would lead to the assumption that the size of the
sand grains might play an important role in the minimum size
of animals to be found in a given spot; large olivellas can ingest
coarse or fine sand while the small individuals would be limited
to fine sand only.
98 NAUTILUS Vol. 73 (3)
The following two observations appear to support this idea.
Columns Via and VIb of Table 3 refer to two areas at Duxbury
Reef; area A may be described as a very large, shallow tide pool
in a sandy area, interspersed with large and small rocks and
many pebbles of various sizes; area B was further seaward in an
area devoid of all rocks, having only fairly uniform fine sand
covering the area to a depth of about an inch. In area B, the
population was somewhat denser and also showed a smaller
average.
Mrs. Fay Wolfson of San Diego is carrying on at present a
growth study on Olivella hiplicata in the Flood Control Chan-
nel of that city. She discovered pockets of fine sand containing
only small shells and she refers in her "unofficial" records to
these areas as the "nurseries." Large specimens of this species are
found elsewhere. Mention of any other facts uncovered so far in
her study is not proper at this time, but this observation cer-
tainly appears to corroborate the assumption expressed above.
As is necessary to mention at this point, these several observa-
tions will have to be considered again a little further below in
connection with the breeding "season" of Olivella.
On the various collecting trips, I noted that, as soon as the
cold waves washed over the previously exposed sand, the buried
and apparently quiescent olivellas became active again; tracks
immediately appeared and I also observed that there was a
tendency for pairs to form, one individual following the other.
Usually, such pairs eventually mated. This mating behavior was
observed almost every time I was at Duxbury Reef, regardless
of the time of the year, except when nightfall prevented the
awaiting of the returning tides. Since this behavior was seen
every month of the year, a safe conclusion is that O. hiplicata
does not have a limited breeding season and that, consequently,
small or rather very young, as well as fully adult individuals
should be found throughout the year. This latter expectation
is borne out by Table 3. Thus from June 1 (indicated as 1 VI)
to December 21 (21 XII), 1950, specimens of over 25 mm in
length were found. The absence of the very small specimens can
be ascribed, I think, to the lack of the very fine sand necessary
for the young animals; I also suggest that the very young olivellas
cannot maintain themselves in the shallower water where the
January, 1960 nautilus 99
tidal changes cause a churning up of the sand and that they may
be expected further down, where the wave action has less effect —
or in protected bays and harbors. The latter seems to apply to
the lot collected in Monterey Harbor, while the former seems
to be the case with the "nurseries" in the Flood Control Channel.
The facts and observations just discussed also may apply in one
way or another to the lot from Point Abreojos in the Oldroyd
collection and designated as O. biplicata parva. On the other
hand, for the time being at least, the possibility of conscious or
unconscious selection on the part of Mr. Hemphill in collecting
the lot cannot be excluded.
On July 29, 1950, 11 pairs of Olivella were collected; these
were at the time actually mating; at first I thought that what had
been termed obese shells might owe this fact to the sex of the
animal, because logically the female with its ovary might require
a more spacious shell than the male. This was, however, not
borne out by the actual measurements. In the mating pairs
(M. P. in Table 3) , the animal plowing ahead is the female
while the one following is the male (when the animals are sep-
arated from each other, the penis is always part of the second,
i.e., posterior animal) . The shell of the females in all 1 1 cases
was longer than that of the male, the differences varying from
0.3 mm to 6.0 mm. As may be seen from the table in column
III b, among the mating pairs the smallest animal was 20.2 mm
long. Smaller mating partners have been observed (as stated
above, a total of 49 mating pairs have been collected, but only
the measurements etc. made on these initial 11 pairs are re-
corded here since the addition of the data from the other 38 pairs
would not have altered the picture and merely added bulk) , the
smallest found being a male of 17.9 mm length mating with a
female 27.0 mm long. Of the 11 pairs reported upon here, the
female was more obese than the male in 6 cases, the male was
more obese than the female in 5 cases. The greatest index differ-
ence between the two sexes was approximatly 9.4 units in the
group where the female was more obese (average of differences
appr. 4.75 units); in the second group the largest index-difference
was only 4.5 units and the average of the differences was about
2.36 units. One of the mating pairs collected on March 22, 1951,
seems interesting; in this pair, the male (20.7 mm long, 12.5 mm
100 NAUTILUS Vol. 73 (3)
wide) had an obesity index of 60.39 while the female (24.0 mm
long, 11.4 mm wide) had an obesity index of only 47.5, a differ-
ence of the two of 12.89; in other words, in this one case the male
is even more obese than the female than in the reverse situation;
but, of course, this is apparently an exceptional case and merely
tends to emphasize the overall conclusion: relative obesity of the
shell cannot be taken as a sex character in O. biplicata.
In Table 1 are recorded also the measurements and indices
obtained from a lot of albino shells, labeled Olivella biplicata
lapilhis (Vanatta) . While this varietal name has previously been
relegated to the synonymy of O. biplicata (Sowerby) based on
a study of color variations within unselected populations (Stoh-
ler, 1952) , these measurements were made, nevertheless, in order
to test the possibility that the white shells really were different in
other ways than in pigmentation. However, as this rather small
lot shows, the range of variation of this group falls well within
the range of our species; thus, as far as size of the shell is con-
cerned, the varietal name lapilhis also appears not justified. In
spite of these facts, in the studies reported on here, a note was
considered desirable whenever a white shell was measured; also
for the mating pairs this particular aspect was recorded. Without
going into a tedious enumeration, the statement may be sufficient
that in the tables of measurements and in the tables of the indices
computed, the letter W (denoting white shell) appears in a
random fashion. As should be stated, the two sets of tables for
each lot were arranged from the shortest to the longest shell and
from the lowest to the highest obesity index, respectively. The
following list is a sample of the distribution of W in the first
100 (smallest) shells of lot XIII from Duxbury Reef (this lot
included 1318 shells, of which 113 were white): Nos. 2, 9, 21,
45, 50, 56, 70, 93, were white, the other shells varied in color as
previously reported. In the table of the obesity indices of the
same lot XIII, the following were white shells among the 100
slenderest specimens: Nos. 5, 13, 52, 62, 84. As seems superfluous
to mention, these order numbers are, of course, not the same as
the ones in the previous list. As to the mating habits of the white
Olivella, I may point out that there seems no color preference
exhibited by the animals; for example in lot no. 9 from Duxbury,
there was a total of 115 individuals collected; of these 12 were
January, 1960 nautilus 101
mating; in one of the 6 mating pairs both partners were white,
in one other pair one partner was white (the male) while the
other partner was olive. The other four pairs showed random
color distribution. On the other hand, in lot 11 (with a total of
137 individuals and also 6 mating pairs) no white animals were
found in the process of mating. As perhaps is amusing to note,
in all the other 37 recorded mating pairs, no white individuals
were involved, which might lead to the erroneous conclusion
that white olivellas don't mate! However, workers in the field of
population genetics are well aware that such chance results are
to be expected in small samples.
The question as to the presence of a real breeding season in
Olivella biplicata at Duxbury Reef is not too readily settled.
From my field notes and from my records, the following data
shed some light on this problem: approximately 20% of all
individuals counted on July 29, 1950, were mating; approxi-
mately the same proportion was counted on November 23, 1950;
somewhat more than 10% were mating on December 21, 1950,
January 23, 1951, February 20, 1951, about 12% on March 22,
1951, but only about 2% on April 24, 1951. Thus, two high
peaks in mating activity were noted; but I cannot ascribe much
validity to these numbers for the simple reason that the instances
of little apparent activity occurred on the dates when either the
lateness of the hour or rough weather prevented me from ex-
tending my observations over a period of about 90 minutes past
the turning of the tide. I believe I may conclude safely that
O. biplicata actually reproduces throughout the year, although
possibly, at certain times, a greater percentage of the individuals
in a given population mate than at other times. Also, as should
be borne in mind, these conclusions are based on observations
made on inter tidal populations; what may take place in the sub-
tidal area is yet to be explored.
The mating behavior, and perhaps consequently the breeding
cycle, may be vastly influenced by meteorological phenomena.
The numbers recorded in Table 3 show amazingly small num-
bers of olivellas collected in August and October of 1950. My
field notes seem to me to contain the answer to this problem;
I quote them verbatim:
"26. VIII. 1950: Apparently the shore was lashed very recently
102 NAUTILUS Vol. 73 (3)
by heavy waves judging by amount of drift on beach and by
cover of fine silt over the sand. Only small number of O. b.
was seen."
"12. X. 1950: Exactly same appearance as on 26. VIII. 1950;
still fewer O. b. seen."
"9. XI. 1950: Water clear; no silt observed in entire area. O. b.
present in large numbers, ca. 25% mating."
That the population did not die out during the 6 or 7 weeks
of adverse conditions in the intertidal area is suggested by the
size distribution of the group collected on November 9; com-
paring the data in Table 3 shows that this group fits in very
well indeed with all the others. If, however, the population
previously present was killed off by the presence of silt, then
we would have to postulate that a very large population moved
in rather rapidly from the subtidal region. This in itself is not
impossible, and I assume that just such a situation did occur,
namely the olivellas present moved out when conditions became
adverse and returned when conditions improved again.
In conclusion, facts, observations and deductions may be sum-
marized as follows:
Olivella biplicata (Sowerby) is a highly variable species
throughout its range of distribution, varying from very obese
to very slender individuals in any unselected population. Also
it varies widely in color from almost entirely white to a very
dark chestnut brown, almost black.
There is no apparent sexual dimorphism in this species,
although in the measured mating pairs the female was, without
exception, larger than the male. Size alone, however, would not
be a satisfactory criterion to distinguish the two sexes as there
is great overlap.
At Duxbury Reef in Marin County, California, there is no
discernible breeding season, mating behavior having been ob-
served whenever tidal and meteorological conditions permitted
the observer to remain in the area for about 1 ^ hours after the
tide turned.
No sexual selection seems apparent as far as the shell color
of the individuals is concerned.
The size of sand grains in the habitat of a particular popula-
tion may influence the size range, i.e., coarse sand prevents very
small individuals from living there. In the search for food,
O. biplicata does ingest sand.
January, 1960 nautilus lOS
During low tide O. biplicata becomes quiescent, tending to
aggregate, sometimes in very large numbers, near larger rocks
or kelpholdfasts; this quiescence is observed even in fairly large
tide pools. With the return of the first cold waves, the animals
resume active locomotion and travel surprisingly rapidly and far.
This behavior is ascribed to a lack of oxygen rather than the
influence of the tidal pressure.
The varieties angelena and fucana, established by Oldroyd,
are considered of no taxonomic validity. The door is left open
for the variety parva, described from Point Abreojos in Lower
California, being possibly a valid geographical variant.
Bibliography
Gifford, D. S., and E. W. Gifford. 1941. Color variation in Oli-
vella biplicata. Naut. 55 (1) : 10-12.
1942. Color variation in Olivella biplicata in various locali-
ties. Naut. 56 (2): 43-48.
1944. California olivellas. Naut. 57 (3) : 73-80.
Keen, A. Myra. 1937. An abridged check list of west North Ameri-
can marine Mollusca. Stanford Univ. Press, pp. 1-84.
Oldroyd, T. S. 1918. Olix'ella biplicata angelena, var. nov. Naut.
32 (1): 34-35.
1921. Some varieties of western olivellas. Naut. 34 (4):
117-119, p. V, figs. 4, 5, 6, 7.
Oldroyd, I. Shepard. 1927. The marine shells of the west coast
of North America. Vol. II, pt. 1: pp. 160-162: pi. 26, figs.
16, 16a, 17, 17a, 20, 20a, 23, 23a.
Stohler, R. 1952. Studies on mollusk populations: II. Naut. 65
(4): 135-137.
NEW PHYLLONOTUS FROM THE EASTERN PACIFIC
Bv A. MYRA KEEN
Stanford University, California
Having in recent years recommended the deletion of Phyllono-
tus (family Muricidae) from the roster of West American
generic names, I now have the privilege of reinstating it, through
a new discovery by an alert collector, Dr. Donald R. Shasky. To
bring out the real significance of this find, one must review a
little the history of the name Phyllonotus and its place in the
Muricidae.
Phyllonotus Swainson, 1833, has, by convention, included sev-
eral handsome muricid species that share a tendency toward
fu^ / ^ '^
104 NAUTILUS Vol. 73 (3)
brown color banding, pink or brown coloration around the
aperture, and heavy varices that are not firmly soldered to the
whorl, especially near the suture. The number of these varices
per whorl and their complexity of form are classificatory char-
acters not usually regarded as important. A few years ago, with
the revival of Hexaplex Perry, 1811,^ complications began, for
its type species would come within the boundaries of Phyl-
lonotus in its customary sense. Nothing is gained, however, by
synonymizing Phyllonotus. A possible way to preserve it would
be to redefine both units; interestingly enough, a division could
be made on the basis of morphology that would separate the
species also into geographic groups. Hexaplex, with its East
Indian type species and 3 West American forms — H. brassica
(Lamarck, 1822), H. erythrostomus (Swainson, 1831), and H.
regius (Swainson, 1821)^ — would be restricted to the Pacific,
whereas Phyllonotus would be entirely Atlantic in distribution.
The type of Phyllonotus has usually been assumed to be the
northern Caribbean P. pomum (Gmelin, 1791), for the actual
monotypical type species, P. imperialis (Swainson, 1831) has
been considered identical with it. Abbott (1958, p. 61) has
challenged such a synonymy and, pointing out that the name
Murex imperialis is a primary homonym, has renamed the type
species as M. (Phyllonotus) margaritensis. His figures of the
southern Caribbean form looked so disquietingly like Hexaplex
regius that I speculated in print (Keen, 1959) on whether this
might indeed be a Caribbean Hexaplex, in which case Phyllono-
tus as a name would be jeopardized more than ever. Opportun-
ity since then to study several specimens and tabulation of all
available size measurements have laid at rest my fears. I am con-
1 Hexaplex was based on several species, none indicated as type. Jous-
seaume in the 1880's cited Murex cichoreus as type, but as this name was not
actually in the original list, the first valid type selection was that of Hertlein
and Strong, 1955: H. foliacea Perry zr Murex cichoreus Gmelin, 1791 [better
known under Lamarck's later name, M. endivia], an Indo-Pacific species.
2 This specific name has been attributed to Wood, 1828, but I can now
report that it was actually validated earlier by Swainson in his Exotic
ConcJiology, ed. 1, pt. 2, October, 1821, in the caption to an unnumbered
plate. The name was overlooked entirely by Sherborn in his lyulex Ani-
maliutn (one of his rare omissions) , although earlier Sherborn and
Reynell (Proc. Malac. Soc. London, 11(5): 279, 1915) had cited Murex
regius as validated by Swainson. This earliest use of the name was confirmed
when Dr. S. S. Berry acquired recently a copy of Swainson's exceedingly
rare book.
January, 1960 nautilus 105
vinced that Phyllonotus margaritensis is distinctive and not a
Hexaplex. New limits, however, will have to be set tor both
groups, thus:
Hexaplex includes those shells with several varices (usually
more than 4i/^) per whorl, with well-developed frills and spines
on the varices, especially along the shoulder of the whorl. Pink
coloration of the aperture is characteristic in all species, from a
trace of color at the posterior notch to complete suffusion of the
entire aperture.
Phyllonotus comprises those shells with fewer than 41/2 varices
(mostly 3 to 31/2) per whorl; varices have subdued spines or none
at all. Pink coloration in the aperture is sporadic (i.e., in only
one species and not invariably in that one) .
Under the new arrangement, Phyllonotus, with P. pomum, P.
margaritensis (or, as Dr. Clench [1959, p. 333] prefers, P.
pomum margaritensis), and one or two West African species,
would still be a group confined to the Atlantic were it not that
a new and unquestionable eastern Pacific Phyllonotus has been
found. Not only that, but the 8 specimens of the new form so
far seen show sufficient morphologic and geographic differenti-
ation to suggest a species and subspecies, which are here des-
cribed.
Phyllonotus Swainson, 1833
Zool. Illust., ser. 2, vol. 3, pi. 100 (generic name only, no
species names cited) .
Type species (subsequent fixation, Swainson, 1833) : Murex
(Phyllonotus) imperialis, var. a = M. imperialis Swainson, 1831
(not Fischer, 1807) ^=z M. (P.) margaritensis Abbott, 1958 (new
name) .
Phyllonotus peratus, new species. PI. 10, fig. 6
Shell of medium size, with about 8 whorls, general color a
dull grayish-brown, with obscure traces of underlying color
bands of cinnamon to chestnut brown below the suture and
on the base, with lighter areas between varices, and with darker
spots on intervarical nodes; aperture white, suffused along the
inner lip (also inside the outer lip of the holotype) with a
warm creamy-yellow, and with four dark brown spots (ends of
discontinuous color bands) showing on the outer lip, of which
the uppermost spot continues above the aperture as a brown
blotch. Sculpture of hollow spines along the varices, simulating
106 NAUTILUS Vol. 73 (3)
those of Hexaplex on a small scale, with 2 (rarely 3) axial rows
of sharply-defined intervarical nodes. Varices regularly 3 per
whorl, each varix ending just short of the corresponding varix
of the previous whorl, so that the rows of varices run diagonally
up the spire. Spiral sculpture of several primary cords (8 on the
last whorl of the holotype) that show best near the varices and
may disappear between the nodes, with 6 to 8 uniform-sized
spiral threads both on and between the primary cords. Aper-
ture somewhat elliptical, the inner lip nearly smooth, rising into
a free-standing parietal w^all; outer lip serrate, with 14 irregular
notches that reflect the primary and some of the secondary
spiral sculpture. Operculum dark brown, ovate, with the nucleus
sub-terminal and the right margin thickened within; outer sur-
face not smooth but showing a radial depression, marking the
inner edge of the thickening, and about 1 1 undulating con-
centric increments of growth. Posterior siphonal notch of mod-
erate size, angular. Anterior canal narrowed, markedly recurved,
nearly closed. Remnants of previous canals present (one on
holotype, two on paratype) , projecting to the left of the pillar.
Dimensions: Holotype, length, 66 mm., width, 41 mm., length
of aperture, 24 mm.; paratype, length, 69 mm., width, 42 mm.,
length of aperture, 25 mm.
Repositories: Holotype, California Academy of Sciences Paleo.
