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VOL. 75 
JULY, 1961 to APRIL, 1962 


Professor Emeritus of Zoology, University of Pennsylvania 

Consulting Biologists Inc., Bethlehem Pike, Spring House, Pa. 

H. A. Pilsbry Chair of Malacology, Academy of Natural Sciences 


Philadelphia, Pennsylvania 


April, 1962 nautilus iii 


Names of new genera, species, etc. in italics 

Acteocina candei distinct from Retusa canaliculata 87 

Adventitious dispersal 94, 124 

Alabama 32, 63, 97, 104, 106, 123, 124, 145 

American Malacological Union 83 

Ancylidae, radulae 97, 145 

Anomia aculeata, variation and sculpture 131 

Arcidens confragosus in Kansas 95 

Arkansas 28 

Atlantic, eastern 50, 109 

western 1, 7, 21, 40, 50, 55, 85, 87, 94, 108, 

109, 127, 128, 131, 138, 149 

Australorbis albicans, anatomy and shell 156 

Beckianum H. B. Baker (genus of Achatinidae or subgenus 

of Leptinaria 84 

Brachidontes recurvus, gonad development and spawning 149 

Bulimulus dealbatus jonesi 166 

California, inland 84 

marines 19 

Canada 103 

Capulus sericeus Burch & Burch 19 

Camaenidae, Puerto Rican 64 

Caracolus marginella mayaguezi H. B. Baker 64 

Catinella pugilator Hubricht 31, pi. 4, 61 

Catinella texana Hubricht 31, pi. 4, 61 

Cerion, anatomy 33 

Cionella lubrica, aggregations Ill 

Clausilia, pallial complex 36 

Corbicula fluminea from Ohio River 126 

Cuba 156 

Cymatium caribbaeum, range extension 94 

Dates of the Nautilus 39 

Directory of conchologists 40 

Engina zonata Gray, type species 107 

Eupleura caudata, sexual behavior 7 

Europse 36, 39 

European Malacological Congress 39 


iv NAUTILUS Vol. 75 (Index) 

Fluorescence, enhancement by glycerin 138 

Florida, inland 63, 142 

marine 94 

Gaeotis nigrolineata 121 

Gardner, Julia Anna, obituary 122 

"Gastrondonta" saludensis to Helicodiscus 105 

Georgia 63, 124 

Geotaxis in Physa 75 

Glyphyalinia junalaskana, anatomy 125 

Gonads and spawning 149 

Haiti 71 

Helicodiscus barri Hubricht 105 

Helicodiscus hadenoecus Hubricht 106 

Helicodiscus jacksoni Hubricht 106 

Helicodiscus multidejis Hubricht 102 

Helicodiscus notius & H. n. specus Hubricht 104, 105 

Helicodiscus saludensis 105 

Helicodiscus shimeki Hubricht 103 

Helicodiscus, subgenus Troglodiscus 105 

Helisoma anceps carried by bug 124 

Holopodopes, infraorder o£ Geophila 116 

Illinois 113, 126 

Ilyanassa obsoleta, mass movement 85 

Indiana 123 

Iowa 103 

Jamaica 142 

Kansas 46, 95 

Kentucky 105, 106 

Lacinaria, pallial complex 36 

Leptinaria, subgenus (?) Beckianum 84 

Louisiana 79 

Maryland 62, 107 

Mexico 31, 84, 142 

Michigan 124 

Mississippi 63 

Missouri 104 

Nenia tridens, anatomy 35 

Neoplanorbinae, radulae 145 

New Guinea 70 

April, 1962 nautilus v 

New Mexico 28 

New York 103 

North Carolina 62, 124, 126 

Notes and news 39, 84, 123 

Nucella 109 

Obeliscus (Stenogyra) terebraster rarisinister H. B. Baker 117 

Oklahoma 46, 104, 124 

Oleacinoids, Puerto Rican 142 

Oliva rejecta Burch & Burch 165 

Oliva spicata & O. venulata 162, 165 

Olivella mutica, reproduction 139 

Pacific, eastern 19, 162, 165 

Papuina ferussaci 67 

"Paravitrea" roundyi to Helicodiscus (?) 107 

Physa, geotactic behavior 75 

"Pilsbryna" tridens to Helicodiscus (?) 107 

Pleistocene 43 

Polydontes incerta & P. lima asperula 66 

Polygyra gracilis Hubricht 26 

Polygyra lithica Hubricht 28 

Pomacea paludosa in Alabama 123 

Practicolella berlandieri campi 29 

Proptera capax in Kansas 95 

Publications received 41, 86, (3) iii, (4) iii 

Puerto Rico 33, 64, 116, 142, 156 

Pupillidae 33 

Radulae of ancylids 97, 145 

Retusa canaliculata distinct from Acteocina candei 87 

Rhodacmeinae, radulae 97 

Rissoellidae 21 

Sagdidae 142 

Schwengel, Jeanne Sanderson, obituary 36 

Sexual behavior, Eupleura and Urosalpinx 7 

Smith, Maxwell, death notice and portrait 84, (4) pi. 18 

South America 16 

South Carolina 63 

Spiraxidae 142 

Stenotrema calvescens Hubricht 28 

Strobilops lonsdalei & S. I. cansasiana Ho &: Leonard 43, 46 

vi NAUTILUS Vol. 75 (Index) 

Strombus canaliculatus 127, 129 

Succinea Indiana 60, 123 

Succinea solastra Hubricht 30 

Succinea urbana Hubricht 32 

Tennessee 29, 102, 105, 106 

Thyasira sarsi, synonymy 50 

Texas 27, 30, 43, 61 

Troglodiscus, subgenus of Helicodiscus 105 

Urosalpinx cinerea, sexual behavior 7 

Valvata windhauseni Parodiz 16 

Vasum globulus nuttingi 1 

Vertigo teskeyae 62 

Virgin Island marines 55, 108 

Virginia 104 

Viviparus subpurpureus in Oklahoma 124 

X-ray diffraction & shell structure in Oliva 162 

Xylophaga atlantica 40 


Allen, J. Frances 149 

Baker, H. Burrington 33, 39, 64, 84, 116, 142 

Basch, Paul F 97, 145 

Bode, William T. (Shuster &) 1 

Branson, Branley A. 124 

Briggs, Rev. H. E. J 39 

Burch, John Q 40 

Burch & Rose L. Burch 19, 165 

Deslandes, Newton (Paraense &) 156 

Dexter, Ralph W 40, 85 

Donohue, Jerry & Kenneth Hardcastle 162 

Dundee, Dee Saunders & Patti Watt 79 

Editors 36 

Eyerdam, Walter Jacob 71 

Fechtner, Frederick R 126 

Gregg, Wendell 84 

Hardcastle, Kenneth (Donohue &) 162 

Hargis, William J., Jr. & Clyde L. Mackensie, Jr 7 

Henrard, J. B 67 

April, 1962 nautilus vii 

Ho, Tong-Yun & A. B. Leonard 43 

Hubricht, Leslie 26, 60, 102, 123, 123, 125, 166 

Leonard, A. B. (Ho &) 43 

Mackensie, Clyde L., Jr. (Hargis &) 7 

McClary, Andrew 75 

Merrill, Arthur S 94, 1 3 1 

Moore, D. R. (Olsson &) 127 

Murray, Harold D 95 

Ockelmann, K. W 50 

Olsson, A. A. & D. R. Moore 127 

Orr, Virginia 107 

Owen, D. F 1 24 

Paine, Robert T 1 39 

Paraense, W. Lobato 8c Newton Deslandes 156 

Parodiz, J. J. 16 

Rehder, Harald A 1 09 

Robertson, Robert 2L 128 

Roscoe, Ernest J. 1 1 1 

Shuster, Carl N., Jr. & William T. Bode 1 

Teskey, Margaret C 83 

Watt, Patti (Dundee &) 79 

Weber, J. A 55 

Wells, Harry J. & Mary Jane Wells 87 

Wilson, Druid 122 

Woodridge, Richard G., Ill 138 


Vol. 75 July, 1961 No. 1 



Department of Biological Sciences, University of Delaware^ and 

El Camino High School, Sacramento, California 

The information in this article is based upon the observations 
by Mr. John B. Henderson, Jr. (Nutting, 1919), and from field 
notes made in connection with the Smithsonian-Bredin Caribbean 
Expedition, 1958, and by Mr. Bode during two years, 1957-1958, as 
a Fulbright Fellow at Antigua.^ 

Information on the distribution, ecology, evolution, and sys- 
tematics of the genus Vasum in the western Atlantic is of particu- 
lar interest to the writers of this article. Since at least three species, 
V. jnuricatum, capitelliim, and V. globulus (Globivasum) nuttingi 
(Henderson), have overlapping ranges in the Lesser Antilles, spe- 
cial care should be made to report the exact locality and observa- 
tions of each find. 