Type Collection, no. 7780; paratype, no. 7780A.
Type locality: Calif. Acad. Sci. loc. 17974, 14 mi. SE of Judas
Point, Costa Rica, depth 42 fathoms (T. Crocker-N.Y. Zool.
Soc. Exped. Sta. 214-D-l, 9° 19' 32'' N. Lat., 84° 29' 30" W.
Long., Mar. 1, 1938; mud and shell bottom) .^
Derivation of specific name: The Greeks had a word for this:
their adjective peratos (transliteration) , meaning, "on the oppo-
site side, west." W^hat more appropriate term could one ask?
Comparisons: Although some Phyllonotiis pomum specimens
are said to be spinose, none among the 40 available for com-
parison or among published illustrations equal P. peratus in
this respect. Perhaps other specimens than the type lot of P.
peratus lie, as they did in the California Academy's collection,
3 After this paper had been submitted for publication, I received a report
from Mr. Martin D. Burkenroad that he had just got some peculiar speci-
mens of a Hexaplex. Mv guess that they might be Phyllonotiis proved cor-
rect. One of his 3 specimens, submitted for study, is a good P. peratus
measuring 55 mm. in height, 36.5 mm. in diameter. The range of the
species is therefore extended to Panama (trawled in 40 fathoms off Parita
Bay, Gulf of Panama, Oct. 16, 1959). Relative proportions and sculpture
of this specimen cast doubt on the validity of P. p. deeoris as more than a
color form, though again the southern specimen is slightly more spinose
and darker in color than those from Mexico.
January, 1960 nautilus 107
mislabelled "Murex regius, juvenile." The 3 varices and lack of
pink coloration, of course, refute this.
Phyllonotus peratus decoris, new subspecies. PI. 10, figs. 4, 5, 7.
This may be only a color form, but the differences seem con-
sistent. The aperture is brighter yellow than that of P. peratus,
s. s., and the spire is a delicate coral pink for the first three or
four whorls in some specimens. The untreated shell is lighter
in color: a grayish-yellow. Etching with acid ( which had hap-
pened to several specimens before they reached their present
owners) reveals an underlying color pattern of bright cinnamon-
brown bands on a white ground, with small flecks of darker and
lighter brown to the left of each node between varices. Sculp-
ture is lower and more subdued. The shell seems also a little
smaller, with a shorter anterior canal and, in most specimens,
a slightly shorter spire.
Dimensions in mm.
Length Width Length of aperture
Repositories: Holotype, Stanford Univ. Paleo. Type Coll., no.
8753, courtesy of Dr. D. R. Shasky; Paratype, collection of Dr.
SJiasky; Hypotypes 1 to 3, collection of Captain Xavier Men-
dosa, Guaymas, Mexico; Hypotype 4, pi. 10, fig. 7, collection of
Dr. Edwin J. Purcell, Tucson, Arizona, from west Mexico, exact
locality and depth unknown.
Type locality: West Mexican coast near the Guatemalan
borcler, depth about 15 fathoms; obtained from a shrimp boat
by Dr. Donald R. Shasky.
Other localities: The hypotypes are also from shrimp boats —
exact localities unknown but unquestionably from the West
Mexican coast between the Gulf of Tehuantepec and Guaymas,
Sonora, probably in the Salina Cruz area, all taken in nets of
the Hector II, Cpt. Juan Rios, skipper.
Derivation of name: The adjective decoris is a Latin w^ord
meaning "elegant", "ornamented", or "beautiful", all appro-
priate terms for this shell.
Comparisons: The distinctions between P. peratus peratus and
P. peratus decoris, though slight, are apparently consistent, the
108 NAUTILUS Vol. 73 (3)
dimensions of the 6 specimens of the latter being almost identi-
cal except for the one slender variant shown in fig. 7. P. peratus,
s. s., geographically a more southern form, seems to be somewhat
the larger, with a higher spire and longer canal. Differences of
both from P. pomum may be summarized thus: In P. pomum
the varices become narrowed at the suture, giving the effect of
being tucked under the preceding varix, whereas in P. peratus,
s. I., they tend to flare and in P. peratus, s. s., to form several
imbricating scales or laminae. In P. pomum the spiral sculpture
consists of alternating primary and secondary ribs, striated by
finer threads. In P. peratus the secondary sculpture is nearly
even, with no additional fine threads. In P. pomum the intervar-
ical spaces rise into one or two rounded axial ribs, but in P.
peratus there are only isolated and sharp-pointed nodes in irreg-
ular rows. Graphical analysis of height-width ratios shows that
although P. pomum is more variable, P. peratus tends to be
longer for the width, to have a larger aperture, and to have a
narrower canal.
Phyllonotus pomum seems to be a stock in which geographic
races are developing into possible subspecies. The southern Car-
ibbean stock, here regarded, for convenience, as a separate
species (P. margaritensis), seems, on the basis of 9 specimens for
which measurements were available (as compared to 18, chosen
as random, for P. pomum), to grow much larger (maximum
length 110 mm., as compared to 85 in P. pomum), and it is
markedly wider for the length, with a larger aperture and a
wider canal. One unnamed variant of P. pomum, represented in
the Stanford Collection by 3 specimens from Colon, Panama, is
nearly as spinous as P. peratus, but the shells are decidedly
thinner and more slender, darker in color (almost a uniform
chestnut brown in one specimen) , with a much longer and nearly
straight anterior canal. All these variations imply that the
Miocene ancestor of the tropical-American Phyllonotus stock
was a vigorous strain capable of a number of geographic varia-
tions that are now being realized in both the western Atlantic
and the eastern Pacific.
Phyllonotus sp. PI. 10, figs. 2-3
A single specimen of a form that one would assume should be
PLATE 10
1 (x 1) ' Phyllonotus pomum (Gmelin), Grand Bahama I., James McLean,
coll. 2 (X 9) and 3 (x 1)' P- sp- 4, 5 (x 1). P- peratus decoris Keen,
holotype. 6 (x 1) > P- peratus Keen, holotype. 7 (x ^) , P- peratus decoris
Keen, hypotype 4.
NAUTILUS 73 (3)
PLATE 11
1-3, Calipyrgula pecosensis Leonard and Ho (1, holotype, 11265, KUMNH.
2 paratype,' 11266 KUMNH. having 7 whorls and proportions approaching
C. hibbardi. 3, paratvpe, 11266, KUMNH., sectioned to show strongly twisted
columella). 4, 5, C. hibbardi Leonard and Franzen, paratypes, 981,
KUMNH., from Lower Pliocene Laverne Formation, bx/o miles south Gate,
Beaver County, Okla. (4, sectioned to show nature of columella). All
figures enlarged approximately 15 times.
January, 1960 nautilus 109
part of the P. peratiis complex was collected by Mr. E. C. Stiles
on the beach at Mazatlan, Mexico .The specimen, although not
taken alive, was but recently dead, for the nuclear tip is com-
plete and perfect, showing 21/2 dark brown, rather cylindrical,
smooth whorls above the cancellate juvenile shell. Having only
6 whorls, the specimen is evidently immature. Careful study of
sculpture, form of varices, and other criteria that separate P. per-
atus from its congeners leaves one with the conviction that this
is a specimen of P. pomum. It is figured here for two reasons:
to show the form of the nuclear whorls, of which I have seen
no good illustration, and to put collectors at Mazatlan on the
alert. Though this seems unlikely, one must not dismiss as im-
possible the survival of P. pomum in both oceans since the sepa-
ration of the Atlantic and the Pacific during the later part of
the Tertiary. Fortuitous introduction of this specimen to the
Mazatlan beach seems more probable, however, perhaps by shell-
collecting tourists who had recently been working the shores of
eastern Mexico.
Repository: Stanford Univ. Paleo. Type Coll., no. 8754, cour-
tesy of Mr. E. C. Stiles, collector. Mazatlan, Mexico, spring, 1958.
Acknowledgments. My special thanks go to the collectors
whose loan and gift of material made this paper possible: Dr.
Donald R. Shasky, of Glendale, California; Dr. Edwin J. Purcell,
of Tucson, Arizona; Mr. Everett C. Stiles, of Seattle, Washington,
and the California Academy of Sciences, San Francisco. I wish
to thank Dr. William Clench for a comparative specimen of
P. margaritensis, Mr. and Mrs. John Q. Burch for the privilege
of studying their holdings of this form, and Mr. James McLean
for the loan of an unusually fine specimen of P. pomum for
illustration.
References
Abbott, R. Tucker. The marine mollusks of Grand Cayman
Island, British West Indies. Monog. Acad. Nat. Sci. Phila-
delphia, no. 11, 138 pp., 5 pis., Dec. 31, 1958.
Clench, W. J. The genus Murex in the Western Atlantic, supple-
ment. Johnsonia, vol. 3, no. 39, pp. 331-334, 2 figs., June 26,
1959.
Keen, A. Myra. Some side notes on "Sea Shells of Tropical West
America." The Veliger, vol. 2, no. 1, pp. 1-3, July 1, 1959.
110 NAUTILUS Vol. 73 (3)
A NEW SPECIES OF CALIPYRGULA (HYDROBIIDAE)
FROM THE PLEISTOCENE OF TEXAS
By a. BYRON LEONARD and TONG-YUN HO
In the course of studies on the fossil molluscan faunas in the
late Cenozoic continental deposits of western and southwestern
Texas, sponsored jointly by the University of Texas Bureau of
Economic Geology, and The National Science Foundation under
the terms of a contract (NSF'-G3481) between National Science
Foundation and the University of Kansas, shells of an unde-
scribed species of hydrobiid gastropod in late Pleistocene depos-
its in the Pecos River Valley were discovered.
The new species is assigned to the genus Calipyrgiila Pilsbry,
because of the general resemblance to other species now assigned
to that genus, and the great difficulty of establishing actual rela-
tionships among hydrobiid snails in the absence of the soft
parts. There is no proof that the species described by Leonard
and Franzen (1944, p. 19) from the early Pliocene Laverne
Formation are in fact congeneric with those described by Pilsbry
(1934, p. 541) or that the species under consideration here is
congeneric with either of the two previously described groups
of species assigned to the genus Calipyrgula. Admitting the
widely separated localities of occurrence and considerations of
stratigraphic occurrence (Leonard and Franzen, 1944; Pilsbry,
1955) , it seems unlikely that all these species are actually con-
generic, but there seems little to be gained by proposing generic
names for the species concerned.
Calipyrgula pecosensis, new species. Plate 11, figs. 1-3.
Diagnosis: A minute hydrobiid gastropod, having an elongate,
narrowly conic, imperforate or narrowly rimate shell of 7 or 8
rounded whorls; small aperture; simple, ovate peristome, re-
flected over umbilicus and adherent to last whorl above; simple
and well incised suture.
Calipyrgula pecosensis (pi. 11, figs. 1-3) most closely resembles
C. hihbardi Leonard and Franzen, (figs. 4, 5) but differs in hav-
ing a more slender shell, one or two more whorls, less elongately
oval aperture (figs. 1, 2, 5) and more strongly twisted and
slightly heavier columellar axis (figs. 3, 4) .
Holotype: Catalogue number 11265, University of Kansas
Museum of Natural History, obtained by A. B. Leonard and
January, 1960 nautilus 111
Tong-Yun Ho, 6 June 1959. Orginal number, ABL 1012A.
Description of holotype: Shell elongately conic, imperforate,
small (total length, 4.42 mm., diameter, 1.27 mm.), last whorl
relatively large, comprising approximately one-half the total
length of shell; spire narrow, slender, tapering gradually to
relatively blunt apex; aperture ovate, higher than wide (height,
0.97 mm., diameter 0.71 mm.) ; peristome simple, inner margin
reflected over umbilicus, and closely adherent to last whorl
above; whorls 8, well rounded at periphery; suture simple,
deeply incised; one and one-half apical whorls finely granu-
lose having waxy texture, remaining whorls having closely
spaced, delicate, vertical growth ridges not clearly visible with-
out magnification, and extremely delicate incised spiral grooves,
visible only with high magnification.
Paratypes: Little variation occurs among the more than 1000
paratypes collected from 6 localities, although a few individuals
(PL 11, fig. 2), are less narrowly elongate than is the holotype
and have proportions that more nearly approach those of
Calipyrgula liibbardi. The extremes of variation are exemplified
by the measurements (in millimeters) of 4 paratypes (catalogue
number, 11266) from the type locality.
Total Height Diameter No. of
length Diameter Aperture Aperture whorls
On a few paratypical shells, the fine, incised spiral sculpture
is somewhat better developed, especially on the last whorl, than
on the holotype, but on none is the sculpture a conspicuous fea-
ture. Rarely, individual shells possess irregularly distributed,
strong, vertical ridges, seemingly produced by fusion of the usual,
fine, more numerous vertical ridges. Still other rare specimens
have the peristome erect, not reflected over the minute umbilicus,
and standing free of the last whorl above. Paratypical specimens
have been deposited at the Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts.
Type locality: Late Pleistocene deposits exposed in left bank
of Pecos River, 3.5 miles northeast of Imperial, in Crane County,
Texas.
Stratigraphic distribution: Each of the six collecting localities
from which Calipyrgula pecosensis has been recovered consists of
late Pleistocene terrace deposits, exposed by subsequent degra-
112 NAUTILUS Vol. 73 (3)
dation of the channel of the Pecos River. If C. pecosensis is bio-
logically congeneric with C. hibbardi, of the early Pliocene La-
verne Formation of northwestern Oklahoma (Leonard and Fran-
zen, 1944, p. 19), the genus is unaccounted for in the region of
its known occurrence east of the Rocky Mountains, through
most of late Cenozoic time.
Areal distribution and ecology: Calipyrgula pecosensis is now
known only from exposures along the Pecos River, southwestern
Texas, distributed from Pecos River at bridge on U. S. Highway
290, 3 miles southeast of Sheffield, Pecos County, to one mile
east of the city of Pecos, in Ward County. Although the latter
locality provided the most abundant population of all the local
deposits from which the species was collected, we were unable
to discover examples farther upstream.
The 6 localities from which C. pecosensis was collected, to-
gether with approximate numbers of specimens taken at each
locality, are:
Right bank of Pecos River, near bridge on U. S.. Highway 290,
3 miles southeast of Sheffield, Pecos County, Texas; 5 specimens.
Right bank of Pecos River, 5 miles northeast of Buena Vista,
Pecos County, Texas; 50 specimens.
Left bank of Pecos River, 3.5 miles northeast of Imperial, in
Crane County, Texas; 400 specimens. (Type locality.)
Left bank of Pecos River, 3 miles northwest of Imperial, in
Crane County, Texas; 300 specimens.
Left bank of Pecos River, 3 miles southwest of Grand Falls,
Ward County, Texas; 250 specimens.
Left bank of Pecos River, one mile east of Pecos, in Ward
County, Texas; 350 specimens.
Little is known of the ecological requirements of Calipyrgula
pecosensis. Judging from the lithology of the sediments in which
it has been found (finely and well sorted silts and fine sand,
having humic stains and marl development) , this snail thrived
in sluggish streams or in marshes; this view is strengthened by
the occurrence with it of numerous individuals of Physa anatina,
Ferrissia parallela, Pisidium sp., and a species of ostracod. So
far as known, however, no other prosobranchiate gastropod
occurs with C. pecosensis.
Literature cited
Leonard, A. Byron, and Franzen, Dorothea S. 1944. Mollusca
of the Laverne Formation (Lower Pliocene) of Beaver
January, 1960 nautilus 118
County, Oklahoma Univ. Kansas Sci. Bull. 30 (pt. 1, no.
2) : 15-39, pis. 4, 5. May 15.
Pilsbry, Henry A. 1935. Mollusks of the fresh-water Pliocene
beds of the Kettleman Hills and neighboring oil fields,
California. Proc. Acad. Nat. Sci. Philadelphia 86: 541-
570, Jan. 29 (issue of 1934)
SUCCINEA AUREA LEA AND S. PYRITES, NEW
Bv LESLIE HUBRICHT
Succinea aurea was described by Lea (Proc. Amer. Philos.
Soc. 2:32, 1841. Trans. Amer. Philos. Soc. 9:4, 1846) from speci-
mens collected at Springfield, Ohio. During the spring of 1958,
the author visited Springfield to collect this species. Two species
of Succineidae, Oxyloma retusa (Lea) and Quickella vermeta
(Say) , were found in abundance in that area.
In his description. Lea states: "This is a beautiful little species,
remarkable for its fine surface and bright golden colour. In
form it is nearly allied to vermeta. (Say.)" Quickella vermeta is
sometimes of a bright golden color, especially when it lives in
sunny situations. Succinea aurea is undoubtedly based on Quick-
ella vermeta, and the Atlantic Coastal species which has borne
this name is a different species.
For this Atlantic Coastal species, the author proposes the
name Succinea pyrites, new species. It is described by H. A.
Pilsbry, Land Mollusca of North America (North of Mexico)
2:815-818, figs. 441a, b; 442E (holotype) , F. Type locality: Cape
May, New Jersey, holotype 67795, paratypes 247364 Sc 189420
A.N.S.P., other paratypes 20614, collection of the author.
Of the localities cited by Pilsbry for Succinea aurea, those from
near the Atlantic Coast from Virginia to New Jersey are prob-
ably all Succinea pyrites. In the authors experience, S. pyrites
is confined to salt marshes and the records from western New
York need to be verified anatomically before being accepted.
The records from Yammasee, South Carolina; Wells, York Co.,
Maine; and Barnstable Co., Masachusetts, are based on imma-
ture shells which are unidentifiable. The record from Oldtown,
Ohio, is based on an immature specimen of Oxyloma retusa
(Lea) . The records from Cincinnati, Ohio, and Clark Co.,
Indiana, are undoubtedly based on Quickella vermeta which is
a very common snail along the Ohio River.
114 NAUTILUS Vol. 73 (3)
THE GREEN LAND MOLLUSK FROM NEW GUINEA
B^ WILLIAM J. CLENCH
Mr. J. B. Henrard (Naut. 13: 63-65, 1959) has questioned the
generic placement of the species I described as Eustomopsis
(Verdichloritis) polingi. He claims that it should be Planispira
(Cristigibba) polingi. As he has not seen my specimen of polingi,
his specimens may be something else, as polingi is not, in any
sense, a Cristigibba and certainly is a ''Chloritis". The few speci-
mens of three species of Cristigibba in the MCZ. are compara-
tively heavy shells, are opaque, and possess a very strong gibbous
crest. E. polingi is thin, translucent and has no trace of a crest.