Habitat notes. Vasum muricatum and V. globulus nuttingi were 
found within a few miles of each other at Barbuda, but in dis- 
tinctly different habitats. Vasum muricatum is generally found in 
or close to turtle grass beds. 
Vasum muricatum (Born) 

Anegada: from Pomato Point in shoal water on Turtle Grass 
flat. Barbuda: south shore (Gavenor's Landing) . 5 specimens 
from shallow water, 3 to 6 feet depths, on sandy bottom in region 
of coral heads and nearby shoreward expanse of turtle grass. Ab- 
bott (1950) noted that "It seems to prefer rather sheltered, shal- 
low waters, but is sometimes found on relatively exposed reefs." 
Antigua: Jumby Bay, Long Island. 
Vasum globulus nuttingi (Henderson) 

Barbuda: windward side of Spanish Point. Specimens found in 
shallow water along coral (dead) rock ledges and on patches of 

1 University of Delaware Marine Laboratories, Contribution No. 7. 

2 For the opportunities afforded him, as an invited member of the expedi- 
tion, the senior author wishes to express his gratitude to the sponsors of the 
expedition, Mr. and Mrs. J. Bruce Bredin and to the expedition leader, 
Dr. Waldo L. Schmitt. 


2 NAUTILUS Vol. 75 (1) 

sand among the rocks. Roughly 40% of the specimens were within 
a few inches of Diadema. Several Vasum were only an inch away 
from the periphery of the Diadema. Depths were 1 to 3 feet. In 
only one case, a specimen was collected in another habitat: turtle 
grass on a sand-covered ledge of coral rock, depth 6 inches. 
Antigua: from areas just inside points of land, in semi-protected 
waters. As at Barbuda, where an offshore coral reef broke the 
direct force of the surf, the localities in Antigua were areas with 
well-circulated seawater but not exposed to direct surf. We found 
no Nutting's vase well within bays. Localities included are: Car- 
lisle Bay: in 3 feet of water. Dead Sands: rocky area on the east 
side, in 5 feet of water. Ding-a-Dong Nook: in 4 to 5 feet of water. 
Exchange Bay and Smith Island at Mill Reef: in one to 21/2 feet. 
Freeman's Bay: inside harbor mouth. Galley Bay: in 3 to 6 feet 
of water. Literally hundreds of Nutting's Vase shells containing 
hermit crabs were observed on the Galley Bay side of the point 
separating Galley Bay and Little Galley Bay. Specimens can be 
collected at Little Galley Bay when the sea is calm. Guana Island: 
at the part of the island called the Headlands, at 3 to 4 foot 
depths. Half Moon Bay: inside rocky area on the Mill Reef 
side of the bay and in 2 to 6 feet of water, along western shore- 
line, midway between the sand beach and the open sea to the 
south. Johnson's Island: northwest side, in 4 to 5 foot depths. 
Rendezvous Bay: in 4 feet of water; this area may be heavily 
populated, but extremely rough surge makes it difficult to collect; 
it was the only locality on Antigua where specimens were found 
directly exposed to the open sea. Windward Bay: in tide pools 
formed by a small reef of dead coral, at the water's edge to one 
and a half feet; in 3 to 6 foot depths outside of the tide pools. 

Natural history of species of Vasum: Little is known about 
the living animal. Vasum muricatum is known to be predatory 
upon worms and clams (Abbott, 1954) ; presumably, the other 
species are also predatory. 

In its habitat, Vasum globulus nuttingi escapes notice from 
all but the experienced collector. Nutting (1919) reported from 
the notes of Mr. Henderson that, "Miss Sykes seemed to be the 
only collector with eyes properly focussed to detect these turbin- 
ellas clinging to the rocks, so well concealed are they by their 
coloration and by the calcareous deposit that further hides them 
from their enemy (but not Miss Sykes). One of the chief enemies 
of these rock-living mollusks is a fish of the grouper family that at 
high tide swims about the reefs examining every inch of their 
surface for little mollusks that have not hidden themselves very 
carefully." Miss Esther Bates, in a mimeographed newsletter, 

July, 1961 NAUTILUS 3 

commented upon collecting at Carlisle Bay, Antigua, on 22 
April, 1958: "The tide was low and they [natives] led us to a 
coral reef at the water's edge. They [Nutting's vase] are difficult 
to see against the rocky background but I came away with a 
goodly numbei" ..." 

Vasiim muricatum may be nocturnal, hiding during the bright 
sunlight hours among a little species of coral and turtle grass; 
traveling across the bottom, in 2 to 3 feet of water, in the late 
afternoon. This may also be the case for V. globulus nuttingi. 
Most specimens appear to be, to the collector swimming about in 
search of them, at rest among growths on rocks or partially cov- 
ered with sand. Possibly visual, audible, or pressure stimuli, 
caused by the swimmer, cause the snails to "freeze." Specimens 
removed from the water frequently come part way out of their 
shells and appear to be quite active. 

Three of the species have been dredged from moderate depths 
{Abbott, 1950; Rehder and Abbott, 1951) : V. latiriforme (18 to 
20 fathoms) , capitellum (10 fathoms), and globulus nuttingi (7 
fathoms) . The type locality of globulus nuttingi was designated 
as 7 fathoms (Abbott, 1950) , whereas the published account of 
the living animals (Nutting, 1919) reported them on the rocks 
in shallow water about the reefs at the entrance to Freeman's 
Bay. There is no doubt that shallow water areas are characteristic 
habitats; it remains to be proved that moderate depths are also 
usual or common. If a range in depth of habitat exists, then a 
seasonal migration pattern, perhaps for feeding or breeding, may 
be involved. In this respect, the collections by Bode and Shuster 
were made only in shallow water, during the period from 9 
March through 25 June, 1958. A false distribution may be caused 
by the activities of predators, such as the grouper (Nutting, 
1919) , or by hermit crabs. 

V. globulus nuttingi has been found at depths ranging from 
the water's edge to, as a rule, not more than 6 or 8 feet of water. 
In every case, except at Rendezvous Bay, the snails were found in 
an area protected from the direct waves of the open sea, yet they 
were always located where there was considerable water circula- 
tion. As a rule, they were not found in sheltered areas, as at the 
head of a bay. The most sheltered habitat observed by Mr. Bode 
was just inside the entrance to Freeman's Bay, where the snails 

4 NAUTILUS Vol. 75 (1) 

were found in shallow water, not exceeding a foot in depth. Their 
distribution in this particular area, as in several others, was 
restricted to a zone covered by a moss-like giowth on the rocks. 
This growth was not collected nor identified, although it can 
be recognized easily again once a person has seen it. It appears 
like a coarse moss, 14 inch to I1/2 inches long, ranging in color 
from a grey to a brown. 

In some areas the Nutting's vase was found in the shallow water 
within a short distance of the beach, among rocks and dead coral, 
with a higher ridge of rock and dead coral between the open 
sea and their habitat. Often a bank of live coral would be located 
at the edge of this "protecting" reef and the water depths quickly 
dropped off to at least 25 or 30 feet. Although examples of V. 
globulus nuttingi were found in shallow water, they were within 
a distance generally not exceeding 50 to a 100 feet from deep 

Specimens of Nutting's vase are frequently found partially 
buried in pockets of coral sand in crevasses of dead coral. When 
collected out in the open, on the top of rocks, they are always 
beneath the water's surface. No examples were ever found on 
living coral, although some, as at Half Moon Bay, were almost 
completely encrusted with what appeared to be a living coral. 
This coral had a very light lavender color when first taken 
from the water. It completely covered and hid the shape of the 
shell, with the exception of the aperture. 

Large numbers of immature V. globulus nuttingi have not yet 
been found. In some areas only adult specimens occuned. In 
others, only immature examples were found and when one or 
two snails were found in an area they were most often immature. 
On the basis of these observations, since in most areas the adult 
specimens considerably outnumber the immature ones, we believe 
that either the young stages are to be found in another habitat, 
perhaps a breeding area, or that the adult populations are main- 
tained by a low number of surviving immature snails. Mr. Bode 
has noted that the local populations can be over-fished. This 
observation suggests that the snails either have a wider range 
of distribution, as into greater depths, or more probably, there 
is a low level of population replenishment due to predation and 
other forms of mortality or to a low reproductive rate. 

July, 196 1 


ligmcs a lu d, J'a.suin {(.'Aabivaauin) globulus nutliugi (Henderson), lo 
illnstrate variation in adult shells: a, the "fossilized" shell frim a kitchen 
midden, b, characteristic shell, c. an elongate specimen, d, the largest shell. 
Scale block 5 cm. in length. 

An area that needs considerably more investigation comprises 
the small islands off the northeast coast of Antigua, and in par- 
ticular, the windward side of Long Island and Guana Island. 
Exchange Island, Little Exchange, Hellgate, Pelican Island and 
Maid Island should also be more closely investigated. In this 
general area, dead shells have been picked up that appear to be 
larger and longer than those found on the south side of the island. 

Among the several dozens of V. globulus nuttingi collected by 
Mr. Bode, 3 are worthy of special mention: 2 are atypical, one 
was found in a kitchen midden. These 3 specimens (figs a, c, d) 
and a more usual one (fig. b) from Barbuda (Shuster!) are de- 
picted on page 5. An elongate specimen (fig. c) was collected 
alive at Windward Bay, Antigua, on 9 March, 1958. It was found 
at a depth of about 3 feet of water in an area consisting of broken, 
dead coral and coral rock, with small patches of sand. A very 
large shell was found in the same area. The third shell (fig. a) 
was found in a kitchen midden, approximately two feet below 
the surface, in conjunction with potshard and broken conch and 
nuuex shells. The midden site was at the western end of a "shell 
beach" on tlie north side of Antigua, between the airjjort and 






EupU'ura caudaiu (Say) . Fig. 1 , male and female in Fig. 2. two males 
attempting copulation with one female. 

July, 1961 NAUTILUS 7 

Long Island. This shell was in a bank approximately 7 feet above 
mean tide and 10 feet back from the water's edge. Two of these 
specimens were pictured at slightly larger than actual size by 
Shuster (1959) : the example with tlie elevated spire and the low- 
spired shell from the midden. 

Abbott, R. T. 1950. The genera Xancus and Vasurn in the 

western Atlantic. Johnsonia, 2 (28) : 20 1-2 18. 

1954. American Seashells. D. van Nostrand, Princeton, N. J. 

Nutting, C. C. 1919. Barbados-Antigua expedition. University of 

Iowa Studies in Natural History, 8(3) : 199-203. 
Rehder, H. A. and R. T. Abbott. 1951. Some new and interesting 

mollusks from the deeper waters of the Gulf of Mexico. Rev. 

Soc. Malacologica, 8 (2) -.5 3-66. 
Shuster, C. N. Jr. 1958. Caribbean adventure. Estuarine Bulletin, 




Virginia Fisheries Laboratoi7 

The observations reported herein are part of an extensive study 
of the biology of Urosalpinx cinerea (Say) , the smooth oyster 
drill, and Eupleura caudata (Say) , the rough oyster drill, which 
is in progress at our laboratory. Results of studies of other aspects 
of the behavior and ecology of these predatory snails will be re- 
ported as they become available. 

Little is known of the copulatory behavior of dioecious marine 
gastropods. Though Stauber (1943) reported a partial pairing of 
E. caudata, pairing of U. cinerea has never been described. Our 
studies show that pairing of both species is a complex process 
involving fairly intricate behavior of both sexes. 

Copulation o^ Eupleura caudata. According to Stauber's (1943) 
description of a partial mating of E. caudata the male mounted 
the right side of the female and formed the anterior part of his 
foot into a copulatory groove through which the penis was ex- 

1 Contributions from the Virginia Fisheries Laboratory, No. 00. This re- 
search was conducted under contract with the U. S. Fish and Wildlife Service, 
Bureau of Commercial Fisheries, No. 14-19-008-2372, Study of Oyster Drills in 
Chesapeake Bay. 

- Present address. Bureau of Commercial Fisheries Biological Laboratory, 
Milford, Connecticut. 

8 NAUTILUS Vol. 75 (1) 

tended into her mantle cavity. Even though disturbed repeatedly 
this pair copulated intermittently for 21 days. Our laboratory 
observations of pairing of captive E. caudata confirm and enlarge 
upon those of Stauber. Eighty-one separate pairings were observed 
but because the average time each couple was together was ex- 
tensive and coitus occurred intermittently, only two copulations 
were seen from beginning to end. 

Prior to coupling, the female usually moved about the aquar- 
ium walls and finally assumed a stationary position with her 
siphonal tip upward. Pairing also occurred in horizontal positions 
on oysters, or on the aquarium bottom, or rarely, on the walls. 
The mantle cavity was then exposed by allowing the shell to 
hang down and away from the posterior part of the foot. Though 
in most cases a male (or males) had already assumed a position 
upon the shell of the female, several unpaired females also ex- 
posed their mantle cavities in the same fashion prior to copulating 
and probably this behavior is normal precopulatory activity. Fol- 
lowing this the male often moved about on the female's shell for 
a short time but eventually assumed a stationary position on her 
right ventral surface slightly posterior to the mantle cavity with 
his siphonal cavity pointing in the same direction as hers (Fig. 1, 
p. 6) . The foot of the male extended to the rim of the mantle 
cavity and a copulatory groove formed in its anterior surface 
through which the penis was protruded into the mantle cavity 
and presumably the vagina, though that orifice was always hidden 
from view by the shell. On termination of copulation, the female 
often twisted from side to side and opened and closed the cavity 
opening. Similar twisting behavior has been observed in other 
gastropods apparently attempting to dislodge predatory snails, 
Odostomia, from their shells (Allen, 1958) . The male then with- 
drew and either remained in position or moved about on the 
shell of the female or to the substrate. 

E. caudata usually remained in position and copulated inter- 
mittently for extensive periods, often up to 48 hours. Two pairs 
copulated continuously for 2 hours 45 minutes ± 15 minutes and 
3 hour 39 minutes ± 10 minutes, and a third for 5 hours ± 10 
minutes, but all were paired longer. Another pair copulated 
intermittently for a total of at least 8 hoins 48 minutes over a 
period of two days, during which the male maintained his posi- 

July, 1961 NAUTILUS 9 

tion on the female. A marked pair copulated 1 1 times in five 
months. During this period the pair often separated and the 
female paired with other males. 

Eupleura caudata showed marked promiscuity. Of 29 marked 
pairs, 12 females copulated with more than one male and 15 males 
with more than one female. One male copulated with 6 different 
females and one female accepted 5 males throughout the season. 
Although these are observations on drills confined in running- 
water aquaria and cages, there is little reason to doubt that, 
depending on density and movement, similar promiscuity occurs 
in nature. 

Usually, males were smaller than their consorts (48 males: 
mean height 18.7 mm., range 14.9-23.5 mm., standard deviation 
2.3; 44 females: mean height 22.6 mm., range 17.5-28.9 mm., 
standard deviation 3.2) . 

Seasonal periodicity of copulation was determined in outside 
cage experiments and running-water aquarium observations. 
Though the frequency of observation was not always constant 
throughout the year, the number of pairings seen in aquaria per 
month roughly corresponded to those in more careful cage exper- 
iments. In 1956 pairings were recorded as follows: March-6, 
April-14, May-15, June-3, July-9, August-1, September-1, October- 
9, November- 1, and December- 1. The first occurred on March 7 

(12.9°) and the first peak occurred in late April and early May. 
A second peak came in July, comparatively few pairings occurred 
from July through September, but an increase took place in the 
last part of October. Copulation ceased after December 10 

(10.3°C) . The lowest temperature at which pairings were ob- 
served was 10.3°C, the highest 28.4°C. Also in 1956, 30 females 
and 30 males, caged in a single large compartment, were examined 
every two days between 1400 and 1700 hours. Onset of the mating 
period was not observed because the experiment was established 
too late in the season. Pairing was first seen in late April at 13.7°C, 
reached a peak on June 7 at 23.7°C, and ceased on July 7 at 
26.1 °C, (Figure 3). A late wave of 28 pairings, about one-third as 
intense as the spring wave, began at the end of September 

(21.7°C) and ended November 11 (16.4°C.) Observations made 
two or three times a week during the winter of 1956-57 revealed 
no copulatory activity but this was expected because all other 



Vol. 75 (1) 



1 1 1 1 1 1 1 1 1 


lOvAuHro 9 1IH01a 

t.joo to on 

:.<I03 JO ON 

overt activity had ceased. Observations in the first half of the 
summer of 1957 on two cage compartments containing 45 speci- 
mens each (sex ratio of entire sample, 50 females to 40 males) 
indicated that copulation was less intense than in 1956. Pairing 
was first observed on April 4 (1 1.1 °C) . There was no distinct peak 

July, 1961 NAUTILUS 11 

and after the week of May 6 activity slowed and ceased on July 
6 (26.1°C). Most activity occurred slightly earlier than in 1956, 
probably as a result of earlier warming. No observations were 
made in late summer. 

Little diurnal periodicity was noticed and nocturnal activity 
was not investigated. Of 56 pairings, 23 (48 per cent) were com- 
pleted before 1200 hours and 29 (52 per cent) occurred after 1200 
hours. Because observational effort varied somewhat these data 
ajre not precise, though they are probably indicative of general 
conditions and comparable to those pertaining to U. cinerea 
gathered in similar fashion, see below. 