It does have a very slight depression behind the lip, a character
found in a multitude of land mollusks.
The shell morphology is that of Eustomopsis, but its final
placement in this genus must await the time when its soft anat-
omy is known.
Regarding the green coloration exhibited by a few genera of
land mollusks, I did not raise the question as to whether this
color was impregnated in the lime of the shell or only invested
in the periostracum. Henrard's statement that "in all those
shells, (i.e. Ligiius, Amphidromus) the coloring matter is situ-
ated in the calcareous layer of the whorls and all the other shells
mentioned by him", is certainly in error. The green color in
Liguus, most Amphidromus, Helicostyla, Papuina and others, is
located in the periostracum.
FAMILY NAMES IN PULMONATA, 4
By H. BURRINGTON BAKER
The following alphabetic index to the subfamilies and syno-
nyms of the family names in Pulmonata gives each combining
stem and, after a hyphen, enough letters to place the family in
the alphabetic list in my first (1) paper, Naut. ^9:128, which
was supplemented by a second (2) , Naut. 76>: 34, and a third (3) ,
Naut. 7^:141. This is the 4th. In these supplements, the names
in the first were corrected or redated (Cf. 2, 3, or 4) and more
names were added (2, 3 or 4) . The families with synonyms are
followed by CAPITAL letters, to assist in recognition of their
abbreviations.
January, 1960 nautilus 115
At the London Congress (1958) : "The rule that a family
name is not to be changed when the type genus is found to be
a synonym" (Cf. 1, footnotes 2 8c 4) "was reaffirmed, with the
clear provision that this was not intended to upset changes of
long standing or wide acceptance" (News letter, Soc. Syst. ZooL,
ser. 2, no. 9, p. 2). Will not the application of this "clear" pro-
vision and/or the "50 year rule" be very difficult? In any case,
please do not quote my 4 lists for the "acceptance" of any of
the changes involved. The family names which are based on
synonyms are Aperidae, Auriculidae (?) , Buliminidae, Coch-
lostylidae (obsolete) or Eulotidae, Gadiniidae, Helicterinae
(Achatinellidae) and Lucerninae (Camaenidae) . The last two
apparently have been in synonymy for 50 years, as is also true of
any name founded on Brachypodella (Urocoptidae), Mesodon-
tinae (Polygyridae) or Scolodens (Scolodontidae or Systrophi-
idae) , and probaby Plectopylididae (Corillidae) . Nevertheless,
"Helicteridae" was used as a heading in the Zoological Record
until 1939.
Acanthinul-Va.i Acav-ACA (cf. 3). Acell-Ly. Achatin-ACH.
Achatinell-Helict. Acrolox-An.^ Acrob-ini Starabogatov (1958)-
Pl. AcTore- (cf. 3) . Adven-Eu. Aegist-Eul (cf. 2). Agriolimac-Li.
Ailly-. Albe-Hv (3) . Alopi-Cl. Amastr-AMA. Ameriann-eae
Zilch (1959) -PI. Ammonitell-Thys., Lu. Amphibol-AMP.^
Amphibulim-Or. Amphidox-Pun. Amphidrom-Lu, Amphipepl-
Ly. Ampull-Ach (3) . Ampullacer-Amp. Anaden-Ar. Ancyl-AN
(cf. 3).^ Ancylastr-An.i Aneite-At. Anoglypt-Aca (3) Anthrac-
opup-inae Zilch (1959) -Au. Aper-AP. Aqueban-Sa. Ariant-He.^
Ariolimac-Ar. Arion-AR.^ Ariophant-Xe, Artemon-Str. (3).
Aspasit-Va. Athoracophor-AT. Aulacospir-inae Zilch (1959)-
Pup. Auricul-AU (cf. 2) . Auriculell-To. Austroselenit-Str.
Azec-idae^ Kennard & Woodward (1926) -Ci.
Bale-Cl.^ Berendti-Br. Binney-Ar. Biomphalari-Pl. Bothiembry-
Or (3) . Brachypodell-BR (= Urocoptidae) . Bradybaen-inae
Pilsbry (1924)-Eul. Bulim-.^ Bulimin-BU. Buliminops-Eul.
Bulimul-Or. Bulin- (cf. 3, but correct "Bulinulinae" to Bulin-
inae) Bullin-Pl (cf. 3).
Caecili-Ach. Caecilianell-Ach. Camaen-Lu. Camptocerat-Pl. Cam-
pylae-He. Carychi-CA.i Caryod-Aca. Cassidul-Au (cf. 3) . Cati-
nell-Su. Cecilioid-Ach.^ Cepol-, Cepoli-Xa (cf. 3) . Cerast-Bu.
Cerastu-inae Wenz (1930) -Bu. Ceri-, Cerion-CE (cf. 3) .^
1 "Type genus'' accepted by ICZN. Ancylus takes as type A. fluviatilis, so
Pseudancylinae is a synonym of Ancylinae, and Acroloxinae replaces my
(1) typical subfamily.
116 NAUTILUS Vol. 73 (3)
Charop-Pun. Chilin-. Chlamydephor-Ap. Chlorit-Lu. Choa-
nomphal-PI. Chondrin-Bu, Pup. Chondrul-Bu. Chron-Eu.
Cionell-CI. Circinari-Va, Ha. Clausili-CL.^ Clavator-Aca. Coch-
licop-Ol, Ci. Cochlodin-Cl. Cochlostyl-Eul (3), but prior (ob-
solete?) . Coeliax-Ach (cf. 3) Coneuplect-Eu (3) . Conovul-Au.
Conul-Eu.2 Coret-Pl. Corill-PIe. Cryptelasm-Ach. Cryptell-Li.
Cylindrell-Br. Cystopelt-Eu.
Daudebardi-Vi. Derocerat-Li (3) . Dipnelic-Lu (2) . Disc-Pun. i
Dorcasi- (cf. 3) . Drepanotrem (at)-eae Zilch (1959) -PI. Durgell-
Xe. Durgellin-Eu (cf. 3) . Dyak-Xe.
Elasmatin-To. Ellobi-Au. En-Bu (cf. 2) .^ Endodont-Pun. Enne-
Str (cf. 3) . Epiphragmophor-Xa. Erept-Helica (cf. 2) . Euca-
lod-Br. Euconul-EU.i Euglandin-Sp. Eulot-EUL (cf. 3). Eu-
paryph-He.-
Fanul-Eu. Ferrissi-An. Ferussac-Ach. Filholi-Cl. Flammulin-Pun
(cf. 3). Fruticicol-Eul, Hy. Fusul-Cl.
Gadini-G (cf. 3). Garnieri-Cl (cf. 3) . Gastrocopt-Pup. Gastro-
dont-Vi. Geomitr-Hy. Geotroch-Helica. Gibb-Str. (2) . Girasi-
Xe (cf. 3) . Glandin-Ol. Godwini-Vi. Gonidom-Str (2) . Gonio-
disc-Pun. Grangerell-Or. Gudeconch-Pun.
Hadr-Lu. Haplotremat-HA. Hedleyell-Aca. Hedleyoconch-Eu.
Helic-HE.i Helicarion-HELICA (ef. 3; Helixarionidae prior) .
Helicell-Vir Hy (cf. 3),^ He.^ Helicigon-He (cf. 3) . Helicodisc-
Pun. Helicodont-He (cf. 3). Helicostyl-Eul (cf. 3). Helicter-
HELICT (= Achatinellidae) . Helisomat-Pl. Helixarion-
Helica (3) . Helminthoglypt-Xa. Hemiplect-Xe. Holospir-Br.
Humboldtian-Xa. Hyalimac-Su. Hyalini-Vi. Hygromi-HY.^
Hypselostomat-inae Zilch (1959) -Pup.
Imer-Va. Isidor-Pl.
Jacost-Hy.- Jamini-Bu. Janell-At. Janul-Vi.
Kaliell-Helica.
Laevapic-An. Laminifer-Cl. Lampadi-Lu (3) . Lane- Laom-Pun.
Lati- (cf. 3). Lauri-Pup.^ Ledoulxi-Ur. Leptachatin-Ama.
Leptax-He. Leucochro-Hy. Liardeti-Eu. Ligu-Or (2) . Limac-
LI (cf. 3) .^ Limn-, Limnae-Ly.- Limnophys-Ly. Lucern-LU
(= Camaenidae) . Lymn-,- Lymnae-LY (cf. 2) .^ Lysino-Xa.
Macrochlamyd-Xe. Macrocyl-. Marpess-Cl. Megalophaedus-Cl
(3) . Megaspir-. Megomphic-Thys., Lu. Meisenheimeri-Va.
Melamp-Au. Mantissoide-Cl. Mesodont-MES (z= Polygyridae) .
Microceram-Br. Microcyst-Eu. Milac-Li.^ Miratest-Pl (3).
Murell-He (cf. 3) . Myotest-Xe.
Nanin-Xe. Napae-Bu. Neni-, Neniastr-Cl. Neoplanorb-An. Neso-
pup-Pup. Nitor-Helica.
Obelisc-Ach (cf. 3) . Odontostom-Or. Oleacin-OL.^ Onchid-,
Oncidi-ON. Oncidiell-On. Oopelt-Ar. Opeat-Ach. Orcul-Pup.
2 "Type genus" rejected by ICZN.
January, 1960 nautilus 117
Oreohelic-Lu. Orthalic-OR. Orthogibb-Str (2) . Ostracoleth-
Xe. Otin-.i Otoconch-Pun.
Pachnod-Bu. Pacificell-To. Pagodin-, Pagodul-, Pagodulin-Pup.
Papuin-Lu. Paralaom-Pun. Parmacell-Li. Parmacellill-Tr.
Parmarion-Xe. Partul-. Paryphant-Rh (c£. 3 & 2) . Patul-Pun.
Patulastr-Pun. Pedinogyi-Aca. Pediped-Au. Peltat-Ur. Peroni-
On. Phaedus-Cl. Phenacohelic-Pun. Philomyc-PHI. Philonesi-
Eu. Phys-. Physastr-inae Starabogatov (1958) -PI. Planispir-Lu.
Planorb-PL (cf. 3) .^ Planorbul-Pl. Platysuccine-Sa. Plectopy-
lid-PLE (cf. 3; = Corillidae). Plesiophys-inae Bequaert &
Clench (1939) -PI. Pleurodisc-PLEU (= Pyramidulidae/
1914) . Pleurodont-Lu. Plutoni-Vi. Poecilozonit-Vi. Polygyr-
Mes. Polygyrell-Thys. Pompholic-Pl. Pompholycode-Pl. Pro-
tancyl-Pl. Pseudancyl-An.- Pseudocharop-Pun. Pseudoplect-
Xe. Pseudotrochatell-Helica. Pseudoveronicell-Va. Ptychotre-
mat-Str. Punct-PUN.^ Pup-Ce (cf. 3) . Pupill-PUP.i Pyramidul-
idae^ Kennard & Woodward (1914) -Pleu. Pythi-Au.
Rathousi- (cf. 3) Rhagad-Lu. Rhodacm-An. Rhytid-RH (cf. 3) .
Rotadisc-Pun. Rumin-Ach.^
Sagd-SA. Sarasinul-Ve. Scalax-inae Zilch (1959) -Ach. Scaph-On.
Scarab-Au. Scolodent-, Scolodont-SC. Segmentin-Pl.^ Selenit-
Str. Semperul-Ve. Serrul-Cl (3). Sesar-Helica. Sheldoni-Ur.
Siphonari-. Sital-Xe. Sonorell-Xa. Sophin-Eu. Spelaeoconch-Ci.
Spelaeodisc-Va. Sphincterochil-inae Wenz (1959) -Hy. Spirax-
SP. Staffordi-Xe. Stenacm- (cf. Naut. 72:68). Stenogyr-Ach.
Stenop-Sc. Stenopyl-Pun. 8treptax-STR. Strepostel-Str (3) .
Streptostyl-Sp. Strobil-, Strobilops- STRO. Strobil-idae Zilch
(1959) not Joos (1911) -To. Succine-SU^ Systrophi-Sc.
Tacheocampyiae-He (3) . Tanychlamyd-Xe. Tebennophor-Phi.
Testacell- Theb-He.i Thyrophorell- (cf. 3) . Thysanophor-
THYS. Thysanot-Pun. Tornatellin-TO.i Trigonochlamyd-TR.
Triodops-Mes. Trimuscul-idae Habe (1958) -G. Triptychi-Cl.
Trochomorph-Vi. Trochananin-Ur. Eu. Trochozonit-Ur.
Troch-Hy. Truncatellin-Pup.^
Urocopt-Br. Urocyl-UR.
Vaginul-Ve. Vallon-VA.^ Varicell-Ol. Veronicell-VE. Vertigin-
Pup.i Vitre-Vi.^ Vitrin-VI. Vitriplutoni-, Vitriniplutoni-
Vi.
Xanthomelon-Lu. Xanthonych-XA. Xerophil-He, Hy.^ Xest-
XE.
Zacole-inae Webb (1959)-Ar. Zaptych-Cl (3; Xaptych-) . Zonit-
Vi. Zoph-Str. Zospe-idae Bourguignat (1856) -Ca.
3 Previously (1) , I did not notice that succinea was a Latin adjective.
NOTES AND NEWS
Lloyd E. Berry, 1892-1959.— On October the sixth, 1959, we
118 NAUTILUS Vol. 73 (3)
lost a very dear and true friend with the passing of Lloyd E.
Berry, a retired U. S. Army lieutenant who saw action in Siberia
during World War I. His passing at the U. S. Government Hos-
pital at Sawtelle was the result of service-connected disability.
We who have known Mr. Berry the last decade knew him as a
conchologist, who specialized on the Cypraeidae, having an out-
standing collection. Few Cypraea collectors could equal his col-
lection in quality, number of species, and number of specimens.
Nor do I know of a private collection that was so well kept. This
fabulous collection was inherited by his surviving sister, Mrs. J.
V. Yankie, 1804 Warwick Road, San Marino, California, and
will eventually be for sale as a unit. — Mrs. John Q. Burch
Sanibel marines. — I understand that some Indo-Pacific shells
are being found on Sanibel Island, Florida, and think that this
is the result of dredging. Some 3 shell collections were lost dur-
ing a hurricane in 1920, and now these shells are being found
on the beach, dead, of course. — Lula B. Siekman (from letter) .
Sinistral shell of Campeloma integrum. — While sinistral
specimens of the uterine young of Campeloma are fairly com-
mon, adult specimens of reversed symmetry are only rarely col-
lected. An adult sinistral specimen of Campeloma integrum Say
was collected by one of my students in field zoology, Mario
Pisanelli, on April 8, 1959. This shell and three dextral shells
of the same species were collected in a swamp at Twin Lakes,
north of Kent, Ohio. The sinistral shell measures 30 mm. high,
and 19 mm. wide. The apex is eroded, and 41/2 whorls are in
view. The species was determined by Dr. William J. Clench,
and the specimen was given to the Museum of Comparative
Zoology at Cambridge, Mass.
Over (Naut. 29: 90-95. 1915) reported a similar sinistral
specimen of this species from the Vermilion River in Clay
County, South Dakota. Sinistral specimens of C. decisum have
been reported by Ancey from New York (Naut. 10: 104-105. 1897),
Hinkley (Naut. 20: 40-44. 1906) from the Tennessee River,
Vanatta (Naut. 26: 96. 1912) from Pennsylvania, and Daniels
(Naut. 26: 38-42. 1912) from Indiana. F. C. Baker (The Fresh-
water Mollusca of Wisconsin, Part I, p. 67. 1928) reported one
January, 1960 nautilus
111
reversed specimen among 160 adults collected. Sinistral shells of
C. siibsoUdum were found in Missouri by Sampson (Naut. 29:
128-129. 1916) and one of C. geniculum by Goodrich (Naut. 52:
129-131. 1939) in Georgia. Webb (United States Mollusca, p.
149. 1942) gives an illustration of a sinistral specimen of C. pon-
derosa and C. decisiim, the latter from New York. Sinistral speci-
mens of an unidentified species of Campeloma were described
by Hand (Naut. 41: 106-107. 1928) and by Meyer (Naut.
41: 107. 1928) from Chicago. Pilsbry (Naut. 10: 118. 1897)
commented in regard to the rarity of reversed adult shells of
Campeloma that, "This doubtless indicates that the reversed
condition is an unfavorable one for maturation." — Ralph W.
Dexter, Dept. of Biology, Kent State Univ., Kent, Ohio.
PUBLICATIONS RECEIVED
EuTHYNEURA. By Adolf Zilch. Handbuch der Palaeozoologie.
Gastropoda, by Wilhelm Wenz, continued Teil 2, Lieferung 1,
pp. 1-200 + I-XII, 701 figs. 1959.— This issue carefully surveys
the shell-bearing (fossil forming) opisthobranchs and Pulmon-
ata, up through the geophile Orthurethra and Heterurethra.
For each genus or subgenus included, the type species, a brief
description of its shell, and its distribution in time and space,
are given. Shell descriptions for the major taxons are necessarily
vague, and notes on the animals, about which little can be
learned from paleozoology, mainly are limited to habitat and
external form. The figures of the type species show the shape ex-
cellently, but sometimes are a little weak on details, such as sculp-
ture and apertural armatures. However, they do serve well to illus-
trate this systematic outline, which, of course, is not meant to
be used for specific identifications. In the limnophiles, the Lan-
cinae are included in the Lymnaeidae, but the Ancylidae are
divided into 5 families. Among the last is the 'Terrissiidae,"
although unfortunately Hannibal, 1912, made both Laevapicinae
and its "type genus" prior. In the pupilloids, Steenberg, 1925,
is followed mainly. Of course, his "Chondrinidae" is subsequent
to Gastrocoptinae Pilsbry, 1918, and even the usage of Chondrina
is open to doubt (Cf. Pilsbry, 1935, Man. Conch. 2<9:203) . In
general, the names of familial groups follow the old "rules;"
for example, "Trimusculidae" is substituted for Gadiniidae,
1840. On the other hand, "Cerastuinae" apparently is preferred
to Pachnodinae because of the supposed homonymy (?) of the
"type genus" of Cerastinae Wenz, 1923. Subclass Euthyneura is
divided into 10 orders, 6 or 7 of which are opisthobranchs
("Suborder" Notaspidea, p. 59) and 2 or 3 are Pulmonata
(Soleolifera omitted, p. 4) . To the latter, I would add Actophila
(p. 63) and Thalossophila (Amphibolidae &: Siphonariidae, but
not Gadiniidae, which are patelliform Limnophila) . — H. B. B.
Gebruder Borntraeger, Berlin. DM 84.
Two NEW PLEURocERiD snails from eastern Mexico. By Fred
G. Thompson. Occ. Papers Mus. Zool. Univ. Mich. 600: 8 pp.,
1 fig., 1 pi. 1959. — Lithasiopsis crassa and L. darnelli are des-
cribed from Rio Sabinas. Radulae are like northern genera. —
H. B. B.
"Taphius PRONUs" (Martens, 1873) Pulmonata, Planorbidae) .
By W. Lobato Paraense & Newton Deslandes. Rev. Brazil. Biol.
i5:367-373, 10 figs. 1958.— Shells and animals are described and
figured clearly. — H. B. B.
WILLIAM H. WEEKS SHELL COLLECTION: New lists now
in preparation; full scientific data. Send name and address
for free copies. Shells also wanted for purchase and exchange.
George E. Jacobs, 853 Riverside Drive, New York, 32, N. Y.
A CHECK LIST OF THE MARINE SHELLS OF ST. CROIX,
U. S. Virgin Islands, with random annotations. Up to date.
Lists 650 species plus 60 subspecies, including 20 new. With
70 illustrations. The "random notes" contain much new infor-
mation. Price: $4.10, post free.
G. UsTiCKE, 1 North Street, Christiansted, St. Croix, Virgin Is.
How TO COLLECT SHELLS: Published by the American Malacological Union.
$1.00. Write:
Margaret C. Teskey, Sect., Route 2, Box 318, Marinette, Wis.
Directory of conchologists. — The 1960 edition will be published the first of
that year. Price will be $2.50, postpaid. For inclusion, write:
John Q. Burch, 4206 Halldale Ave., Los Angeles 62, Calif.
THE NAUTILUS
Vol. yi April, 1960 No. 4
SEVEN PUERTO RICO CONES: NOTES AND RADULAE
By GERMAINE L. WARMKE
Institute of Marine Biology, University of Puerto Rico, Mayaguez
The cones are among the most beautiful and most interesting
gastropod mollusks. At least part of this interest comes from the
fact that they possess a poison gland and are capable of injecting
a venom into their prey by means of detachable, spear-like
radula teeth (p. 123) . Painful and even fatal bites are reported
to have been inflicted on humans (Abbott, 1950) . None of the
West Indian species, however, is believed to be dangerous to man.
The present paper describes the radulae of seven species of
cones found in Puerto Rico. Most of these species occur through-
out the West Indies; but to our knowledge, the radulae have
not been described or figured previously.
All the radulae studied are toxoglossate. Each tooth is made
up of a sheet of chitin rolled up like a paper spill. The teeth,
from 20 to 70 in number, depending on the species, are enclosed
in a radula sac. These are detached one at a time from the
radula, through the proboscis, to paralyze the prey. A highly
toxic venom flows from the large poison gland through the hol-
low tube of the tooth when the tooth is emitted by the animal.
The radula teeth studied appear to fall into four distinct
groups, as follows:
1. Point armed on one side with a barb and on the other with
a blunt-ended blade, (fig. 4) . The tooth is serrate for about half
of its length. There is usually a minute cusp at the end of the
serration. In addition, there is a spur at the base of the tooth.
C. daucus, C. spurius, C. mus, and C. juliae belong to this group.
These four species can easily be separated by the following key:
A. Serration just over I/2 length of tooth. Blade very short
C. spurius (fig. 5)
B. Serration less than i/^g length of tooth.
1. Blade slightly longer than serration, serration usually with
17-22 notches C. juliae (fig. 6)
2. Blade shorter than serration.
a. Blade about I/2 length of serration C. daucus (fig. 4)
119
120 NAUTILUS Vol. 73 (4)
b. Blade almost as long as serration, serration usually with
8-15 notches C. miis (fig. 7)
II. Tooth rather short and broad, the head armed on one side
with a small barb at the tip and three minutely serrated blades
on the other. Conus regins belongs to this group (fig. 3) .
III. Tooth small and lacks any kind of serration. The tip is
armed with a sharp barb on each side, one being about twice
as long as the other. The rest of the tooth is plain, except for
a rather large spur at the base. C. jaspedius and its forms belong
to this group (fig. 1).
IV. Tooth large, lacking serration; the shaft is cylindrical with
a simple base and no spur. The armature consists of two large
barbs, one on each side of the tip, and a long backward-point-
ing process with curved tip. C. ranunculus is in this group
(fig- 2) .
Peile (1939) suggested that the presence of the projecting spur
at the base of the tooth might serve to retain the tooth within
the proboscis when prey is attacked. However, Kohn (1956)
showed that Conus striatus, which does not possess the spur,
retains the tooth in the proboscis in feeding.
Kohn (1959) showed that the presence of the spur at the base
of the cone is generally correlated with feeding on eunicid and
other tube-dwelling polychaete worms. Since eunicids burrow
in the coral and coral-reef rocks, Kohn suggests that the spur
may help the predator in extracting the worm from its burrow.
At least one of the Puerto Rican species, Conus regius, was found
to contain a large number of polychaete bristles in its alimentary
tract, and the radula teeth of this species do have spurs at their
base. (Table 1 and fig. 3) .
About 100 live or preserved cones were available for this study.
Notes which may prove of interest are included for each species.
Conus regius Gmelin
A common species found living under rocks and in coral reefs
in 4 to 10 feet of water. The animal is blood red in color. The
cones studied ranged in length from 19 mm. to 67 mm. The
radula sac contained from 25 to 32 teeth, most of them stained
yellowish brown. The teeth increased in length with the size
of the shell. In the 19 mm. cone the teeth measured 0.525 mm.
in length; in the 67 mm. cone they measured 2.1 mm. The
largest variation in size of teeth within one shell was 0.07 mm.
In the 67 mm. cone, the poison gland measured 20 mm. in
April, 1960 nautilus 121
length, the radula sac 10 mm. in length, and the tube leading
from the poison gland to the buccal mass was 290 mm. long
when stretched out.
CoNUs Mus Hwass
This is one of the most common cones in Puerto Rico. It is
found crawling on rocks and in crevices in a few feet of water.
The animal is dark red, speckled with purplish red. The
cones studied ranged in size from 15 mm. to 36 mm. in length.
The radula sac contained from 32 to 35 teeth, which ranged in
length from 0.379 mm. (15 mm. shell) to 1.106 mm. (36 mm.
shell) . The teeth within the same cone vary little in size, 0.03
mm. being the largest variation observed.
In the 36 mm. cone the poison gland measured 9.5 mm. in
length, the radula sac 5.5 mm. in length, and the tube leading
from the poison gland was 95 mm. in length when stretched
out.
During March, 1959, Conus mus was found with egg cases.
The cones were in a rock depression, in about 2 feet of water,
and the cases were attached to the rock. The egg capsules were
light cream colored, rectangular in shape, and measured approx-
imately 10 mm. in length by 7 mm. across. Each case contained
over 500 eggs, the majority still in the 1-cell stage, but some
had divided into 2 cells, and a few had reached the 4-cell stage.
Conus jaspedius Gmelin
Most of the specimens used for this study were obtained from
shallow dredgings on sandy or mud bottoms. The animal is
yellow, speckled with black. The cones ranged in length from
19 mm. to 30 mm. and varied from smooth (without pustula-
tion) , to intermediate (some degree of pustulation) , to verrucose
(pustulose) .
The radula sac contained from 52 to 70 teeth. The teeth of
these three forms are identical in shape, and the length of the
teeth is proportional to the size of the cone, regardless of its
form. (See Table 1) . This similarity in tooth structure would
seem to indicate that Conus verrucosus is not a distinct species
from Conus jaspedius, but rather a genetic form as suggested
by Abbot (1958).
122 NAUTILUS Vol. 73 (4)
Table 1. Comparison of tooth and shell length in smooth,
intermediate, and pustulose forms of Conus jaspedius Gmelin.
Shell length Average length of
in mm. teeth in mm,.
Smooth 19 0.31
Pustulose 20 0.34
Intermediate 21 0.34
Smooth 21 0.36
Intermediate 22 0.38
Pustulose 26 0.41
Smooth 30 0.44
Conus ranunculus Hwass
This is one of the rarer species of cones from Puerto Rico.
All our information is based on one preserved specimen collected
by M. McDowell, on Mona Island, 40 miles west of Puerto Rico.
The specimen measured 50 mm. in length and the operculum
11 mm. The animal had a bright orange proboscis, the edge of
the mantle was yellow-orange, and the rest of the animal was
cream colored, splotched with brown. The periostracum was
thin and Hght yellow in color. The radula sac measured 13 mm.
and contained 34 teeth, averaging 5.17 mm. in length.
In this species, the muscle at the base of the tooth is very
long and strong.
Conus spurius Gmelin
Dead shells of this species are fairly common on the west
coast, where there are reefs offshore. Our studies are based on
a freshly-killed specimen, 44 mm. in length, which had a 14
mm. operculum. The periostracum was light brown, thin, flaky,
and transparent. The animal was cream-white in color. The
radula sac contained 38 teeth, measuring from 1.085 mm. to 1.15
mm. in length.
Conus daucus Hwass
Like C. spurius, dead shells are often found on the beaches,
but live shells are hard to find. The animal is bright orange
throughout. The tiny black eyes are almost at the tips of the
tentacles. The radula teeth measured from 0.995 mm. to 1.20
mm. in length in the 45 mm. shell.
In this species, the muscle at the base of the tooth appears to
be weak, since not one of the teeth retained the muscle when
April, 1960
NAUTILUS
123
Camera lucida drawings of single radula tooth from 7 species of Puerto
Rican cones. 1, Conus jaspedius (actual size of tooth .34 mm., extracted
from a 21 mm. cone) . 2, C. ranunculus (tooth 5.17 mm. from 50 mm. shell) .
Part of shaft omitted in figure because of extreme length. 3, C. regius (tooth
2.1 mm. from 67 mm. shell). 4, C. daucus (tooth 1.2 mm. from 45 mm.
shell). 5, C. spurius (tooth 1.15 mm. from 44 mm. shell). 6, C. juliae
(tooth .63 mm, from 20 mm. shell) . 7, C. mus (tooth .75 mm. from 30
mm. shell) .
124 NAUTILUS Vol. 73 (4)
they were teased away from the radula sac.
One specimen found at Aguadilla remained alive in a dish of
salt water for over 2 weeks. The animal was most active at
night. When the shell was placed with the aperture down on
the bottom of the dish, the animal would turn a somersault by
pushing with its foot until it flipped up in an almost verticle
position, with the spire of the shell resting on the bottom of
the dish.
CoNus juLiAE Clench
This little cone is usually found on the west and northwest
coasts, where there are reefs offshore. Specimens of this species
remained alive for several days in the laboratory. The color of
the animal is bright orange-red. The specimens studied were
20 and 27 mm. in length. The radula sac contained 20 teeth,
which measured from 0.63 to 0.69 mm. in length.
In the 20 mm. cone, the poison gland measured 5.5 mm. in
length and the radula sac 2.0 mm. in length.
Acknowledgements: This work was made possible through the
kindness of many persons, who donated or loaned their shells
to the Institute of Marine Biology for this study. The author
is most grateful to Ted and Lois Arnow, F. St. Clair, Merrill
Arbuckle, M. McDowell, Genie Garrison, Amy and K. O. Phares,
Gordon Usticke, General Merwin H. Silverthorn, Jorge Rivera
Lopez, A. J. Ostheimer, 3rd., A. L. Laurence, James and Betty
Bradford, and Charles Cover.
Christine Boyce, of the College of Agriculture and Mechanic
Arts, University of Puerto Rico, made all the drawings.
Literature Cited
Abbott, R. Tucker. 1950. The venomous cone shells. Science
Counselor. Dec. 1950.
1958. The marine mollusks of Grand Cayman Island, British
West Indies. Mon. Acad. Nat. Sci. Philadelphia No. 11: 1-38.
Kohn, Alan J. 1956. Piscivorous gastropods of the genus Conus.
Proc. Nat. Acad. Sci., 42: 168-171.
1959. The ecology of Conus in Hawaii. Ecological Mon.
2P(1): 47-90.
Peile, A. J. 1939. Radula notes VIII. Conus. Proc. Malacol. Soc.
London 23: (6): 348-356.
April, 1960 nautilus 125
NEW CALIPYRGULA FROM PLEISTOCENE OF TEXAS
AND NOTES ON COCHLIOPA RIOGRANDENSIS
By a. B. LEONARD and TONG-YUN HO
Continued studies of the large assemblages of fossil shells from
Pleistocene deposits in the Pecos River Valley in southwestern
Texas, jointly sponsored by The University of Texas Bureau of
Economic Geology and The National Science Foundation (con-
tract number NSF-G3481), have brought to light another un-
described species, here assigned to the genus Calipyrgula for
reasons which were discussed in a previous paper (Leonard and
Ho, 1960). In the course of these studies, both fossil and living
examples of Cochliopa riograndensis Pilsbry and Ferriss were
obtained. We also include in this paper our observations on this
little-known species.
Calipyrgula circumstriata, new species. Plate 12, figs. 1-3
Diagnosis: A minute hydrobiid gastropod, having an elongate,
narrowly conic, imperforate, or narrowly rimate shell of 6 or 7
rounded whorls; aperture small, ovate, slightly angulate above;
peristome simple, slightly reflected over minute umbilicus and
adherent to last whorl above; suture simple, well incised; proto-
conch finely granulose, remaining whorls generally bearing 5 or
more narrow spiral ridges of somewhat irregular size and
spacing.
Calipyrgula circumstriata closely resembles C. hihbardi Leon-
ard and Franzen (1944, p. 19) , but the latter lacks the surface
sculpture characteristic of C. circumstriata. It differs from C.
pecosensis Leonard and Ho (1960, p. 110) in having fewer
whorls, less slender form, and spiral sculpture. From species of
Tryonia, C. circumstriata differs by having spiral, rather than
vertical sculpture, and by having well-rounded, rather than
shouldered whorls.
Holotype: (PI. 12, Fig. 1) Catalogue no. 11301, University of
Kansas Museum of Natural History, obtained by A. B. Leonard
and Tong-yun Ho, 4 June, 1959. Original no.: ABL 1004.
Description of holotype: Shell minute (less than 5 mm. in
length) , narrowly conic, imperforate, having 7 well-rounded
whorls; aperture ovate, narrower and angulate above; peristone
simple, slightly reflected over umbilicus and adherent to last
whorl above; protoconch of one and one-half whorls finely gran-
ulose, remaining whorls bearing narrow, somewhat irregularly
spaced spiral ridges ranging in number from 3 on second whorl
126 NAUTILUS Vol. 73 (4)
to 12 on last; last three whorls having a few indistinct vertical
ridges crossing spiral sculpture; extremely fine and numerous
vertical growth lines apparent on all whorls save protoconch;
suture simple, well incised.
Paratypes: No. 11302, UKMNH. (Plate 12, figures 2, 3). Con-
siderable variation occurs among the several hundred paratypes
from the same locality; the extremes are illustrated. Fig. 2
exemplifies the obsolescence of the spiral lines on some shells,
while fig. 3 shows another extreme, but relatively rare, variation
in which vertical ridges are conspicuously developed. Inter-
gradation occurs among the variants; most shells, however,
closely resemble the holotype. Variations in dimensions are illus-
trated by the following measurements, in millimeters.
Height Diameter No. of
Length Diameter aperture aperture whorls
Holotype 4.9 2.1 1.5 1.2 7
Paratype 4.3 1.8 1.3 1.0 61/2
(fig- 2)
Paratype 3.7 1.4 1.2 0.7 6
(fig- 3)
Paratypical specimens have been deposited at the Museum of
Comparative Zoology, Harvard University, Cambridge, Massa-
chusetts, and at the United States National Museum, Wash-
ington, D. C.
Type locality: Late Pleistocene terrace deposits in right bank
of Pecos River, one-fourth mile above mouth of Independence
Creek, on Chandler Ranch, Terrell County, Texas.
Areal and stratigraphic distribution: We found Calipyrgula
circumstriata at: the type locality; on W. C. Dunlap Ranch,
about 12 miles south-southeast of Sheffield, in Terrell County;
and near bridge on U. S. Highway 290, 3 miles southeast of
Sheffield, Pecos County, Texas. At the last mentioned locality,
a single specimen of C. circumstriata was found associated with
C. pecosensis, although the latter species occurs abundantly in
the late Pleistocene terrace along the Pecos River, as far north
as northern Reeves County (Leonard and Ho, 1960, p. 112).
Each of the three localities of occurrence of C. circumstriata is
in the late Pleistocene terrace in which the Pecos River is
presently shallowly entrenched.
Inasmuch as C. circumstriata has been found in deposits along
the Pecos only at places where that river has begun its entrench-
April, 1960 NAUTILUS 127
ment in Cretaceous rocks as it approaches the deeply entrenched
Rio Grande, we may infer that C. circumstriata inhabited the
clear, cool, rocky streams characteristic of certain tributaries
(such as Independence Creek) of the Pecos River. This inference
is strengthened by the observation that at the Chandler Ranch
(type) locality, C. circumstriata occurs with fossil Cochliopa
riograndensis Pilsbry and Ferriss; the latter is presently living in
Independence Creek. Efforts to find living C. circumstriata have
not been successful.
Cochliopa riograndensis Pilsbry and Ferriss. PI. 12, figs. 4-7,
text figs, 1-4.
Pilsbry and Ferriss (1906, p. 171) described Cochliopa rio-
grandensis from shells found at two localities near the Rio
Grande, Val Verde County, Texas. The original description
was based on "dead" specimens taken from drift debris. For
this reason, they gave no information about habitat, or about
anatomical details of radula and operculum. To our knowledge,
neither the radula nor the operculum has been described and
figured. We deem it advisable, therefore, to contribute our ob-
servations to a further knowledge of this little-known species.