On two occasions, pairs of males were observed in copula. 
Careful external examination and studies of gonad smears showed 
all four animals to be normal males with no detectable female 
characteristics. Five trios in copula, each composed of a female 
and a male in the usual position with an extra male in copulatory 
position on the first male, were also observed in aquaria. The 
extended penes of both males were seen twice. Usually the male 
next to the female copulated with her while the intromittent 
organ of the second male extended into the mantle cavity of the 
first, but at times both males attempted copulation with the 
female (Fig. 2, p. 6) . Copulations involving two functional males, 
the middle one acting as both male and female, and another 
female has been reported for the hermaphroditic species, 
Lymnea stagnalis, (Crabbe, 1927) but never for dioecious gas- 
tropods. Once a quartet with 3 males, all situated chain-fashion 
on a female, was observed. The penes of at least two of these 
males were extended into the mantle cavity of the animal before 
them. In dioecious animals, like drills, these abeirrant copulatory 
groups have little reproductive significance, but seem to support 
the hypothesis of exocrine stimulation or attraction of males to 
females. Possibly the males were attracted to the females, or to 
each other, by release of female exocrine in the vicinity and, 
without being able to discriminate further, established and main- 
tained contact with each other. 

Copulation of Urosalpinx cinerea. Observations of 76 pairs of 
U. cinerea showed their copulatory behavior to be generally the 
same as that of E. caudata, but with several distinct differences. 
The female usually assumed a stationary position with her 

12 NAUTILUS Vol. 75 (1) 

siphonal tip upwards, allowed the shell to fall away from the 
substrate, thus exposing the right rear corner of the mantle 
cavity and twisted the shell from side to side several times in a 
sort of "precopulatory dance." Most often a male was already on 
the female's back when this precopulatory play was observed, but 
several times males appeared to be attracted from afar to the 
demonstrating female. Sometimes unattached males, often several 
at a time, were attached to a demonstrating female or copulating 
pair, possibly drawn to the receptive female or the pair by some 
stimulus, probably an exocrine. Following the precopulatory dem- 
onstration by the female the male assumed a position on her right 
ventral side, formed a copulatory groove of the anterior portion 
of his foot and inserted his penis through the groove into the 
female's mantle cavity. On completion of coitus, the intromittent 
organ was withdrawn, and in contrast to the slow separation or 
intermittent resumption of E. candata, the male moved quickly 
away. The shortest complete copulation observed lasted 4 min- 
utes, the longest 32 min. Including preplay time, one complete 
pairing occupied one hour and 25 minutes, but actual copulatory 
contact required only 1 1 minutes. Usually U. cinerea copulation 
occupied only 3 to 4 minutes. This characteristic short contact, 
seldom more than 4 or 5 minutes, probably explains why U. 
cinerea mating has not been previously reported. 

According to our observations, only one pair of 20 marked 
pairs recopulated and the pairings were U/o months apart. Of 
the marked pairs 7 males and 2 females coupled with different 
mates. Thus, E. caudata seemed more promiscuous than U. 
cinerea. However, the incidence of promiscuity in U. cinerea 
may be actually greater than these data indicate because of the 
unlikelihood of observing its brief sexual contacts as readily as 
the much longer ones of E. caudata. 

The male is usually the smaller of the pair: 41 pairing Seaside 
(from the ocean coast of the Eastern Shore of Virginia) males 
averaged 29.8 mm. in height, range 19.5-38.7, standard deviation 
5.1; 46 females averaged 34.4 mm. in height, range 21.9-44.6 mm., 
standard deviation 4.7; six York River males averaged 19.3 mm. 
in height, range 16.5-24.6 mm. and five females averaged 19.5 mm. 
(17.3-21.2 mm.). 

Pairing was seen in late October and early November, 1955, 

July, 1961 NAUTILUS 13 

until the temperature dropped to 13.7°C. In 1956, it began on 
Februai-y 29 (8.2°C., increased in March with 10 pairings, reached 
a peak in April with 17 pairings, and diminished in May to three 
pairings (20,3°C) . Pairing was not observed again until Oc- 
tober, (20.8°C), none in November and 2 on December 7 
(11.5°C). Of 59 pairings, nine (15 per cent) were observed 
before 1200 hours and 50 (85 per cent) after 1200 hours. In 
contrast to E. caudata, U. cinerea seems to exhibit marked 
preference for the afternoon. 

Two pairs of structurally normal males were observed in copula 
and, on another occasion, two males were observed copulating 
with one female. 

Sperm viability and storage. One U. cinerea male, forcibly 
separated from a female, exuded sperm in a continuous, viscous 
stream, thus indicating that discrete spermatophores are prob- 
ably not employed. Examination of seminal receptacles of at 
least 50 females of both species support this conclusion. Seminal 
receptacle smears of two females taken immediately after copula- 
tion contained immotile sperm while those in another were 
motile; therefore, it is not clear whether the sperm are immotile 
when passed. Microscopic examinations of seminal receptacles of 
several hundied individuals of both species revealed that many 
mature females contained fully or partially motile sperm at all 
times of the year though the sperm in some were entirely immo- 
tile. Whether they are stored in the motile condition is not clear 
because the mechanical action of smearing may have stimulated 
the sperm to activity. Apparently spermatozoal energy is sustained 
in some way, either through conservation of energy by immotility 
or special nourishment because both species can store viable sperm 
for considerable periods of time. 

Stauber (1943) reported that a U. cinerea female isolated from 
April to October of the same year deposited e^^ cases containing 
viable embryos through the period. In our experiments 4 females 
isolated in November and December of 1955 produced viable 
eggs the following spring and summer. Of these, one deposited in 
May after 5 months, two spawned in August, after somewhat less 
than 9 months, and one in September after 9 months. Two post 
copula E. caudata females isolated in 1955 produced egg cases 
through 1956 and into 1957, but the embryos produced in 1957 

14 NAUTILUS Vol. 75 (1) 

did not develop. Unless parthenogenesis occurred — which is most 
unlikely — or the so-called "embryos" included in these 1957 egg 
cases were merely unfertilized ova and not really embryos at all, 
the sperm must have been vigorous enough to affect fertilization 
even after a storage period of over 14 months. In any case, how- 
ever, the embryos (or unfertilized ova) produced in 1957 failed 
to develop even though handled in the same fashion as others 
which survived. Eight other females, isolated in late fall or early 
winter of 1956 produced viable cases for periods of at least six 
months after isolation. 

Discussion. Several points of biological interest have arisen 
during the present study. Because of the nature of the photo 
receptive organs it is doubtful, but not certainly so, that the 
precopulatory "dance" or movements of the female attracts the 
male. More feasible is the chemical stimulation of the male by 
the female. If female oyster drills actually attract males during 
their "premating ritual" by release of an exocrine, such a chemical 
might be useful as an experimental or control tool because it is 
probably highly specific and powerful enough to be effective in 
extreme dilution. 

It has been shown that females of both species are able to store 
sperm in their seminal receptacles for extensive periods. The 
mechanisms by which they are sustained should be of interest in 
studies of cell culture and nutrition. 

Oyster drills are promiscuous and have the facility for sperm 
storage; therefore, unless sperm from previous pairings are dis- 
carded or resorbed completely before another mating occurs, a 
female contains viable sperm from several different males in her 
seminal receptacle. Under these circumstances, embryos produced 
therefrom might have different paternal hereditary materials. 
Thus, differences in rate of embryonic development, time of 
hatching, appearance, etc. may be due to varied paternity and 
not the usual genetic difference found among siblings. Until the 
precise nature of sperm storage, syngamy and egg case formation 
is understood, experimenters working with drill embryos cannot 
safely assume that all embryos in a unimaternal cluster or even a 
single egg case are of the same parentage. 


1. Eupleura caudata and U. cirierea exhibited fairly complex 

July, 1961 NAUTILUS 15 

mating behavior involving definite precopulatory movement pat- 
terns by the female, stimulation of nearby males, possibly by 
exocrine activity, assumption of relatively constant copulaiory 
positions by both sexes and copulatory groove formation by males. 
Females often twisted violently immediately prior to cessation of 

2. Urosalpinx cinerea completed copulation in a matter of 
minutes, but E. caudata often persisted intermittently for days, 
remaining in position all the while. 

3. Eupleiira caudata was apparently more promiscuous than 
U. cinerea but possibly this disparity may have been a function of 
a species different in frequency of a pairing, i.e., if E. caudata 
normally pairs more often, its comparative incidence of promis- 
cuity would naturally be greater. Or, it may also have been a 
function of the length of time that pairing consumes. For ex- 
ample, E. caudata pairing takes much longer than U. cinerea 

(12 to 20 hours vs. 3 to 4 minutes or up to 200 times longer) ; 
therefore, U. cinerea copulation would more easily be overlooked, 
resulting in a numerical bias in favor of E. caudata in any com- 
parison of frequency of pairing. 