Shortly after the discovery of fossil shells of C. riograndensis
(UKMNH. no. 11343) at the type locality of Calipyrgula cir-
cumstriata, we found nearly 30 specimens of living C. riogran-
densis in Independence Creek, near the bridge on Texas High-
way 349, about 16 miles south of Sheffield, in Terrell County,
Texas (UKMNH. no. 11636, 4 June, 1959, original no. TYH
30) and no more than a mile from the place where the fossil
shells were collected. The snails were living in relatively slug-
gish water about six inches deep in the clear, cool, fast-flowing
stream. C. riograndensis was here restricted to the edge of the
stream, and was fairly abundant under cobbles of limestone,
logs, and in aquatic vegetation. Associated with C. riogranden-
sis were Lymnaea hulimoides techella (Haldeman) , Gyraulus
similaris (Baker), Physa anatina Lea, and Ferrissia shimekii
(Pilsbry) . To our knowledge, this occurrence of C. riograndensis
is a northern extension of its previously known range.
Operculum (PI. 13, fig. 1): Horny, thin, roundly ovate, all
sides regularly rounding upward to a semiarcuate apex; whorls
5, the last about triple the width of penultimate whorl, inner
128 NAUTILUS Vol. 73 (4)
whorls narrower and increasing regularly in width; sculpture
consisting of coarse unequidistant, obliquely radiating growth
lines crossed by very fine spiral lines that are visible only on
the last half of the last whorl; nucleus small, circular, more or
less granular, placed about 6/13 the distance from base to apex,
somewhat toward the left side; attachment to operculigerous
lobe indicated by a thickening about a third of the distance from
border to nucleus; thickening raised above the general level
of the operculum; greater diameter from 0.8 millimeters to 1.3
millimeters.
Measurement of figured specimen: greater diameter, 1.3 mm.;
lesser diameter 1.1 mm.
Radida (PL 13, figs. 2, 3, 4): Central tooth (fig. 2): Width
about twice height; lateral angles produced and sharply tri-
angular; ventral margin having large, triangular process extend-
ing from base; lateral ridge large, wide, having two basal den-
ticles, anterior one larger than posterior; reflection wide and low,
having 11 sharply elongate cusps, central cusp about three times
as wide as lateral cusps, reaching nearly 2/^ distance from dorsal
margin to ventral margin; lateral cusps smaller, gradually in-
creasing in size toward center. Lateral tooth (fig. 3) : Body
slightly wider than high, having sharply rounded lobe extending
from inner basal part of body; large, U-shaped, pit-like depres-
sion in center of body; peduncle almost twice as long as body,
gradually narrowing toward end, having elongate ridge parallel
with long axis, but displaced toward outer margin; reflection
wide and low, bearing two inner cusps, one large central cusp
and four outer cusps, all cusps sharply elongated and diminish-
ing in size toward margin of tooth. Inner marginal tooth (fig. 4) :
Falcate, having long and narrow body; peduncle spatulate, less
than twice length of body, bearing elongate centrally placed
ridge parallel to long axis and branching near base of body;
reflection wide and low, bearing 20 to 30 fine, sharply elongate
cusps; cusps near central portion of reflection relatively large
and acute, those on lateral margin becoming smaller.
Measurements of figured radula in microns
Central tooth. Width 32
Height 16
Length of cusp row 20
Lateral tooth. Height 58
Length of cusp row 20
Marginal tooth. Height 51
Length of cusp row 24
We found no significant differences between the fossil shells
of Cochliopa riograndensis (PI. 12, figs. 4, 5, 6), and those from
living individuals (PI. 12, fig. 7) . The spiral color bands on
NAUTILUS 73 (4)
PLATE 12
Figs. 1-3, Calipyrgula circumstriata Leonard & Ho: 1, holotype shell.
2, 3, paratypes from same locality, to show extremes of variation in size, pro-
portions and surface sculptine. Figs. 4-7, Cochliopa riograndensis Pilsbry &
Ferriss: 4-6, fossil shells. 7, apical view of shell from living animals.
NAUTILUS 73 (4)
PLATE 13
Figs. 1-4, Cochliopa riograndensis Pilsbry & Ferriss: 1, outer view of oper-
culum, X 26. 2-4, radular teeth (x 1000) : 2, central; 3, lateral; 4, inner
marginal.
Figs. 5-7, Choanopoma radiosum (Morelet) : normal adult (5) and two
elongate freaks (6, 7) . Fig. 8, Coelocentnun fistulare (Morelet) nuclear
whorls.
April, 1960 nautilus 129
shells from living snails differ in number and in width; a few
individuals had broad colorless bands on the ventral aspect of
the shell. The protoconch is invariably colorless, and lacks also
the spiral ridges characteristic of the remaining whorls. Color
bands are, of course, absent on fossil shells since the epiostracum
is not preserved, and the spiral ridges are often faint, as they are
on the figured specimen.
Several of the living examples of C. riograndensis from Inde-
pendence Creek contained embryos in advanced stages of devel-
opment; these varied from 30 to 50 in number in adult snails.
Literature cited
Leonard, A. Byron, and Franzen, Dorothea S. 1944. Mollusca
of the LaVerne Formation (Lower Pliocene) of Beaver
County, Oklahoma. Univ. Kansas Sci. Bull., 30, pt. 1, no. 2:
15-39, pis. 4, 5. May 15.
Leonard, A. Byron, and Ho, Tong-yun. 1960. A new species of
Calypyrgula (Hydrobiidae) from the Pleistocene of Texas.
Nautilus, 73 (3): 110-113, pi. 11. January 15.
Pilsbry, Henry A., and Ferriss, J. H. 1906. Mollusca of the
southwestern states. H. Proc. Acad. Nat. Sci. -Philadelphia,
58: 123-175, pi. 9. July 24.
^ I ^^>5- '
NON-MARINE MOLLUSKS FROM BRITISH HONDURAS
By fritz HAAS and ALAN SOLEM
Chicago Natural History Museum
Outside of the few records listed by von Martens (1890-1901),
no non-marine mollusks have been reported from British Hon-
duras. The availability of some material collected by Ivan T.
Sanderson in 1940 and W. D. Thomas in February, 1959, has
prompted this short list. Mr. Thomas collected at the Rio Frio
Cave East, about two miles from Augustine on the Mountain
Pine Ridge, Cayo District, British Honduras. Mr. Sanderson
collected at several localities as listed under the specific names.
The material is all deposited in Chicago Natural History Mu-
seum, with a set of duplicates of Mr. Thomas's shells in the
Museum of Comparative Zoology, Harvard.
Most of the species are common Guatemalan forms, but sur-
prisingly, some species previously known only from the Bay
130 NAUTILUS Vol. 73 (4)
Islands off Honduras (see Ancey, 1886 and Richards, 1938)
were represented. Some peculiarly incrusted fresh-water shells
are reported on separately (liaas, 1959) . A few forms are figured
as an aid to future workers.
Pomacea belizensis (Crosse and Fischer) . Stann Creek Valley.
Pomacea ghiesbreghti ghieshreghti (Reeve) . Kate's Lagoon.
Neocyclotiis dysoni dysoni (Pfeiffer) . Rio Frio Cave, Kate's
Lagoon, Bokowina.
Choanopoma (C.) rigidulum (Morelet) . Rio Frio Cave.
Choanopoma (Colobostylus) radiosum (Morelet). (PI. 13,
figs. 5-7) Rio Frio Cave.
Two non-decorticated, elongate freak specimens are shown
with a normal individual.
Choanopoma (C hoanopomops) largillierti (Pfeiffer) . Che-
tumal.
Helicina (H.) flavida (Menke) . Rio Frio Cave.
Helicina (Oxyrhombus) amoena (Pfeiffer) . Rio Frio Cave,
Double Falls, Bokowina.
Lucidella {Poeniella) lirata (Pfeiffer) . Rio Frio Cave.
Pyrgodomus simpsoni (Ancey) . Rio Frio Cave.
Described from Utilla Island, Honduras, by Ancey (1886:
253-254) , this species differs from the Guatemalan P. microdina
(Morelet) only in having a more angulate periphery. Possibly
they are only subspecifically distinct.
Pachycheilus (P.) planensis Lea Rio Frio Cave.
Living examples were collected from the stream issuing from
the cave and mummified specimens (see Haas, 1959) were found
inside the cave.
Pachycheilus (Glyptomelania) obeliscus (Reeve) . Sibun R.,
tributary of Eastern Branch, Cayo District (W. D.
Thomas!) .
Synopeas beckianum (Pfeiffer) . Rio Frio Cave.
Leptinaria (L.) fordiana (Ancey) . Rio Frio Cave.
Leptinaria (L.) lamellata (Potiez and Michaud) . Stann Creek
Valley.
Spiraxis (Volutaxis) sulciferus (Morelet) . Rio Frio Cave.
Streptostyla thomsoni (Ancey) . Rio Frio Cave.
Streptostyla dysoni (Pfeiffer) . Rio Frio Cave.
Euglandina sp. Rio Frio Cave.
Euglandina cylindracea (Philippi) . Chetumal.
Hyalosagda haldemaniana (C. B. Adams) . "British Honduras."
Unless Sanderson's locality is in error, this species has
probably been imported on plants from Jamaica.
April, 1960 nautilus 131
Microceramus (M.) consisus (Morelet) . Chetumal.
Brachypodella (B.) sub tills (Morelet) . Rio Frio Cave.
Coelocentrum (C.) fistulare (Morelet). (PL 13, fig. 8). Rio
Frio Cave. The opportunity is taken to figure the nuclear
whorls.
Orthalicus princeps crossei Martens. Rio Frio Cave, Double
Falls.
Drymaeus (D.) alternans honduranus (Pfeiffer) . Rio Frio
Cave, Kate's Lagoon.
Bulimulus (B.) corneus (Sowerby) . Chetumal.
Averellia {Trichodiscina) coactiliata (Deshayes) . Rio Frio
Cave, Chetumal.
We are indebted to Mr. W, Donald Thomas of Balboa, Canal
Zone, for the opportunity to study this material and to Mr.
A. H. Anderson, Archaeological Commissioner of British Hon-
duras, for information concerning the Rio Frio Cave site.
Bibliography
Ancey, C. F. 1886. Une excursion malacologique sur le versant
atlantique de Honduras. Ann. de Malacologie 2:237-260.
Haas, Fritz. 1959. Ueber Schneckenmummien. Arch. f. Mollusk.
(95(4/6) : 159-161, 5 textfigs.
Martens, E. G. 1890-1901. Land and freshwater MoUusca. Biol.
Cent. Americana.
Richards, H. G. 1938. Land mollusks from the island of Roatan,
Honduras. Proc. Amer. Phil. Soc, 79 (2) : 167-178, 3 pis., 1 fig.
QUANTITATIVE SAMPLING
By CHARLES B. WURTZ
Consulting Biologists, Philadelphia 2, Pa.
In this paper, the term quantitative is used in the sense of the
statistician and not in the lay sense of simply large numbers of
organisms taken in the field.
Mollusks are collected, as are all other organisms, for two
reasons only. First, and of no interest here, they are collected
simply as curiosities of natural history. Second, they are col-
lected for the purpose of studying all, or some aspect of the
population. In this latter kind of collecting the only practical
method of collecting is by means of samples. The samples rep-
resent observations. In so far as it is practicable to do so, and
only in so far as practicable, samples should be collected quanti-
132 NAUTILUS Vol. 73 (4)
tatively. Actually, an ideal quantitative sample must fulfill three
fundamental criteria (cf. Simpson and Roe, "Quantitative Zool-
ogy" (1939) :
1) Collections must be made from a single, well-defined
population only.
2) Collections must include all the variations within the
population.
3) The variations must occur in the sample with the same
relative frequency they have in the whole population.
Good sampling starts with a definition of the population to
be sampled, and good collecting involves as complete a collection
of all the elements in the definition as possible.
The first criterion of the ideal sample is commonly achieved.
It is easy to limit our collections to a single, well-defined pop-
ulation. All that is necessary is to specify the geographic area,
to include all the ecological habitats, and to limit the material
to be collected to a feasible group. The crucial part of this
criterion is the exact description of the population to be studied.
The second criterion is a product of technique. This ideal may
or may not be achieved by a persistent collector with nimble
fingers and a supply of sampling devices. The problem en-
countered in resolving this criterion is the ability to recognize
objectively when effective satisfaction of the criterion has been
realized. Subjectively, no one can recognize whether or not this
criterion has been fulfilled.
The third criterion is the area in which we move from biol-
ogy into statistics. We cease dealing with natural populations
as such and begin working with frequencies. The statistics of
quantitative analysis fall beyond the scope of this paper.
The first criterion of the ideal sample is almost wholly biolog-
ical, and the biologist can resolve it to his complete satisfaction.
The third criterion is almost wholly statistical, and the statisti-
cian can resolve it to his complete satisfaction. But the criterion
falling between these two appears to be in a never-never land
of inarticulation. The collector who works intensely goes home
convinced that he has collected all the species with all their
variations that occur within the defined population. The statis-
tician who receives this material for processing is inclined to
doubt that this has been accomplished. This, apparently, is in
accord with the experience of each.
April, 1960 nautilus 133
In any quantitative sampling, there are only three basic ap-
proaches. These are: 1) samples based on a fixed and predeter-
mined unit of area, 2) samples based on a fixed and pre-
determined unit of mass, and 3) samples taken through a fixed
and predetermined period of time. Everything else is only a
modification of one or more of these.
Plankton can be measured volumetrically. Time can be used
for organisms that can be trapped or for organisms that will
invade and populate a sterile surface. However, these are re-
stricted samples. The techniques based on area are the only ones
that are applicable to all kinds of organisms.
Two years ago the molluscan population of a New Jersey
river was sampled. The area collected was a shaded, rubble-
bottomed, low-gradient stretch of a small coastal river. Although
one pool had a depth of about 15 feet the stream held a littoral
fauna only. Rooted aquatic vegetation occurred in extensive
patches. The stream was not polluted, although some organic
enrichment was evident. From the collected area, the following
13 species of mollusks were taken:
Campeloma decisum Gyraulus parvus
Amnicola limosa Physa heterostropha
Goniobasis virginica Laevapex fuscus
Lymnaea humilis Elliptio complanatus
Pseudosuccinea columella Anodonta cataracta
Helisoma trivolvis Sphaerium transversum
Helisoma anceps
At this location on the eastern seaboard in the latitude of
New Jersey, any experienced collector would assume that all
the species of the defined (molluscan) population that might
be anticipated were, in fact, found. The only exception might
be Pisidium casertanum.
The sample was taken on an areal basis. The collection was
made so that material from every ecological habitat was included.
The qualitative collecting was quantitatively tempered by insert-
ing a time factor. Each habitat was collected until 30 minutes
elapsed without the addition of a species not previously taken.
If a purely quantitative sample had been sought, only two
techniques, or a modified combination of them, could have
been used. The first would be the quadrat method; the other
the transect method. However, the physical structure of the
134 NAUTILUS Vol. 73 (4)
collected area would have eliminated the proper use of either
of these.
In quantitative collecting, the sampling devices must be uni-
form. No sampling device ever invented would have taken a
uniform and constant sample from this locality because the
physical structure of the stream militated against this. Collecting
was done until the collectors were satisfied subjectively that
no further additions to the fauna could be found.
It is significant to note that the accumulated field experience
of most collectors is such that the possibility of designing a
single piece of equipment that would uniformly sample even a
small percentage of all the areas collected in a field season is
virtually nil. This is a very real handicap for absolute quantita-
tive sampling. Because of this, all collecting must, of necessity,
be biased. Furthermore, because of this, it is theoretically
impossible to take samples that are amenable to quantitative
techniques.
Aside from the inability to take quantitative samples because
of mechanical limitations, there is another aspect that must be
considered. This is the concept of randomness. The heterogeneity
of the environment and the behavioral characteristics of each
species are opposed to random collecting. A good quantitative
sample must be chosen at random, otherwise bias is introduced.
Actually natural waters are not disposed at random and aquatic
organisms are not dispersed at random. There are real causes
behind their occurrence and distribution. This is a built-in
bias that cannot be overcome.
An experienced biologist would anticipate the presence of the
species actually collected from this locality, but he certainly
would not seek them by random collecting. He would look for
them in the habitats where they live. The three species of clams,
along with Campeloma decisum, are burrowing forms. They can
only be taken by digging into the substrate. The limpet,
Laevapex fuscus is virtually sedentary. Conversely, the other pul-
monate snails are quite peripatetic in their foraging, as are
Amnicola limosa and Goniobasis virginica. Some of these tend
to be gregarious while others tend to be solitary. The lymnaeids
do not hesitate to leave the water and forage on sloping mud
banks, etc. Yet they certainly must be included in the fauna of
April, 1960 nautilus 135
an aquatic habitat. Evidently the collector resolves the second
fundamental criterion of the ideal sample to his own satisfaction,
but he does not do this quantitatively in a statistical sense, and
his data should not be used to resolve the third criterion.
To sample, randomly, a population composed of burrowing,
sedentary and peripatetic species, as well as solitary and gre-
garious species, and have the results mean anything quantita-
tively, is a highly dubious procedure.
The second fundamental criterion for the ideal quantitative
sample cannot be mechanically realized as a standard practice.
Quantitative samples can only be taken if the population is
defined with those restrictive terms which, by their very nature,
eliminate the total faunal complex from the population. A com-
plete fauna cannot be included in the definition.
The collector who wishes to handle his material quantitatively,
and the statistician who wishes to analyze such material, must
keep two thoughts constantly in mind. First, collections produced
by competent qualitative collecting will yield approximate
quantitative data. This approximation is sufficiently accurate
for use in a textual description of the population. Second, a
perfect sample in the quantitative sense is impossible of collec-
tion because the introduction of bias in sampling cannot be
avoided, and this defeats the concept of randomness.
In addition to these very severe limitations, it is worth men-
tioning that statistical analyses are very time-consuming, and
therefore expensive. Furthermore, only very rarely do statistical
analyses reveal facts that are not already apparent from really
competent qualitative collecting. Usually such studies do not
produce results that are sufficiently advanced over qualitative
work to justify the time and expense involved.