4. Though U. cinerea paired at lower temperatures than E. 
caudata (8.2°C vs. 10.2°C) , mating activities of both increased 
during April. U. cinerea ceased copulatory activity in June while 
E. caudata persisted through the first week in July before stopping 
temporarily. Both resumed mating activities in September which 
increased in October, diminished in November and ceased alto- 
gether in early December. 

5. In our laboratory aquaria, U. cinerea copulated more fre- 
quently in the afternoon and evening while mating activities of 
E. caudata were more evenly distributed throughout the day. 

6. Females of both species stored viable sperm in their seminal 
receptacles for periods of at least 6 to 9 months. 

7. Several items of general biological interest, sperm nutrition 
and multiple paternity of embryos within a single eg^ case, etc., 
have been discussed. 

Literature cited 
Allen, J. Frances. 1958. Nautilus 72:11-15. 

Bushland, R. C, A. W. Lindquist and E. F. Knipling, 1955. Sci- 
ence 722:287-288. 

16 NAUTILUS Vol. 75 (1) 

Crabbe, E. D, 1927. Biol. Bull. 55:67-109. 

Stauber, L. A. 1943. Ecological studies on the Oyster Drill, Urosal- 
pinx cinerea in Delaware Bay, with notes on the associated 
drill, Eupleura and with special consideration of control 
methods, Unpubl. Typescript, Oyster Res. Lab., Rutgers 
Univ. New Brunswick, N. J.: 1-180. 




The living Valvatidae are widely distributed in the northern 
hemisphere, with only two species known south of the equator in 
New Caledonia. Fossils are known from the Jurassic and Tertiary 
of the Old World, as well as from the Pliocene of California and 
Pleistocene of central United States. The Neotropical region has 
three species described from Guatemala and one from Jamaica. 

However, from South America, where living species of Vnhiata 
are unknown, C. H. Fritzche in 1924 (1) described several fossil 
forms, discovered by the geologist G. Steinmann in very early Ter- 
tiary strata (Puca Formation) of Bolivia and N.W. Argentina. 
These are: 

Valvata humilis Fr. (op.cit p. 23, pl.II, f.6) , limestone of Mira- 
flores, near Potosi, Bolivia. 

Valvata yaviana Fr. (p. 23, pl.II, f.7) , and V. satira Fr. (p.24, 
pl.II,f.8) , limestone of Yavi, N. of province of Jujuy, Argentina, 
close to the Bolivian border. 

According to the original descriptions and figures, there are 
very little specific differences among Fritzche's forms, without 
sculpturing or carination, and no indication to which of the 
present subgenera they may be related, but all seem to be of regu- 
lar planorboid shape as in Valvata s.s., differing also from V. 
Q-uatemalensis Morelet or strebeli Cross, and Fisch. 

A very different form, found in strata of southern regions but 
same relative age, presents the characteristic strong keel recalling 
the subgenus Tropidina. It was previously an unknown taxono- 
mic element in the extinct Patagonian fauna, and belongs to a 
new species, as follows: 
Valvata windhauseni new species. PI. 1, figs. 1-6 

Shell very large, with almost 5 whorls (the last portion of the 

July, 1961 NAUTILUS 17 

body whoii not completely preserved) , increasing regularly, and 
very convex, but at middle zone of the last whorl an angulosity 
is marked by a strong carina or marginate keel, which starts at the 
end of the suture, above the apertural angle; below the keel, the 
body becomes convex again. The best preserved portions of the 
last two whorls, are conspicuously marked with oblique striae 
very regularly spaced (approx. 0.4 mm. apart), also visible on the 
base of the shell. Although partially covered with a hard calcar- 
eous matrix, a wdde umbilical area is observed. The median alti- 
tude of the last whorl, from suture to keel is about the same as 
the rest of the spire, and the altitude of the shell is almost equal to 
its width. The apex, well preserved, is rather obtuse. 

Dimensions: total height 6 mm.; height from apex to carina 
4.5 mm.; maj. diam. 8.5, min. diam. 7.5 mm. 

Holotype*, and 4 paratypes, from Nahuel Niyeu (25 miles W. 
of Valcheta) , Rio Negro province, Argentina, in lacustrine de- 
posits of the Jahuel Formation (Windhausen 1918), of Danian 
age (Paleocene) ; collector Dr. R. Wichmann 1923 (2) . 

The species is named in memory of Dr. A. Windhausen, who 
contributed greatly to the knowledge of the Cretaceous-Tertiary 
boundary stratigraphy in Patagonia, and its fossils. 

The smaller paratype, is a well preserved inner cast, crystallized 
by infiltration of silica solution, a characteristic of many of the 
fresh-water fossils found in these strata. This silicification is also 
present in the apex of the holotype. The other paratypes are 
larger than the holotype, showing the carina, filled with tuffaceous 
matrix, and deformed by lateral pressure, probably due to the 
strong diastrophism which subsequently affected the sediments 
during the Tertiary. 

The area where the specimens were found, south of the Rio 
Negro River, corresponds to a great depression in the territory, 
partially filled with lacustrine and tuffaceous sediments, parallel, 
and sometimes mixed, or underlying, with the Roca marine 
beds (3) , covering in unconformity the Senonian substratum. 
From this area also, an abundant fresh-water fauna is known, with 
Diplodon bondenbenderi D-J., Physa doering D-J., Physa wich- 
manni Parodiz, several pleurocerids under the old name of 
"Melania", Lioplacodes, and others (4) . This fauna shows resem- 
blance with that of Upper Cretaceous and Lower Tertiai^ of 
western United States. 

* Type and paratypes in Museo Argentine Nat. Ciencias and Carnegie 
Musuem respectively. 

18 NAUTILUS Vol. 75 (1) 

This new species is the only well carinate Valvata, fossil or 
living, known from the Neotropical region. To assign it to the 
subgenus Tropidina, on account of its strong keel, might be, how- 
ever, a taxonomic risk. The apex is not flattened but somewhat 
elevated, as in Pleurovalvata, although the spaced radial striae are 
very different from the coarse rib-like lamellae of that subgenus, 
and the general outline, on the other hand, resembles Cincinna. 
By all these combinations of features, V. windhauseni cannot be 
restricted to any of the known subgenera. Valvata is a genus very 
plastic to the enviroment, and Haas (5) reached the conclusion 
that lacustrine habits are in someway related with sculpturing in 
fresh-water mollusks and, in the case of Tropidina, the presence of 
a keeled sculpture is probably due to a genotypic feature acquired 
during prolonged lacustrine life and become invariable. For very 
variable living species, as V. utahensis, Morrison (6) questioned 
if the specimens should be distributed in thirds to the subgenera 
Cincinna, Pleurovalvata and Tropidina, or should one disregard 
these names as long as they are based on superficial characters 

Notes and references 

(1) C. H. Fritzche, Neues Jahrb, fur Min. Geol. und Palaont., 
50:1-56, taf.1-4, Stuttgart 1924. Paper continues on pages 313-334 
about older fauna which are of no concern here, but the explana- 
tions of plates all on pag. 334. 

(2) R. Wichmann, 1927, published an account of this strati- 
graphic series: Boletin Academia Nacional Ciencias, Cordoba, 
50:384-407. Recent studies proved that the moUuscan fauna is 

(3) Roca (Rocanean or "Rocanense") now considered as 
northern member of San Jorge Formation (Montian) , and it is 
synchronic with Puca Formation of Peru and Bolivia (with Val- 
vata) which Fritzche formerly placed in the Upper Cretaceous. 
See also references to other fresh-water mollusks of the Puca For- 
mation in H. A, Pilsbry, John Hopkins Univ., Studies in Geol. 
ii: 69-72, 1939; and F. Ahlfeld, Revista Museo La Plata (Geol.) 
3:1-370, 1946. 

(4) See M. Doello Jurado, BoL Acad. Nac. Cienc. Cordoba, 
50:407-416, 1927. 

(5) F. Haas, Zool. Ser. Field Museum Nat. Hist., 24, No. 8, 

(6) Naut. 55:140, 1940. 

NAin ILUS 75 (I) 

PLA 11. 1 

I'ah'ala windhauseni Paiodiz, from tvpe lot. Figs. 3 to 6, holotvpe. 

NAUTILUS 75 (1) 


Capulus sericeus Burch & Burch. Figs. 1 & 2, holotype, ventral and dorsal 
views. Figs. 3 & 4, additional examples, mentioned in text 

July, 19G1 NAUTILUS 19 



Among other interesting species taken on the Ariel Expedition 
to the Gulf of California, Aug. 30 to Sept. 6, 1960, were several 
specimens of a species of Capulus which appears to be new. 

Capulus sericeus, new species. Plate 2. 

Shell capshaped, oval; apex spiral, turned towards the pos- 
terior side, curling downwards, and twisted to the left as in some 
species of the genus Crepidula; aperture transversely oval, with 
an irregularly sinuated margin, the posterior expanded; exterior 
surface with definite and distinct axial and radiating raised lines, 
a velvety periostracum extending beyond the margins; interior 
rose color with darker rays extending from the apex to the anter- 
ior margin; shell not symmetrical but modified in shape by its 
sessile habit conforming to the surface on which it is attached; 
growth marks conspicuous but irregular. Length of holotype 14.8 
mm.; width 12.3 mm.; height 6.3 mm. 

This new form was taken commensal on Pecten sericeus Hinds, 
1845. It is obviously related to Capulus californicus Dall of Cali- 
fornia, which is commensal on Pecten diegensis Dall, but differs 
from that species in the sculpture of the exterior surface and the 
color and rays of the interior. The axial lines are much more 
pronounced than are those ©n Capulus californicus Dall, and the 
radial lines are much more prominent. The periostracum on 
Capulus sericeus is much more like overlapping shingles than on 
Capulus californicus Dall, which seems to be nearer to Capulus 
ungaricoides (Orbigny) in that it is smooth under the periostra- 
cum. On Capulus californicus Dall, the radiating lines are not as 
uniform, but seem to be more as irregularities in the shell due to 
conformation to the host shell. A study of the protoconch shows 
little variation in species, all being of the same general form, but 
a comparison indicates that Capulus sericeus has a more sunken 
protoconch. Capulus sericeus resembles Capulus hungaricus Lin- 
naeus, type species of the genus, in having radiating raised lines, 
but lacks the fine ribs that radiate from the apex toward the mar- 
gin in the latter species. The only other species described from the 
eastern Pacific is Capulus ungaricoides (Orbigny, 1841), type 
locality, Payta, Peru. Orbigny desaibed his species under the 
name Pileopsis Lamarack, 1812 which is in the synonymy of Capu- 

20 NAUTILUS Vol. 75 (1) 

lus Montford, 1810. This species is smooth except for radiating 
raised lines on the periostracum. We wish to thank Dr. S. P. 
Dance, British Museum (Natural History) for his advice that 
Capulus ungaricoides (Orbigny) does not seem to him to be the 
same species. In addition to the shell characters mentioned above, 
we are convinced, from a study of ocean currents, that it is un- 
likely that this occurrence constitutes a range extension from 
Peru. Capulus chilensis Dall, 1904, from the Antarctic, is possibly 
a species related to Capulus ungaricoides (Orbigny) . 

The type specimen (pi. 2, figs. 1 & 2) was trawled off Cabo 
Haro, near Guaymas, Sonora, Mexico at a depth of 100 fathoms, 
and will be deposited in the Stanford University Type Collection, 
no. 8519. This specimen is from the collection of Dr. Homer King. 

Other specimens here named paratypes are: one specimen 
trawled off Cabo Haro, Sonora, Mexico in 20-40 fathoms, Dec. 27, 
1959 on the ship General Yanes, collection of Dr. Donald Shasky; 
one specimen trawled off Cabo Haro, Sonora, Mexico, collection 
of Mr. Mark Rogers. 

The following are mentioned as hypotypes, all from the Gulf 
of California, but not the type locality; one specimen trawled off 
Espiritu Santo Island in 40-90 fathoms, fig. 3, collection of Dr. 
Donald Shasky; one specimen trawled off Monserrate Island in 
20-40 fathoms, Sept. 1, 1960, fig. 4, collection of Dr. Donald 
Shasky; one specimen trawled off MonseiTate Island in 75 fath- 
oms, collection of Mr. Mark Rogers. 


Carcelles, Alberto. 1944. Comm. Zool. del Museo De Hist. Nat. de 

Dall, W. H. 1889. Marine mollusks of the southeastern coast. U.S. 

Nat. Mus. Bull. 37:152. 
Dall. W. H. 1900. Naut. 13 (9): 100. 
Dall, W. H. 1908. Mollusca and Brachiopoda. Albatross Rep. Bull. 

Mus. Comp. Zool. ^i (6) :329. 
Dall, W. H. 1927. In Eastman-Zittel, Textbook of Paleontology, 

2nd ed., London. 
Orbigny, A. d' 1811. Amer. Merid. (Moll.), p.457. 
Thiele, Johannes. 1931 Handbuck der systematischen Weichtier- 

kunde, vol. 1, p. 246. 
Tryon, George W. 1886. Man. of Conch. (1) 5:105,131. 

July, 1961 NAUTILUS 21 


(Concluded from April no.) 

Genera, subgenera and species in the family Rissoellidae 

The genera and subgenera Laciinella (family Lacunidae) , 
Fairhankia (Micromelaniidae) , Hyala, "Dardania" (=:Darda- 
nula), Tatea (Rissoidae) , and Diala (Cerithiidae) have, at one 
time or another, been wrongly included in the Rissoellidae by 
various malacologists. Some of the species listed below (notably 
those named by A. Adams and W. H. Turton) probably are also 
not rissoellids. All species originally named as rissoellids are in- 
cluded in the list, as well as species which have been rightly or 
wrongly transferred to rissoellid genera. 

alhella, Rissoa. Alder (1844) . See also Thompson (1844) . An 
older name for R. glabra Alder non Brown than R. diaphana. 
The name R. albella Alder never has been adopted and the "50 
year rule" is here invoked to suppress it (I.C.Z.N. ruling 
pending) . 

alderi, fjeffreysia. Carpenter (1856). "Mazatlan" [W. coast 
Mexico]. According to Bartsch (1920) a Barleeia (Rissoidae) . 
See also Baker, Hanna & Strong (1930) . 

angulijerens, Rissoa. Folin (1869) . "Bale de Panama." Re- 
ferred by Bartsch (1920) to Rissoella. 

atlantica, Jeffreysia. Smith (1892). "St. Helena." 

bakeri, Rissoella (?) . Strong (1938). "9 to 15 fms. off Guada- 
lupe Island, Mexico." 

becki, Jeffreysia. Turton (1932) . "Port Alfred, South Africa." 

bifasciata, Jeffreysia. Carpenter (1856). "Mazatlan" [W. coast 
Mexico]. See Bartsch (1920). 

caffra, Rissoa (Cingula). Sowerby (1897). "Port Elizabeth" 
[South Africa]. Referred by Bartsch (1915) and Turton (1932) 
to Jeffreysia. Specimens collected by Turton at Port Alfred 
(M.C.Z. 101515) are not rissoellids. 

californica, fRissoella. Bartsch (1927), "San Clemente Island, 

capensis, Rissoa (Cingula). Sowerby (1892) . "Port Elizabeth" 
[South Africa]. Apparently this is the species referred by Bartsch 
(1915) io Jeffreysia. See Turton (1932). 

caribaea, Rissoella (Phycodrosus). Rehder (1943a) . "Bonefish 
Key, Fla." [One of the Crawl Keys between Grassy Key and V^aca 
Key, Florida Keys.] 

cylindrica, Jeffreysia. Jeffreys (1856) . ". . . in about 12 fathoms 
at Spezia" [Italy]. Referred by Monterosato (1878) to Aclis 

22 NAUTILUS Vol. 75 (1) 

(Aclididae) . Specimens from Sicily (A.N.S.P. 247781), which 
presumably are correctly identified, have hyperstrophic proto- 
conchs and are pyramidellids. 

diaphana, Rissoa? Alder (1848. New name for R.? glabra 
Alder non R. glabra (Brown) . Referred by Forbes & Hanley 
(1850) to Jeffreysia and incorrectly by Clark (1850) to "Chem- 
nitzia" (Pyramidellidae) , who claimed, wrongly, that the apex 
is reflexed (i.e., hyperstrophic) ; see Jeffreys (1851) and Alder 
(1851a) . See albella and nitidus. 

duperrei, Paludestrina. Velain (1877). [Misspelled 'Duperei' 
p. 144.] "He Saint-Paul; dans I'ouest du banc Roure, par les 
fonds de 35 metres a 50 metres." [S. Indian Ocean]. Referred by 
Thiele (1912) to Jeffreysiopsis. 

eburnea, Rissoa. Stimpson (1851a, b) . ". . . in thirty fathoms, 
off Cape Ann" [Massachusetts]. Doubtfully and wrongly referred 
by Gould (1870) to Rissoella. An "Odostomia" (Pyramidellidae); 
see Bartsch (1909) . 

edzvardiensis, Jeffreysia. Watson (1880). [Challenger] "St. 145a 
. . . Lat. 46° 41' S., long. 38° 10' E. Prince Edward Island, between 
Cape of Good Hope and Kerguelen. 50 to 150 fms." Figured Wat- 
son (1886); possibly from 310 fms., or from Sta. 145. Referred 
doubtfully by Thiele (1912) to Jeffreysiella, and definitely by 
Iredale (1912) to Heterorissoa. 