MOUNTING MINUTE RADULAE
By R. D. turner
Museum of Comparative Zoology
The preparation of radula mounts of minute snails (i.e.,
those 5 mm. and less in size) is generally a time consuming,
extremely tedious task, and, in the process of moving the radula
from one solution to another, it is often lost. This is particularly
136 NAUTILUS Vol. 73 (4)
frustrating when only one or two specimens are available. The
following technique is practically foolproof and produces excel-
lent results.
If the specimens are preserved in alcohol, the animal can
often be extracted from the shell using a minuten pin mounted
in a glass rod and hooked at the point. Using the slide on which
it is planned to mount the radula, start at the left and place
in a line, first a small drop of KOH or NAOH, then a drop of
water, a drop of 70% alcohol and a drop of polyvinyl alcohol
mounting medium. Place the specimen in the NAOH to soften
the animal until it is clear but can still be moved without falling
apart. The length of time required for this varies with the
material and it must be watched under the scope. The specimen
is then slid from the NAOH to the drop of water to remove
the base, then on to the drop of 70% alcohol for a couple of
minutes. From here you can slide it into the drop of mounting
medium. The radula can be seen inside the head and can be
easily teased out by using "minuten pin dissecting needles."
Move the radula to a clear spot and, if sufficiently large, one
possibly may dissect a portion of the radula in the medium.
The operculum, if present, can also be removed easily and safely
and moved to a clear spot. The remainder of the animal can
then be removed from the mounting medium if desired.
If the animal cannot be removed from the shell, only two
small drops need be added to the slide so that, beginning at
the left, there is a drop of weak acid to dissolve the shell, a
drop of water to get rid of the acid and then a drop of weak
base to soften the animal and continuing as before.
By this procedure, the entire process may be carried through
to completion without ever lifting the specimen or taking your
eye from it. To this point, all work is done under the highest
power of a binocular dissecting scope. Once the cover slip is in
place, the remainder of the slide can be wiped clean and the
radula can be examined under the high power of a compound
microscope. Since PVA does not harden rapidly, one can, with
a needle and light pressure on the cover glass, rotate the radula
and examine it in various positions while making illustrations.
When the study is completed, the slide should be ringed with
duco cement or fingernail polish. Staining the radula usually
April, 1960 nautilus 137
is not necessary when using PVA mounting medium but if this is
desired, a drop of acid stain can be added to the line just prior
to the drop of PVA. This procedure has reduced a process, which
may run into hours devoted mainly to groping for lost radulae,
to less than 20 minutes and so far, since using this method, I
have not lost a specimen.
LAND SNAILS FROM CENTRAL SOUTH FLORIDA
By DRUID WILSON
The area around Lake Moody, two miles north of Frostproof
in Polk County, is in a part of the Central Highlands or High-
lands Ridge of Florida locally called "the Ridge." Originally,
the open pine woods of the high surrounding hills graded in
places and at slightly lower elevations into black-jack oak
(Quercus laevis) woods and near the lake into a narrow zone of
palmetto thickets. Most of the area is now planted in citrus
groves. The soil is mapped as Norfolk fine sand (F'owler and
others, 1932, Soil map of Polk Co.), but some minor variation
is evident. The soil is noticeably lighter wherever black-jack
oaks predominate. During the last 25 years, changing cultural
practices doubtlessly have caused fluctuations in the populations
of the snail species. The soil is naturally deficient in calcium
and application of such materials as basic slag and dolomite to
the soil of citrus groves appears to affect favorably some of the
species. Use of basic slag in the 1930s was never really common,
but dolomite is now almost universally used. Desultory spraying
with lime-sulfur solution or dusting with sulfur in these depres-
sion years probably had little or no effect on species living in
citrus gi'oves. Present day intensive spraying with oil emulsions
and compounds of copper and zinc and application of copper
sulfate to the soil apparently are injurious to some of the species
or inhibit the growth of their food.
Polygyra cereolus (Muhlfeld) . The species was first noticed
in 1940 in a young citrus grove adjacent to the highway near a
roadside fruit stand. All the individuals examined have been
of the "normal" form Polygyra cereolus carpenteriana (Bland)
except one within the size range of the historically typical and
138 NAUTILUS Vol. 73 (4)
larger form Polygyra cereolus cereolus (see Pilsbry, 1940, p. 584-
586) having a diameter of 12 mm. Since 1940 the species has
spread to groves nearby and along the highway which has a base
and shoulders of limestone. Pilsbry (p. 589) reports that Polygyra
cereolus occurs on calcareous soils and Polygyra septemvolva Say
on acid soils in the same districts. Establishment of a spreading
colony of cereolus, in an area of naturally acid soil deficient in
calcium, some distance from any reported occurrence, is prob-
ably due to chance introduction and the presence in the treated
citrus grove soil of sufficient calcium carbonate. All the specimens
examined have the internal lamina characteristic of cereolus.
The data given for Hubricht's conclusions (1953, p. 116) that
"the internal lamina is not a valid specific character, and P.
septemvolva and P. cereolus are one species" are unconvincing
and directly opposed to the data and opinions on the two species
given by Pilsbry (1940, p. 582-590) ,
Praticolella jejuna (Say) . A high, pine timber habitat for
this species was noted long ago (Simpson, 1885-89, p. 66; Webster,
1892) . It apparently is the only land snail endemic on the high-
est hills of the Ridge district. Before most of the land was
cleared, the species was rare and never found in colonies, but
it is now common in citrus groves. In the early 1930s, it was
excessively abundant in two citrus groves several miles apart in
which the soil had been treated with basic slag. Either dolomite
now used is less favorable, or current spraying practices and the
use of copper sulfate on the soil affect the species adversely.
Practicolella bakeri Vanatta. No living specimens have thus
far been found, but "bones" have been collected in two small
areas. A few in a black-jack oak woods one half mile northeast
of Lake Moody, now cleared, and in greater numbers about 1.5
miles further north in a citrus grove bordering on woods of
the same kind.
Euglandina rosea (Ferussac) . This species is probably native
to the thickets of the lake edge and has also been taken around
houses. None of the individuals collected exceeds the measure-
ments given by Pilsbry (1946, p. 194) for the small ecological
variety E. rosea minor (W. G. Binney) which according to Pilsbry
inhabits dry places.
Zonitoides arboreus (Say) . Specimens have been found living
April, 1960 nautilus 139
only in the crowns of the ornamental palm "Cocos plumosa,"
where it is sometimes abundant. This restricted ecological niche,
sometimes as much as 20 feet above ground, has little except
temperature in common with its surroundings. It supports along
with the snails an amazing community of which blattids, sow
bugs, earthworms, and ants are the more obvious members.
Pupisoma dioscoricola (C. B. Adams) . Formerly present in
vast numbers in citrus trees, these minute snails are now common
only in trees not subject to repeated spraying.
These species are the only land mollusks, except an unidenti-
fied slug, that I have found in the area during a period of over
35 years.
I am grateful to Dr. J. P. E. Morrison of the U. S. National
Museum for identifying the specimens of Zonitoides arboreus
and Pupisoma dioscoricola.
Bibliography
Fowler, E. D., and others, 1932, Soil survey of Polk County,
Florida: U. S, Dept. Agriculture, Bur. Chemistry and Soils,
ser. 1927, no. 39, 39 p., soil map (1927) .
Hubricht, Leslie, 1953, Land snails of the southern Atlantic
Coastal Plain: Naut., 66 (4) : 1 14-125.
Pilsbry, H. A., 1939-1948, Land Mollusca of North America
(north of Mexico) : Philadelphia Acad. Nat. Sciences, Mon.
3, V. 1, pt. 1, p. 1-574, 1939, pt. 2, p. 575-994, 1940; v. 2, pt. 1,
p. 1-520, 1946, pt. 2, p. 521-1113, 1948.
Simpson, C. T., 1885-1889, Contributions to the Mollusca of
Florida: Davenport Acad. Sci., Proc. 5 (1885-1889): 45-72,
63*-72*.
Webster, G. W., 1892, [Note on Helix jejuna]: Naut. 5 (10) : 119.
SOME OCEANIC SUB-TIDAL OYSTER POPULATIONS
By harry W. wells and L E. GRAY
Department of Zoology, Duke University
Durham, North Carolina
The oyster, Crassostrea virginica, is usually regarded as an
estuarine organism, that is, one which lives in estuaries, where
seawater is mixed with fresh water. It thrives especially well
under estuarine conditions, forming natural beds. Churchill
(1920) placed the optimum salinity for oysters between 14
140 NAUTILUS Vol. 73 (4)
and 28 o/oo. Several reasons have been suggested for this char-
acteristic distribution. A greater quantity of planktonic food
is available to oysters there than in a more marine environment.
Fresh water flowing into the estuary may bring elements that
are important to the oyster's successful completion of its life
history, such as copper which has been credited with an impor-
tant role in settling (Prytherch 1934) . Probably more important
is the inability of many oyster enemies to live in brackish waters.
The boring sponge, Cliona celata, and the oyster drill, Urosal-
pinx cinereus, are two such oyster enemies that are barred from
upstream beds by reduced salinities.
In high salinity waters of the southeastern states, oysters are
usually restricted to the intertidal zone, the deep water beds
occurring in areas of reduced salinity. This intertidal distribu-
tion has been attributed to the selection of this zone by oyster
larvae (Galtsoff & Prytherch 1927), or to a flow of chemical
elements from fresh water that stimulates larvae to attach in
the intertidal zone (Prytherch 1934), or to the failure of oyster
spat to survive at lower levels. Lunz (1943) blamed the boring
sponge, Cliona celata, for eroding shells below the low tide mark
to such a degree that development of natural oyster beds in this
lower zone is prevented. Similarly, Chestnut and Fahy (1953)
suggested that the predation of oyster spat by the oyster drill,
Urosalpinx cinereus, offers a partial explanation for this distri-
bution of oysters in high salinities.
This report describes an unusual occurrence of oysters below
the intertidal zone in coastal North Carolina waters. Certain
environmental factors which appear to be involved are discussed.
Observations: A study has been made of marine organisms
attached to metal wrecks near the outer coastline in the vicinity
of Cape Hatteras, North Carolina. The location of each collect-
ing station is indicated in Figure 1 and described below.
1 — Overlook. This unidentified wreck is located 100 yards
from the beach 2 miles north of the Pea Island National Wild-
life Refuge Headquarters, formerly the Pea Island Lifesaving
Station. Its boiler stands above high tides; other sections are
exposed only by lowest tides.
2 — Rodanthe. The wreck of Landing Ship 232, 1948, is located
in the surf on the beach 600 yards south of Chicamacomico
April, 1960 nautilus 141
Lifesaving Station. The ship is largely intact, with super-struc-
ture extending well above mean high water.
3 — Waves. The wreck of a landing ship, 1948, is located 50
yards from the beach opposite the village of Waves. The ship
is broken into sections, several of which project above mean
high water.
4 — Salvo. An unidentified wreck is located 80 yards from the
beach opposite the village of Salvo. Several pieces extend above
mean high water.
5 — Ocracoke. Wreck of the Steamer "Ariosto," 1899, is located
300 yards from the beach 2 miles southwest of the ferry slip at
the eastern end of Ocracoke Island. Its mast extends less than
two feet above mean low water.
6 — Portsmouth. Wreck of the Freighter "Miget," 1952, is
located 300 yards from the beach 3 miles south-southwest of
Portsmouth village. A large section extends above mean high
water.
Oysters were collected in numbers at the four stations situated
north of Cape Hatteras. However, no living oysters were found
on the Ocracoke wreck. At the Portsmouth wreck, living oysters
and many dead shells of Crassostrea virginica were present. While
oysters are present on wrecks in the Hatteras area, they were
scarce on wrecks south of the Cape.
On the wrecks at Rodanthe (#2) and Waves (#3) living
oysters were abundant, attached to metal parts below the low
tide level. Immediately above them was a zone of blue mussels,
Mytilus edulis, which formed the lowest truly intertidal zone.
Only abnormally low tides would expose any oysters in this
habitat; most are never exposed. Oysters occupied most of the
surfaces in this subtidal zone, with only occasional openings
where the corrosion of underlying steel or the erosion of oyster
shell by boring sponge had so weakened attachment that a cluster
of oysters had been dislodged. This layer of oysters provides
attachment for some organisms and crevices suitable for the
colonization of other species. Oyster distribution on other wrecks
followed these general lines, with the principal segment of the
population occupying subtidal surfaces, and only a minor repre-
sentation in intertidal zones. The intertidal individuals were
restricted to the lowest fringe of the intertidal zone.
142 NAUTILUS Vol. 73 (4)
On the Portsmouth wreck (#6) , Crassostrea virginica did not
constitute a large part of the community. Oysters have success-
fully attached to this wreck and survived, but they are far out-
numbered by specimens of the horse oyster, Ostrea equestris.
On these southern wrecks, O. equestris effectively replaces C.
virginica, both in numbers and in its contribution to suitable
substrate for the colonization of other organisms. On wrecks
north of the Cape, individuals of O. equestris occur, but in less
abundance than on the Ocracoke and Portsmouth wrecks.
Discussion: An analysis of the various environmental factors
helps explain this unusual distribution.
PHYSICAL FACTORS: Animals attached to these wrecks are
regularly exposed to the action of surf, its aeration, its tendency
to dislodge insecurely attached forms, and to the abrasive effects
of suspended sand. At each station oysters are exposed to oceanic
conditions — relatively stable high salinities and moderate tem-
peratures that change gradually with the seasons. Neither salinity
nor temperature exceed the tolerances of the oyster. Tempera-
ture gradients along the coast have been recorded for the Cape
Hatteras region (Parr 1933, Wells & Gray 1960) , but they seem
unlikely to be directly responsible for permitting sizeable pop-
ulations of oysters north of the Cape and none or few south of
the Cape in comparable situations. Instead, temperature may
have an indirect effect by controlling the distribution of oyster
enemies. Physical factors do not provide a satisfactory explana-
tion for the observed distribution.
BIOLOGICAL FACTORS: A large number of oyster enemies
are recognized in different parts of the world (see Korringa
1952) , They include competitors for food and space, and organ-
isms that invade the oyster meats or shell, as well as the more
obviously effective predators.
Competitors: Mussels (Mytilus edulis and Brachidontes
exustus) and barnacles (Chthamalus and Balanus species) occur
in abundance on these wrecks. By sheer numbers, they would
make attachment to a firm substrate difficult for oyster larvae.
However, they are usually attached at higher levels on the
wrecks than are the oyster populations, and the oysters are suc-
cessful in maintaining themselves in spite of their presence.
The horse oyster, Ostrea equestris, is much more abundant on
April, 1960 nautilus 143
wrecks south of Cape Hatteras. Its abundance might present
serious competition with Crassostrea virginica for suitable at-
tachment.
''Invaders": Apparently the same species of boring sponge
(Cliona species) are present on wrecks south of the Cape as on
wrecks north of the Cape. There is a greater abundance of boring
sponge colonies, however, on southern wrecks. This is probably
a reflection of the greater abundance of Ostrea equestns there,
for this species of oyster may be more heavily infested by boring
sponges, as Wells (1959) has reported at Beaufort, N. C. Shells
of oysters are usually securely attached to the wrecks, otherwise
surf action would pull them loose. Apparently Cliona has not
been so effective in preventing the development of sizeable pop-
ulations of oysters here as Lunz (1943) found in waters of
comparable salinities in South Carolina.
Predators: Although starfish of the genus Asterias are impor-
tant enemies of oysters in Long Island Sound (Galtsoff &
Loosanoff 1939) , few starfish have been found and those only
on the Rodanthe wreck where the oyster population is best
developed. Apparently, starfish have little effect upon these
populations of oysters.
Lunz (1947) has pointed out the importance of the blue
crab, Callinectes sapidus, as a predator of oysters in South Caro-
lina. Numbers of large blue crabs were often observed feeding
among organisms attached to the wrecks. They were present on
wrecks both north and south of the Cape, and were observed
feeding directly on oysters at the Rodanthe wreck. Although
crabs are abundant in this habitat and they prey on oysters, the
distribution of these oyster populations is not related to the
distribution of crabs.
Predaceous gastropods, recognized oyster enemies in many
parts of the world, are represented on these wrecks by two
species: Thais fioridana, the Gulf oyster drill, and Urosalpinx
cinereus, the Atlantic oyster drill. In the Gulf of Mexico, Thais
fioridana is a serious predator of oysters (Burkenroad 1931,
Butler 1954) . In the Hatteras area, this species was collected in
numbers from the two wrecks south of the Cape, but none were
found north of the Cape.
Although individuals have been recorded from Chincoteague,
144
NAUTILUS
Vol. 73 (4)
CAPE HATTERAS
Fig. 1. Eastern North Carolina with the Cape Hatteras region enlarged to
show the location of collecting stations.
Virginia, Thais fioridnna does not occur in numbers north of
the point of Cape Hatteras. The populations on the southern
wrecks were producing typical clusters of e^g capsules in early
June, 1959, indicating that they are capable of reproduction in
this locality. Planktonic larvae of this species could be carried
northward, but apparently have not populated the northern
wrecks. Conditions seem to be unfavorable for reproduction
north of this point. Probably temperature gradients in the
region of Cape Hatteras are responsible for preventing the
successful colonization of otherwise suitable habitats to the
north. Apparently the scarcity of oysters on wrecks south of the
Cape can be correlated with the occurrence of sizeable popula-
tions of this predator.
The Atlantic oyster drill, Urosalpinx cinereus, is recognized
as the most important enemy of oysters in many areas (Carriker
1955) , but because it cannot tolerate low salinities, oyster beds
in the upper part of many estuaries are free of this pest. The
isolation from drills provided by low salinities permits the
April, 1960 nautilus 145
development of large natural oyster beds in these areas.
Urosalpinx cinereus was abundant on the Ocracoke wreck
(#5) , while none were found on the other wrecks studied. Ap-
parently here predation by this species has prevented the main-
tenance of a sizeable oyster population below the low tide mark.
On wrecks north of Cape Hatteras, however, subtidal oyster pop-
ulations survive and thrive in the absence of this species.
Of the major oyster enemies, only drills are absent on wrecks
north of the Cape. All other species mentioned — mussels, barna-
cles, oysters, sponges, starfish, and crabs — have pelagic larvae
which could be carried by currents to a newly-foundered wreck
where they could attach and produce a thriving population.