excolpa, Rissoella. Bartsch (1920) . "Concepcion Bay, Lower 
California" [Mexico]. See Baker, Hanna &: Strong (1930). 

julgida, Rissoella. "A. Ad." Dunker (1882). [Japan]. 

fiiscotincta, Jeffreysia. Turton (1932) . "Port Alfred, South 

galba, Rissoella. Robertson, this paper. 

glabra, Rissoa?. Alder (1844). "Dalkey Island near Dublin 
[Ireland], and at Cullercoats, Northumberland [England]." A 
misapplication of the name Rissoa glabra (Brown) Brown (1844), 
a pyramidellid. R. glabra Alder referred to Rissoella by Gray 
(1847) . Weinkauff (1868) wrongly referred i?. g/o 6 ra (Brown) 
to Jeffreysia. See diaphana. 

globularis, Jeffreysia. "Jeffreys, MS." Forbes & Hanley (1852). 
". . . at Skye [Scotland] and the Shetlands . . ." Figured (1853) . 

gulsonae, Chemnitzia. Clark (1850). ". . . prope ostia Iscae 
Danmoniorum." [Exmouth, England]. Referred doubtfully by 
Jeffreys (1851) to Jeffreysia and later (1867) to Aclis. Type 
species of Pherusina (Aclididae) . 

hera, Jeffreysia. Turton (1932). "Port Alfred, South Africa." 

hertleini, Rissoella. Smith & Gordon (1948). "10 fathoms . . . 
off Cabrillo Point, Monterey Bay, California." 

Heterorissoa Iredale (1912). Type species (original designa- 
tion) : H. secunda. Synonym of Jeffreysiella according to Thiele 

July, 1961 NAUTILUS 23 

hydrophana, Rissoella. A. Adams (1860) . "Tabu-Sima [Tobi 
Shima]; 25 fathoms" [Japan]. 

indistincta, Jeffreysia. Turton (1932) . "Port Alfred, South 

infiata, Jeffreysia. Monterosato (1878). Nom. nudum. "Pal, 
[ermo] e Trap, [ani] (Monts.) ; Messina (Granata) ." [Sicily]. 

Jeffreysia Alder in Forbes & Hanley (1850) . Type species (or- 
iginal designation) : /. diaphana. Objective junior synonym of 

Jeffreysiella Thiele (1912) . Type species (original designation): 
/. notnbilis. 

Jeffreysilla Thiele (1925) . Type species (monotypy) : Rissoella 

Jeffreysina Thiele (1925) . Type species (subsequent designa- 
tion, Winckworth, 1932) : /. globularis. Ranked as genus by 
Rehder (1943a). 

Jeffreysiopsis Thiele (1912) . Type species (here designated) : 
Paludestrina duperrei Veiain. Synonymized with Rissoella, s.s., 
by Thiele (1925). 

joJtnstojii, Rissoella. Baker, Hanna &: Strong (1930) . "Cape 
San Lucas, Lower California" [Mexico]. 

nialayensis, Rissoella? Thiele (1925). "Station 211 . . ." [805 
meters, 7° 48.8' N., 93° 7.6' E., near Nicobar Islands]. 

minima, Rissoella. A. Adams (1860) . "Tsu-Sima; 26 fathoms" 

mundula, Rissoella. A. Adams (1860) . "Tsu-Sima; 26 fathoms" 

nitidus. Turbo. J. Adams (1797). Nom. dubium. "Pembroke- 
shire" [Wales]. Considered by Jeffreys (1867) a possible older 
name for Rissoa? diaphana Alder. 

nitida, Jeffreysia. "Sars" Friele (1876). Hydrobia nitida Sars 
(1859), nom. nudum. "Bergen" [Norway]. Close to "Rissoella" 
eburnea (Stimpson) , a pyramidellid, according to Sars (1878) . 
See eburnea. 

notabilis, Jeffreysiella. Thiele (1912) . "Observatory Bay . . . 
Kerguelen." [S. Indian Ocean]. See also Thiele (1925). 

omphalotropis, Rissoella. A. Adams (1860) . "Sado; 30 fathoms" 

opalina, Rissoa (?) . Jeffreys (1848). "Guernsey and Sark" 
[Channel Islands]. Referred by Forbes & Hanley (1850) to Jej- 
freysia and incorrectly by Clark (1851) to "Chemnitzia" (Pyram- 
idellidae; see Alder (1851b). 

pauli, Rissoella. See sancti-pauli. 

Phycodrosus Rehder (1943a) . Type species (original designa- 
tion) : Rissoella caribaea. Subjective synonym of Jeffreysilla (this 
paper) . 

Rissoella Gray (1847). Type species (monotypy): "Rissoa? 

24 NAUTILUS Vol. 75 (1) 

glaber, Alder." [z=Rissoa? diaphana Alder]. 

sancti-pauli, Rissoella [ (Jeffreysia)]. Velain (1877). Velain 
(1876), nom. nudum. "He Saint-Paul . . . au niveau de la basse 
mer . . ." [S. Indian Ocean]. Emended to R. paidi by Crosse (1879). 

secunda, Heterorissoa. Iredale (1912). "Kermadec Islands." 
[S. Pacific Ocean]. 

simoniana, Jeffreysiopsis. Thiele (1912) . "Simonsbai" [prob- 
ably Simonstown, nr. Cape Town, South Africa]. 

spiralis, Rissoella. A. Adams (1860). "Sado; 30 fathoms" 

sulcosa, Phasianella. Mighels (1843). "Casco Bay [Mame]. 
Wrongly referred by Gould (1870) to Rissoella. An "Odostomia" 
(Pyramidellidae) ; see Johnson (1915) . 

tenuis, Jeffreysia. Turton (1932) . "Port Alfred, South Africa." 

translucens, ? Jeffreysia. Carpenter (1866). Carpenter (1864), 
nom. nudum. "S. Diego" [California]. Referred by Bartsch (1920) 
to "Syncera" (=Assiminea, Assimineidae) . 

tumens, Jeffreysia. Carpenter (1856). "Mazatlan" [W. coast 
Mexico]. Referred by Carpenter (1866) to "Cythna" (PStilifer- 
idae) . Retained by Bartsch (1920) in Rissoella. See also Baker, 
Hanna & Strong (1930) . 

turgidula, Rissoella. A. Adams (1860). "Korea Strait; 46 fath- 

vesicalis, Rissoella. A. Adams (1860) . "Sado; 30 fathoms" 

vitrina, Rissoella. A. Adams (1860). "Tabu-Sima [Tobi Shima]; 

25 fathoms" [Japan]. 

wilfredi, Jeffreysia. Gatliff & Gabriel (1911). "Ocean beach, 
near Point Nepean" [Victoria, Australia]. Referred by Iredale 
(1912) to Heterorissoa (misspelled 'wilfridi') . 

zebra, Rissoella. Thiele (1925) . "Deutsch-Ostafrika . . . Dar- 
essalam" [Tanganyika]. 

Adams, A. 1860. Ann. Mag. Nat. Hist., (S) 6: 332, 333. 
Adams, J. 1797. Trans. Linn. Soc. [London], 3: 65. 
Alder, J. 1844. Ann. Mag. Nat. Hist., (1) 13: 325, 326, pi. 8, figs. 

1-4. 1848. Trans. Tyneside Nat. Field Club, 1: 149. 1851a. Ann. 

Mag. Nat. Hist., (2)7: 193-196. 1851b. Ibid., pp. 460-465. 
Baker, F., G. D. Hanna Sc A. M. Strong, 1930. Proc. California 

Acad. Sci., (i) 19: 36-38, fig. 3, pi. 1, figs. 10, 13, 16. 
Bartsch, P. 1909. Proc. Boston Soc. Nat. Hist., 34: 109. 1915. Bull. 

U.S. Natl. Mus., 91: 134, 135, 225. 1920. Proc. U.S. Natl. Mus., 

^8: 159-164, 175, 176, pi. 12, figs. 1-3, 6, 7, 9. 1927. Ibid. 70(11) : 

31, pi. 4, fig. 2. 
Brown, T. 1844. Illustr. Rec. Conchology Gt. Britain and Ireland, 

ed. 2, p. 13, pi. 9, fig. 37. 
Carpenter, P. P. 1856. Cat. Mazatlan Shells British Mus., pp. 

July, 1961 NAUTILUS 25 

361-363. 1864. Rept. British Assoc. Adv. Sci. "1863," pp. 613, 

657. 1866. Proc. California Acad. Sci., (1) 3: 219. 
Clark, W. 1850. Ann. Mag. Nat. Hist., (2) 6: 454, 455, 459. 1851. 