The pelagic larvae of T. floridana are similarly capable of reach-
ing new wrecks, but are apparently barred from penetrating
north of the Cape. On the other hand, instead of pelagic larvae,
Urosalpinx cinereus produces a "crawl-away" larva which emerges
from its egg capsule as a miniature of the adult. Crawling con-
stitutes the chief means of dispersal, although Carriker (1957)
has shown that these larvae may attach to moving objects or
seaweed or be carried by the surface film. The present-day dis-
tribution of this species is attributed to its unintentional intro-
duction to new areas by man (Carriker 1955) . Urosalpinx
cinereus has been recorded from hard bottoms off the North
Carolina coast (Pearse & Williams 1951, Chestnut, in Carriker
1955) , and has been collected from hard bottom off Ocracoke
Inlet. In view of the exposure of these wrecks to wave action,
crawling over the bottom seems the most likely means of inva-
sion. However, this species would have to traverse miles of sand
bottom to reach these wrecks from known hard bottoms. The
possibility of individuals reaching one of the wrecks would
be very small, but the probability of a successful invasion would
increase with time. The Ocracoke wreck now populated by U.
cinereus, where oysters no longer thrive, is one of the oldest
wrecks. It has been subject to such invasion for sixty years.
It appears as though the abundance of oysters on wrecks north
of Cape Hatteras can be correlated with the absence of oyster
drills, which is in turn correlated with the age of a particular
wreck. In this case, the isolation from oyster drills provided
oysters on these wrecks is similar to the isolation from drills
146 NAUTILUS Vol. 73 (4)
provided oysters in upstream estuarine oyster beds through the
agency of a salinity barrier. In both instances, oysters may flour-
ish in the absence of drills. The presence of mussels, boring
sponges, and crabs has had little obvious effect on these oyster
populations.
The abundance of Ostrea equestris on the southern wrecks
may be explained by a preference of these gastropod drills for
C. virginica as their prey, and their avoidance of O. equestris as
prey. Such a preference between these two oysters is exhibited by
another predaceous gastropod, Fasciolaria hunteria (Wells 1958) .
The presence of Thais floridana on the southern wrecks sug-
gests that it is partly responsible for the observed scarcity of
subtidal oysters there. Probably this species also contributes to
restricting oysters to intertidal situations in higher salinities in
the southeastern states. Unlike Urosalpinx cinereus, Thais flori-
dana can quickly populate new wrecks by means of its pelagic
larvae, thus making the likelihood of oysters establishing sub-
tidal populations in the ocean very small south of Cape Hatteras.
Acknowledgements: The authors wish to express their grati-
tude to Mrs. Mary Jane Wells, Mr. Alex Marsh, and Mr. Bruce
Welch, who were our assistants in this study, and to the Cape
Hatteras National Seashore of the National Park Service, which
supplied useful information and facilities. This study was sup-
ported by a grant (G-5838) to Duke University from the Na-
tional Science Foundation.
Summary
The unusual occurrence of subtidal populations of oysters
in the ocean has been noted on five wrecks on the outer coast
in the region of Cape Hatteras, North Carolina. Comparisons
were made with wrecks where oysters are scarce. The occurrence
of oysters has been correlated with an absence of the oyster drills,
Thais floridana and Urosalpinx cinereus, which are apparently
most important in limiting oysters to intertidal situations in high
salinities in some areas.
References
Burkenroad, M. D. 1931. Ecology 12: 654-665.
Butler, P. A. 1954. Proc. Nat. Shellfish. Assoc. 1953: 67-75.
Carriker, M. R. 1955. U. S. Fish %c Wildl. Serv., Spec. Sci. Rept.
Fish. 148: 1-150.
1957. Jour. Elisha Mitchell Sci. Soc. 7i:328-351.
April, 1960 nautilus 147
Chestnut, A. P. & W. E. Fahy. 1953. Nat. Shellfish. Assoc. Conv.
Addr. 1952: 79-89.
Churchill, E. P. 1920. Rept. Comm. Fish. 1919, App. VIII (Bur.
Fish. Doc. 890): 1-51.
Galtsoff, P. S. & V. L. Loosanoff. 1939. Bull. Bur. Fish. 49: 75-132.
H. F. Prytherch. 1927. Bur. Fish. Econ. Circ. 61: 1-8.
Korringa, P. 1952. Quart. Rev. Biol. 27: 266-308, 339-365. /
Lunz, G. R. 1947. Jour. Elisha Mitchell Sci. Soc. 63: 81. ^
Parr, A. E. 1933. Bull. Bingham Oceanogr. Coll. 4 (3) : 1-90.
Pearse, A. S. & L. G. Williams. 1951. Jour. Elisha Mitchell Sci.
Soc. ^7; 133-16L
Prytherch, H. F. 1934. Ecol. Monogr. 4: 47-107.
Wells, H. W. 1958. Bull. Mar. Sci. Gulf Carib. 5:152-166.
1959. Jour. Elisha Mitchell Sci. Soc. 75 (2) : 168-173.
1. E. Gray, 1960. LimnoL Oceanogr. 5(1). (In press).
NOTES ON THE PRODUCTION OF EGGS IN
ONCOMELANIA NOSOPHORA AND O. FORMOSANA^
By ARIEL A. ROTH and EDWARD D. WAGNER
Department of Microbiology and School of Tropical and Preventive Medicine,
College of Medical Evangelists, Loma Linda, California2
In a previous publication (Roth and Wagner, 1957) the
anatomy of the reproductive system of Oncomelania nosophora
(Robson) has been described. The present investigation was
pursued to determine the changes taking place as the ^g^ de-
velops while passing through the reproductive system of the
female.
For this study, snails which had been reared under laboratory
conditions of near constant temperature and light were used.
The snails were fixed in Bouin's embedded in paraffin, serially
sectioned at 6 to 10 microns and stained usually with Harris'
hematoxylin and tryosin.
The anatomy of the female reproductive system of Oncome-
lania formosana (Pilsbry & Hirase) was studied and found to be
similar to that of O. nosophora. Wagner and Chi (1959) have
shown that these two species will cross with each other produc-
ing fertile offspring. The development of the oocyte appears
1 This investigation was supported by research grant E-644 (C3) from
the National Institute of Allergy and Infectious Diseases of the U. S. Public
Health Service.
2 Present address of senior author: Department of Biology, Emmanue^^ Mis-
sionary College, Berrien Springs, Michigan. ' ^»ir«*; c
148 NAUTILUS Vol. 73 (4)
to follow the same process in both O. nosophora and O. formo-
sana. The following description is based on O. formosana.
The early stages up to the completion of the formation of the
yoke take place in the ovary. The first indication that one of the
cells along the wall of the diverticulated ovary is becoming an
oocyte is an increase in size especially of nucleus along with the
appearance of a large nucleolus. The cytoplasm which is at
first reduced, finely granular and strangely basophylic (Figs. 1
and 2) , becomes acidophylic as the oocyte reaches full size. At
this stage, it is composed mainly of yolk globules of various
sizes tending to be arranged in chains (Figs. 3 and 4) . These
are enclosed in a vitelline membrane and since the oocyte takes
on many varied shapes obviously the contents as well as the
surrounding membrane are very pliable. In many of the medium
and large oocytes, the nucleolus, which in earlier stages appears
single, becomes an amphinucleolus (Fig. 2) with both parts
basophylic; however, one part does not stain as deeply, and, in
Mollory's connective tissue stain, one half is red while the other
is blue.
The following changes usually take place in the parts of the
oviduct which lies between the ovary and the gonopericardial
diverticulum; however in several specimens these changes were
noted in the ovary, while in others these changes were not noted
even in the oocytes found near the gonopericardial diverticulum.
The nucleus, which previously was nearly spherical, now be-
gins to flatten conspicuously. The amphinucleolus also flattens
but maintains its two characteristic parts. A large depression
develops at the outer surface of the oocyte (Figs. 3 and 4) . The
nucleus becomes associated with this depression and usually
lies against it, but is separated from the vitelline membrane by
a thin dense layer of cytoplasm, which apparently is devoid of
yolk (Figs. 3 and 4) . In some instances on the side of the oocyte
opposite from the depression and nucleus can be found a small
mass of cytoplasm, also devoid of yolk (Fig. 3) . This may
correspond to the mass of cytoplasm found at the vegetative
pole of the egg of Lymnaea stagnalis L. as described by Raven
(1945) .
The oviduct proceeds anteriorily beyond the gonopericardial
diverticulum, passes by the opening of the seminal receptacle
April, 1960
NAUTILUS
149
CO
cy^/m
NO
NU
SOJi
All figures at same magnification. Abbreviations used: AN, amphinucleolus:
CO, cytoplasm found on side opposite from nucleus; CY, cytoplasm; NO,
nucleolus; NU, nucleus; SP, sperm; YO, volk.
1, section of young oocyte of O. formosana found along wall of ovary.
2, section of more mature oocyte of O. formosana found along wall of
ovary. 3, section of large oocyte of O. formosatia found in ovary, composite
of two serial sections. 4, section of large oocyte of O. formosana found in
accessory gland (composite of 3 serial sections) .
and bursa copulatrix, and then goes lengthwise through the
elongate, accessory gland opening into the mantle cavity. Eggs
are very rarely found in these anterior portions of the oviduct.
To enhance this possibility, 80 male-female pairs were isolated
in as many petri dishes provided with mud, filter paper, and
maple leaves. The dishes were inspected for the presence of eggs
five to six times a week. We found that the feinales, that lay
any eggs, tend to lay for periods of a few days, which alternate
with longer periods when no eggs are laid. Those females which
appeared to be most active reproductively were examined at
150 NAUTILUS Vol. 73 (4)
frequent intervals throughout the day to try and ascertain when
each was about to lay an egg, and when such was suspected the
temale was immediately placed in the fixative. The shell was
crushed to permit rapid penetration ol the fixative.
The eggs of O. nosophora are laid singly and are enclosed
in a layer of mud, with which entire faecal pellets can be found.
Some of the specimens were fixed when the female appeared to
have selected a site for the building of the egg capsule; others
were fixed after the building of the egg capsule had actually
started. The female used her proboscis to lay the foundation
for the mud capsule. The rest of the process of egg laying was
not observed.
Of the females involved in this study 28 were fixed at a time
when oviposition was suspected. They were serially sectioned,
stained and examined for the presence of eggs. Eggs were fre-
quently found in the posterior part of the oviduct, but in no
instance could they be found in the parts of the reproductive
system anterior to the gonopericardial diverticulum. The mantle
cavity of each of these snails was also examined for the presence
of eggs, but none could be found there. Possibly the last steps
in the production of an egg are rapid ones.
While pursuing another study on the function of other parts
of the reproductive system, an egg was found in the posterior
region of the accessory gland of O. jormosana. The specimen in
which this egg was found had been raised in isolation until 24
hours prior to fixation when it had been mated to a male.
Apparently egg laying already was taking place. The oocyte (Fig.
4) appears to be of the same size and general structure as those
found in the first part of the oviduct. The nuclear contents are
not so distinct. No extra layers are present on the outside of the
oocyte such as are found in eggs that have been laid. Several
sperm are attached to the surface of the oocyte while many more
are in close proximity to it. No sperm appeared to be definitely
inside the oocyte at this stage.
The presence of eggs in the accessory gland of Oncornelania
has not been previously reported, to the writers' knowledge.
Dundee (1957) who studied Po?natiopsis lapidaria (Say) which
belong to the same subfamily, reported that no eggs were found
in the anterior parts of the oviduct in many specimens collected
April, 1960 NAUTILUS 151
from the field. She did state that a fellow worker found an egg
in a specimen, and mentioned a report on it which has not been
published yet.
Since the oocyte found in the accessory gland had a maximum
length of less than 150 microns while Chi and Wagner (1957)
have found the egg, including its mud capsule, had an average
diameter of 1.0 mm., obviously much must be added before the
egg is completed. About a 14 of this addition consists of the mud
capsule. The rest is composed of a clear, spherical, gelatinous
mass in which the small, whitish oocyte is readily detectable. The
origin of this gelatinous mass could not be determined in our
study. Finding an oocyte in the anterior part of the accessory
gland might help elucidate this question. Perhaps fixation of
specimens earlier or later than was done in this study might
produce more positive results; also both Abbott (1946) and
Dundee (I.e.) suggested that the eggs are laid at night. This cycle
might be used as an aid in obtaining eggs in the anterior part of
the oviduct. This was not a factor in our study since the constant
source of light, under which the snails were kept, eliminated the
possibility of such a cycle.
Literature cited
Abbott, R. T. 1946. The egg and breeding habits of Oncomelania
quadrasi Mlldff., the schistosomiasis snail of the Philippines.
Occ. Pap. On Mollusks, Mus. Comp. Zool., Harvard Univ.,
7:41-48.
Chi, L. W. and Wagner, E. D. 1957. Studies on reproduction
and growth of Oncomelania quadrasi, O. nosophora, and O.
formosana, snail hosts of Sehistosoma japonieum. Amer.
Jour. Trop. Med. and Hyg., 6: 949-95.
Dundee, D. S. 1957. Aspects of the biology of Pomatiopsis lapi-
daria (Say) (Mollusca: Gastropoda: Prosobranchia) . Misc.
Pub., Mus. Zool., Univ. Mich., No. 100.
Raven, C. P. 1945. The development of the egg of Lymnaea
stagnalis L. from oviposition till first cleavage. Arch. Neerl.
Zool., 7:91-121.
Roth, A. A. and Wagner, E. D. 1957. The anatomy of the male
and female reproductive systems of Oneomelania nosophora.
Trans. Amer. Microsc. Soc, 76: 52-69.
Wagner, E. D. and Chi, L. W. 1959. Species crossing in Oncome-
lania. Amer. Jour. Trop. Med. and Hyg., 8: 195-198.
152 NAUTILUS Vol. 73 (4)
ECOLOGIC NOTES ON PUERTO RICAN MOLLUSCA
By frank B. GOLLEY
AEG, Savannah River Project,
Department of Zoology, University of Georgia
While on an ecological expedition^ to study the functional
ecology of the montane rain-forest and mangrove forest com-
munities in May, 1959, I was able to make several observations
on molluscan ecology. Although some of these notes may be
repetitive, they are offered here in hope that they may prove a
not unwelcome contribution to the ecology of neotropical mala-
cology. One objective of my study was to determine the biomass
or weight of the herbivores and carnivores in the community.
To do this, a count necessarily was made of all animals in repre-
sentative plots within the forests, at various periods during the
day and night. During these very intensive examinations of the
habitat, mollusks were encountered and the following observa-
tions recorded.
Montane Rain-Forest: Observations were made in two loca-
tions within the montane rain-forest. These were both within
the Luquillo National Forest, on the north-east end of the island
of Puerto Rico. The first study area was on the eastern front of
the mountains above the Sabana Ranger Station, at an altitude of
340 meters, in a relatively undisturbed region of the forest. The
study area was representative in that it included both ravine and
ridge habitats. Vegetation here included the dominant tabunuco
(Dacroydes excelsa) and palms, tree ferns, lianas, and bromeliads
characteristic of tropical forests.
The second study area was located within the mountain massif,
at El Yunque. Here, at the transition between the montane rain-
forest and the sierra palm forest (altitude approximately 700
meters) a moist ravine was investigated. Tabunuco and tree ferns
were present at this location, and the sierra palm (Euterpe
globosa) increased greatly in density. A few additional observa-
1 Supported through a grant from the Rockefeller Foundation to the Insti-
tute of Marine Science, University of Texas. I am indebted to Dr. H. T.
Odum, expedition leader, and Mr. Ronald Wilson, Institute of Marine
Science, for their assistance in making the observations which form the basis
of this paper. I am also grateful to Dr. J. P. E. Morrison for identifying the
mollusks and for his suggestion that I prepare these notes on molluscan
ecology, and to Dr. Grace J. Thomas, University of Ga., for critically reading
the manuscript.
April, 1960 nautilus 153
tions were also made in the mossy forest or elfin woodland,
crowning the mountain peaks, in the Mt. Brittian area.
Rainfall is quite heavy in these regions, averaging at study
area one, 135 inches per year, and increasing as one ascends
the mountains. Temperature is very constant, averaging 23° C.
during the day, and dropping to approximately 19° C. at night.
At the Sabana study area, mollusks were observed in tree trunks
and in the thin leaf litter overlying the soil. The long, thin
Nenia tridens appeared to be the dominant tree snail on the
ridges. Close examination of ten tabunuco trees of different
diameters indicated that the ratio of Nenia to the other abund-
ant snail, Pleurodonte carocolla, was seven to one. In the ravine,
the round Pleurodonte increased in abundance and appeared to
be codominant with Nenia. In the leaf litter, Megalomastoma
(Neopupina) croceum was most abundant. The average number
of snails on the ground was 0.4 per square-meter, and on the
trees, L8 per tree. In this community, snails, when considered
either on the basis of numbers or biomass per square-meter,
are less abundant than are ants or Orthoptera.
On El Yunque, Pleurodonte was decidedly the most abundant
snail and was present on most tree trunks and on the concrete
walls of cabins. As many as a dozen individuals could be col-
lected within 7 feet of the ground on a single palm tree. Over 100
individuals were collected and measured in this locality. The
major diameters of the shells (exclusive of lip of aperture) are
shown below. If shell diameter is a crude approximation of the
Midpoint of diameter class in mm.
Diameters 18 21 24 27 30 33 36 39 42 45 48 51
No. of snails 1 0 3 1 4 4 6 1 6 23 35 17
age of the snail, these data indicate that there may be continual
additions to the population. Diameters of a few of the Pleuro-
donte were also taken at study area one; however, none of these
measured less than 40 mm.
In the mossy forest on Mt. Brittian (approximately 1000 meters
altitude) one square-meter quadrat was carefully inspected for
its animal fauna. Because of exceptionally heavy rainfall, con-
tinual erosion occurs and the soil is washed from between the
tree roots. The result of this erosion produces a thick and very
dense mat of roots on top of the mineral soil and rocks. This
154 NAUTILUS Vol. 73 (4)
root mat is approximately 6 cm. in depth. In addition, the trees
in the mossy forest are covered with a thick growth of mosses
and lichens. The combination of epiphytic covering on the tree
trunks and the root mass on the ground surface appears to pro-
duce an adverse environment for invertebrate life. No snails were
found on the quadrat or on the trees bordering the trail through
the forest. And only 3 small insects were collected in the square-
meter plot.
Mangrove Forest: The mangrove forest was studied at La
Paquerre, the location of the Laboratory of Marine Biology,
University of Puerto Rico. The study area in the forest was
located approximately 20 meters from the open water, where
the vegetation was completely composed of red mangrove,
Rhisophora mangle. At the time of the study, only one-half of
the study area was submerged at high tide. Techniques similar
to those used in the rain-forest were used in the mangrove for-
est. Three one-square-meter quadrats were observed at various
periods during the day and night and all animals seen were
counted. Samples of the thin layer of mud and the mangrove
peat beneath the mud were handsorted for residents. Finally,
tree trunks, prop roots, and leaves were carefully examined for
animals.
The snail, Melampiis cofjeus, was the most numerous animal
present on the surface of the mud. An examination of 9 square-
meter plots revealed that 54 snails were present per square-meter.
Melampus was the second most important animal in the com-
munity on the basis of biomass (.254 grams dry weight/m^) , the
crab, Uca mordax, having the greatest biomass (.425 grams dry
weight/m^) .
The Melampus could be observed during the day, at low tide,
on surfaces of decaying leaves and on the mud. However, during
the night, at high tide, many of these snails had ascended the
young seedlings of Rhisophora or the prop roots of mature
trees. For example, in one night quadrat, 26 individuals of
Melampus were observed on seedlings, 2 on prop roots and 7 on
the ground surface; in another 15 were on seedlings, 22 on
prop roots and 4 on the mud.
No large Melampus were observed in the study area, all speci-
mens being less than 6 mm. in total length. Very small examples
April, 1960 nautilus 155
of Melampus were abundant in every plot examined. Unless the
large adults were present in some portion of the forest not
studied, the size distribution of Melampus would indicate two
generations, with a turn-over in the snail population once every
two years.
NOTEWORTHY RECORDS OF WYOMING MOLLUSCA
By DOROTHY E. BEETLE
Fourteen species in 9 genera have been found during the past
5 years of intensive collecting in Wyoming.
Pupoides hordaceus (Gabb) was collected on the sandstone
cliffs overlooking Guernsey Reservoir, elevation 4400 feet, in
Piatt County. The area is very dry, supporting only scattered
Yellow pine and cedar. Yucca, grasses, and a few herbaceus plants
form a scant ground cover. A single fresh, dead, specimen of P.
hordaceus (D. Beetle 56-289) was found in a crevice in the rocks.
A broken Gastrocopta, probably G. armifera (Say) (D. Beetle
56-298) was sorted out of drift at the edge of the reservoir.
Pupilla syngenes dextroversa (Pils. & Vanatta) was collected
in Hot Springs County, 5 miles east of Thermopolis, on the
road to Lysite. In the leaf mold of cottonwoods and shrubs grow-
ing along a dry stream, 5 specimens (D. Beetle 57-374) were
found. The soil was fine red sand, overlying sandstone. The
elevation is 4400 feet.
Vertigo ovata Say was found by Paul Freytag in Albany County
along the banks of the Little Laramie River near Centennial,
elevation 7800 feet. The snail was found in willow debris, and
brought to the author for identification.
Vallonia excentrica Sterki was discovered at two localities in
Wyoming. At Thermopolis, some individuals (D. Beetle 57-370)
occurred under sedimentary rocks on the hillside opposite the
hospital. This is a south facing slope, dry, supporting cedar,
a few grasses and shrubs. The elevation is 4350 feet. In Sheridan
County this mollusk was found in willow leaves along the
Tongue River at Ranchester, elevation 3775 feet. (D. Beetle
57-430) .
Vallonia costata (Mueller) (D. Beetle 56-276) was collected
at Ayers Natural Bridge near Douglas, Converse County. La
156 NAUTILUS Vol. 73 (4)
Prele Creek flows through a hollow where it has carved out
a natural arch in the sandstone. Cottonwoods, box elder, willows,
grape, and poison ivy grow in dense thickets. V. costata was
living in the leaf mold and grass of the caretaker's lawn. The
elevation is approximately 4850 feet.
Lymnaea (Radix) auricularia (L.) was picked up by Dr. George
Baxter at Alsop Lake, Albany County. This is a shallow, alkali
pond on the Laramie Plains, elevation 7000 feet. The author
has found L. auricularia in two Colorado localities; Woodland
Park in Teller County, and in the lake at City Park at VValsen-
berg, Huerfano County. Where this introduced mollusk has
become established, it is very abundant.
Lymnaea (Galba) doddsi (Baker) was discovered at three lo-
calities along the western slopes of the Big Horn Mountains.
Dry Fork Canyon above the Elkhorn Ranch, near Shell, has a
tiny creek tumbling down it. Near the mouth of the canyon,
at an approximate elevation of 4300 feet, the creek disappears
into a small hole in the rocks. It falls through a cave in which
it has hollowed out a basin in the sedimentary rock. The walls
and floor of the cave are overgrown with moss. The Lymnaea
(D. Beetle 58-219) was around the edge of the basin and in the
moss. It also occurs in Washakie County, in Leigh Canyon, near
the Fish Hatchery at an elevation of 4450 feet. The snails were
crawling on sedimentary rocks at the edge of the small creek
(D. Beetle 58-157), and on grass along a slough farther up the
canyon (D. Beetle 58-147) .
Lymnaea (Galba) dalli Baker was found living in Barber Lake
in Albany County at an elevation of 8700 feet. This is a tiny,
shallow, silt bottomed pond. The animals were crawling on
aquatic vegetation near the shoreline (D. Beetle 50-358) .
It was previously reported (Beetle, D. 1957. Nautilus 71 (1):
12-22.) that Gyraulus articus (Miiller) was found in Leigh Lake,
Teton County. Additional material, identified now by Dr. Dwight
W. Taylor as Promenetus umbilicatellus (Cock.) , has been dis-
covered in Yellowstone Lake as well as in Bighorn, Johnson,
Goshen and Albany Counties. The mollusks are usually crawling
on vegetation in shallow muddy areas. Inclusive elevations range
from 4300 to 9200 feet.
Three species of Pisidium, identified by Rev. H. B. Herring-
April, 1960 nautilus 157
ton, can be included in Wyoming records.
Pisidium compressiim Prime was noted by J. Henderson at
Evanston many years ago. Recently it has been found in Goshen
County in a backwater of the Springer Reservoir, elevation 4250
feet. The clay bottom contained many clams (D. Beetle 56-158) .
Empty shells were found in drift in Guernsey Reservoir, and in
the Laramie River in Albany County. In Laramie County it was
found in two ponds in the south eastern corner of the county.
Pisidium ferrugineum Prime was found in the mud of a beaver
pond on Libbey Creek near Bear Lake in Albany County (D.
Beetle 55-537) . The elevation is approximately 9100 feet.
Pisidium nitidum Jenyns (D. Beetle 56-130) was collected
in a beaver pond on Texas Creek at the Williams Ranch near
Tie Siding, Albany County, elevation 7750 feet. It occurred in
similar situations in Carbon County, both on South Bruiser
Creek near Ryan Park (D. Beetle 56-91), and near Baggs (D.
Beetle 54-404) .
Physa skinneri Taylor, described from the Berends fauna of
the Pleistocene, was collected in Yellowstone Lake by Dr. A. A.
Beetle. This small Physa was living on vegetation and rocks in
shallow water around the shoreline.
The author wishes to acknowledge the assistance of the Wyom-
ing Chapter of the Society of Sigma Xi for two grants-in-aid that
were used in studies of the molluscan fauna of the Big Horn
Mountains.
References
Baker, F. C. 1911. The Lymnaeidae of North and Middle
America. Recent and Fossil. Chicago Acad. Sci. Spec. Publ. 3.
Beetle, D. E. 1954. Nautilus, 67 (4): 121-129.
1957 Ibid., 77 (1): 12-22.
Henderson, J. 1924. Univ. Colo. Studies 13 (2).
1933. Nautilus, 47 (l):l-3.
1936. Supplement Univ. Colo. Studies 23 (2).
Pilsbry, H. A. 1939-1948. Land Mollusca of North America.
North of Mexico. Acad. Nat. Sci. Philadelphia. Monographs
158 NAUTILUS Vol. 73 (4)
HUGH WATSON
1885-1959
With the death of Hugh Watson on 21 January, 1959, Eng-
land lost one of its most meticulous and experienced malacolo-
gists, one whom it will be very difficult to replace for a long time.
Born in Newcastle-on-Tyne on 1 June, 1885, Hugh Watson
was educated privately, and later read the Natural-Sciences
Tripos at Trinity College, Cambridge, where, after a First in
Part I in 1910, his health unfortunately deteriorated. Possessed
of private means, he devoted the rest of his life to the study of
Mollusca, and lived a very retired life in Cambridge.
In his early days, while he was still capable of travel on the
continent, he collected widely both marine and non-marine
Mollusca, but later he concentrated almost entirely on the study
of non-marine gastropods. He was equally interested in both
anatomy and shell form, as is shown by many of his papers.
Among British Mollusca he was especially interested, early on,
in such genera as Vallonia and Vertigo, while later on he be-
came mainly interested in the Zonitidae, especially in the British
species of slugs, although his publications ranged widely. But
his interests extended far beyond this, especially to the study of
certain African genera of Mollusca, notably the Planorbidae.
In fact, his opinion was continually being consulted by malacolo-
gists from all over the world with whom he maintained a volumi-
nous correspondence.
He was equally interested in problems of nomenclature and
a series of papers on the correct names for the British non-marine
Mollusca were published in 1943. Almost all of the names he
suggested have been adopted in the latest British census. At the
time of his death, he had just finished an exhaustive enquiry
into the correct names for the European species of Viviparus.
To a lesser degree, he was concerned with the history of
British species discovered by the study of the Mollusca of Pleisto-
cene deposits and was always hoping for the discovery of new
records both fossil and living, especially in the east of the
country, of species found in adjacent parts of the continent. Un-
fortunately, his ill health prevented him from looking for these
himself, although the care with which he studied doubtful Pleis-
tocene specimens submitted to him was largely instrumental in
April, 1960 nautilus 159
the establishment of several new British records.
Because of his retired life, Hugh Watson was not known per-
sonally to many other malacologists. He worked with extraordi-
narily great care, was fundamentally modest about his own
knowledge, and would only commit himself after an exhaustive
consideration of all literature and data bearing on the subject.
He would not tolerate careless work by anyone, and would go to
great pains to ensure that any work submitted to him for ap-
proval was as thorough and as accurate as his detailed knowledge
could make it. This passion for accuracy made him sometimes
a little unpopular with impatient prospective authors, but it
was completely offset by the kindness which he showed in devot-
ing much of his valuable time to their problems and instruc-
tion.
By his will, Hugh Watson left his specimens and his library
to the University of Cambridge as well as a considerable sum of
money for the furtherance of malacological research.
A list of his publications has been published in Arch. Mol-
luskenk. 88:77 -7S. — B. W. Sparks, University of Cambridge Dept.
Geography, England.
NOTES AND NEWS
James Zetek. 1886-1959 — James Zetek died June 2, 1959, at
his home in Panama City. He married in 1914, and they are
survived by a daughter, Ella Zetek, of Ecuador. Mr. Zetek was
born Dec. 12, 1886, in Chicago, Illinois, and graduated from the
state university in 1911. He went to the Canal Zone in the same
year as entomologist for the zone's Sanitary Department, and
later served similarly the Republic of Panama, which (1915-17)
bestowed on him gold medals and a National Grand Prize. He
also was connected with the Board of Health Laboratory and
the U. S. Department of Agriculture. He helped found the
scientific laboratory of Barro Colorado, in Lake Gatun, and
became its curator in 1923. The island was set aside as a natural
park in 1943, and from 1941 until near the time of his death he
was Resident Manager of the Canal Zone Biological Area. (Cf.
Smithsonian Rept. for 1947: 126-151) .
Although primarily an authority on termites, he also was
160 NAUTILUS Vol. 73 (4)
interested in mollusks and contributed much of the material that
was described by Dr. Pilsbry in 1926, as Scolodonta zeteki com-
memorated. Among Mr. Zetek's own publications on mollusks
are:
1918. The Mollusca of Pratt, Champaign and Vermillion Coun-
ties, Illinois. Trans. 111. Acad. Sci. i7;151-182.
1918. Los moluscos de la Repiiblica de Panama.
1931. (With H. A. Pilsbry) . A Panamic Cyrenoida. Naut. ■^5;69,
pi. 3, fig. 4.
1934. A new Drymaeus from Barro Colorado Island, Panama
Canal Zone. Naut. ^7:93-94, pi. 13, fig. 1. (D. pilsbryi.)
1936. (With R. A. McLean) . Hiata, a new genus of the family
Pholadidae from the Pacific at Panama, with a description
of a new species. Naut. 49: 1 10-1 1 1, pi. 8, figs. 1-4.
— H, B. B. and Stewart H. Jadis.
American Malacological Union. — The 1960 meetings will
be held August 9 to 12, in the Redpath Museum of McGill Uni-
versity, in Montreal, Canada. Rooms will be available in the
Royal Victoria College dormitories. A field trip to Mont St.
Hilaire is planned.
Robert Robertson appointed Assistant Curator of mollusks
at the Academy of Natural Sciences of Philadelphia — Dr. Robert
Robertson, who has recently received his degree from Harvard
University working under Dr. Clench, has been appointed to
the staff of the Academy. His doctorate thesis was on the ecology
of the marine mollusks of Bimini, Bahamas. At the Academy,
he plans among other things to monograph the Indo-Pacific
Phasianellidae and to continue work on Bahamian marine mol-
lusks. He would welcome the loan or gift of specimens of Tricolia
and Phasianella from the Indo-Pacific — R.T.A.
PupoiDES INORNATUS Vauatta was collected in Larimer County,
Colorado, along Route 287 opposite the turnoff to Red Feather
Lakes. A limestone ridge is exposed some two hundred feet
above the highway. It is a dry situation; a few cedar grow on
the rocks, along with poison ivy and grasses. The Pupilla were
found alive in the crevices and on the undersides of loose rock.
(D. Beetle 53-74) . — Dorothy E, Beetle.
Vallonia perspectiva Sterki in Maryland. — On June 28, 1959,
Leslie Hubricht and the writer visited Elliott Island, Dorchester
April, 1960 nautilus iii
County, Maryland, to collect land shells from this isolated sta-
tion. From an Indian shell heap, over 20 species of land shells
were secured, mostly dead, but the most interesting discovery
was the collecting of live juvenile and adult Vallonia perspectiva
Sterki. This small snail has only been previously reported from
the coastal plain by Dr. Pilsbry who collected specimens near
Atlantic City, New Jersey, in August, 1909 and May, 1910 (Naut.
25 ;35) . This station no longer exists (Land Mollusca of North
America, Vol. 2, pt. 2, p. 1033, footnote) . — Ralph W. Jackson,
Route No. 1, Cambridge, Maryland,
Zachrysia provisoria (Pfr.) in Homestead, Florida. — Dr. G.
W. Dekle, entomologist of the State Plant Board of Florida, sent
several specimens of this species which had been collected alive
by P. E. Briggs and J. H. Knowles on November 10, 1959. Appar-
ently this Cuban land snail will shortly become common over
much of southern Florida where suitable situations occur. It
probably is being distributed on plants, either as eggs in the
soil about their roots or as young snails on the plants. As far as
I know, no species in Zachrysia is considered a pest in Cuba.
They feed mainly at night, probably on dead vegetation. (See also
Naut. 7i;76, 1959) — W. J. Clench.
PUBLICATIONS RECEIVED
Two NEW SPECIES of wcst North American marine gastropods.
By Rudolf Stohler. Proc. Calif. Acad. Sci. 29 (11): 423-444, 10 figs.
1959. — Astraea (Uvanilla) rupicollina and Macrarene coronaden-
sis are described and exquisitely figured as new from Lower Cali-
fornia.—H. B. B.
Pleistocene molluscan faunas of the Humboldt deposit,
Ross County, Ohio. By Martin B. Reynolds. Ohio J. of Sci.
5^:152-166. 6 figs. 1959. — Mainly on stratigraphic grounds, this
fresh-water deposit is considered to be of "early" Wisconsin age.
— H. B. B.
Land and freshwater mollusks of Great and Little Inagua,
Bahama Islands. By William J. Clench. Bull. Mus. Comp. Zool.
Harvard 72i:29-53. 1 pL, 1 map. 1959.— Eutrochatella klinei,
Drymaeus bahamensis, subsp. salina (nounl), and Cerion (May-
nardia) ericksoni are new. — H. B. B.
INDO-PACIFIC MOLLUSCA
Monographs of the Marine Mollusks of the
Tropical Western Pacific and Indian Oceans
A new series devoted to taxonomic revisions o£ the marine
mollusks found from East Africa to Polynesia and Japan to
Australia. All species are illustrated either in full color, black-
and-white photographs or line drawings. With descriptions,
synonymies, anatomy, recent and Tertiary records, habitats, etc.
Editor: R. Tucker Abbott; co-editors: William J. Clench and
Harold A. Rehder. Vol. 1, no. 1 (Introduction and Vasidae) now
available. Vol, 1, no. 2 (Stromhidae, with 6 color plates) in press.
Issued in pamphlet or loose leaf form. State preference. Attrac-
tive, gold-stamped, post binder protects your copies, saves you
cost of binding. $4.25 (foreign: add 50 cents) . Subscription: $5.00
per 100 pages. Not sent as a library exchange.
Department of Mollusks,
Academy of Natural Sciences of Phila., Philadelphia 3, Pa., U.S.A.
WILLIAM H. WEEKS SHELL COLLECTION: New lists now
in preparation; full scientific data. Send name and address
for free copies. Shells also wanted for purchase and exchange.
George E. Jacobs, 853 Riverside Drive, New York, 32, N. Y.
A CHECK LIST OF THE MARINE SHELLS OF ST. CROIX,
U. S. Virgin Islands, with random annotations. Up to date.
Lists 650 species plus 60 subspecies, including 20 new. With
70 illustrations. The "random notes" contain much new infor-
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Nearly 300 pages, about 2,000 figures, some in color. First
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