Ibid., (2) 7: 292-297. 1855. British Mar. Test Moll., pp. 7, 

Crosse, H. 1879. Jour. Conchyl., 27: 55. 
Dunker, G. 1882. Index Moll. Mar. Japonici, p. 117. 
Folin, L. de. 1869. Fonds de la Mer, 1: 134, pi. 20, fig. 6. 
Forbes, E. & S. Hanley. 1850. Hist. British Moll., 3: 151-155, pi. 

J.}., figs, 1, 2, pi. 76, figs. 1, 3, 4. 1852. Ibid., 4: 267-269, pi. 133, 

figs. 5, 6. 1853. Ibid., pi. M.M., fig 2. 
Fretter, V. 1948. Jour. Mar. Biol. Assoc. U.K., (2)27: 597-632, 

figs. 1-6, pi. 4. 1956. Proc. Zool. Soc. London, 126: 380. 1954 

(& A. Graham) . Jour. Mar. Biol. Assoc. U.K., (2) 33: 577-583. 
Friele, H. 1876. Forh. Vid.-Selsk. Christiania, "1875," p. 61, pi. 1, 

fig. 6. 
Gatliff, J. H. & C. J. Gabriel. 1911. Proc. Roy. Soc. Victoria, 

{2)24: 188, 189, pi. 46, fig. 3. 
Gould, A. A. 1870. Rept. Invert. Massachusetts, ed. 2, pp. 296, 297, 

figs. 564, 565. 
Gray, J. E. 1847. Proc. Zool. Soc. London, 15: 159. 
Gray, M. E. 1850. Figs. Moll. Anim., 4: 86. 
Iredale, T. 1912 [Oct. 30]. Proc. Malac. Soc. London, 10: 221, 

222, 1 fig. 1915. Ibid., 11: 332. 
Jeffreys, J. G. 1848. Ann. Mag. Nat. Hist., (2)2: 351. 1851. Ibid., 

(2)7: 465-469, pi. 15. 1856. Ibid., (2)17: 184, pi. 2, figs. 8, 9. 

1867. British Conch., 4: 58-63, 106, 107, pi. 1, fig. 3. 
Johnson, C. W. 1915. Occas. Papers Boston Soc. Nat. Hist., 7: 99. 
Lebour, M. V. 1936. Jour. Mar. Biol. Assoc. U.K., (2)20: 552, 

pi. 1, fig. 17. 
McGinty, T. L. 1948. Mollusca (Paul H. Reed, Tavares, Florida) , 

2: 63. Mimeogr. 
Mighels, J. W. 1843. Boston Jour. Nat. Hist., 4: 348, pi. 16, fig. 4. 
Monterosato, T. A. di. 1878. Gior. Sci. Nat. Econ. Palermo, 13: 

87, 91. 
Rehder, H. A. 1943a. Proc. U.S. Natl. Mus., 93: 194, pi. 20, fig. 7. 

1943b. Naut., 57: 33. 
Robertson, R. 1960. American Malac. Union Ann. Repts. "1959," 

pp. 22, 23. In press. Collecting minute mollusks which live in 

marine algae. In How to collect shells. Publ. American Malac. 

Union, ed. 2. 
Sars, G. O. 1878. Moll. Reg. Arcticae Norvegiae, pp. 206, 347, 

pi. 10, fig. 13, pi. 34, fig. 8, pi. vi, fig. 16. 
Sars, M. 1859. Forh. Vid.-Selsk. Christiania, "1858," p. 85. 
Smith, A. G. & M. Gordon, Jr. 1948. Proc. California Acad. Sci., 

(4)26: 224, 225, pi. 3, fig. 15. 
Smith, E. A. 1892. Ann. Mag. Nat. Hist., (6) 10: 130, pi. 12, fig. 7. 
Sowerby, G. B. 1892. Mar. Shells South Africa, p. 38, pi. 2, fig. 41. 

26 NAUTILUS Vol. 75 (1) 

1897. Appendix Mar. Shells South Africa, p. 17, pi. 6, fig. 15. 
Stimpson, W. 1851a. Proc. Boston Soc. Nat. Hist., 4: 14. 1851b. 

Shells New England, p. 34, pi. 1, fig. 1. 
Strong, A. M. 1938. Proc. California Acad. Sci., (4)25: 211, 212, 

pi. 15, fig. 5 
Thiele, J. 1912 [Aug.]. Deutsche Sudpolar-Exped. 1901-1903, 13, 

Zool. 5: 239, 240, 260, 272, 276-278, fig. 13, pi. 15, fig. 5, pi. 16, 

figs. 3, 4, pi. 19, figs. 15, 21. 1925. Wiss. Erg. deutschen Tiefsee- 

Exped. "Valdivia," 17: 87, 91, 92, 363, pi. 16, figs. 13-15, pi. 19, 

figs. 13-15. 1929. Handb. syst. Weichtierkunde, /: 178, 179, figs. 

162, 163. 
Thompson, W. 1844. Rept. British Assoc. Adv. Sci., "1843," p. 

Turton, W. H. 1932. Mar. Shells Port Alfred S. Africa, pp. 151- 

153, pi. 34, figs. 1080, 1083-1087. 
Velain, C. 1876. C. R. Acad. Sci. [Paris], 83: 285. 1877. Arch. Zool. 

Exper. Gen., 6: 115, 116, 144, pi. 3, figs. 18-20. 
Watson, R. B. 1880. Jour. Linn. Soc. [London], Zool., 15: 99. 

1886. Rept. Voy. Challenger. Zool., 15: 584, 701, pi. 43, fig. 5. 
Weinkauff, H. C. 1868. Conch. Mittelmeeres, 2: 275. 
Winckworth, R. 1932. Jour. Conchol., 19: 217, 223. 



PoLYGYRA GRACILIS, new species. PI. 4, figs. N & O. 

Shell pale brown, base lighter, translucent, shining, of 5 to 5.5 
whorls. Spire slightly convex, of slowly increasing whorls, the last 
shortly descending in front; periphery above the middle, rounded. 
First whorl smooth, following whorls becoming inaeasingly stri- 
ate; last two whorls weakly ribbed above, base smooth with 
growth lines only, or with weak ribs behind the lip. Umbilicus 
well-like, expanding in the last whorl to about 4 times its earlier 
diameter, contained about 3.5 times in the diameter of the shell; 
last whorl with a shallow furrow parallel to the umbilical suture. 
Aperture very oblique, deeply furrowed behind, outer and basal 
margins of lip strongly reflected, thickened within, its inner edge 
bearing two teeth; one on the basal margin and one on the outer 
margin. Ends of the lip joined by a parietal callus bearing a 
V-shaped tooth. A short distance within there is a callous tubercle 
on the columellar axis. 





3.9 mm. 

8.3 mm. 


0.47 Holotype. 

4.1 mm. 

9.1 mm. 


0.45 Paratype. 

3.6 mm. 

7.8 mm. 


0.46 Paratype. 

3.2 mm. 

6.7 mm. 


0.48 Medina River drift 

2.7 mm. 

6.7 mm. 


0.40 Nueces River drift. 

NAUTILUS 75 (1) 


Figs. 1 & 2, Xotia tridcns. 1, internal view of pallial complex. 2. penis and 
accessories; optical longitudinal .section of flattened mount in glycerin jelly; 
semi-diagrammatic. Figs. 3 & 4, Cerion uva. 3, penis, cut open and pinned 
out. 4, terminal genitalia, dissected apart. .Scales represent 1 mm. Drawn with 
aid of camera lucida. 



Holotypcs: A-C, Succinca solaslra Hubricht. D & E, .S'. urbana Hubricht. 
F-H, Calinella lexana Hubricht. I-K, C. pugilntor Hubricht. L & M, Pol\g\ra 
litliira Hubricht. N & O, P. gracilis Hubricht. P & Q, Stciiotreiua calvescens 
Hubricht. Pliotographs by John B. Burch, University of Michigan. 

July, 1961 NAUTILUS 27 

Distribution. — Texas: Comal Co.: Guadalupe River bluff, 2 
miles south of Sattler, holotype 205894 and paratypes 205893, 
University of Michigan Museum of Zoology, other paratypes 
14834, collection of the author; drift, Guadalupe River, New 
Braunfels. Kerr Co.: 12 miles southwest of Kerrville; river bluff, 
5 miles southwest of Hunt. Bandera Co.: drift, Medina River, 4.7 
miles northwest of Medina. Medina Co.: Hondo. Real Co.: Frio 
River bluff, 5.5 miles north of Leakey. Uvalde Co.: drift, Nueces 
River, 7 miles west of Uvalde; drift, Nueces River, 1.5 miles south 
of Laguna; Garner State Park. 

Polygyra gracilis is a species of river bluffs and ravines in the 
Edwards Plateau of Texas. It is most closely related to Polygyra 
mooreana (W. G. Binney) , which is found in the same region. It 
differs in its more slender whorls, more depressed shape, and more 
glossy and translucent shell. Of the specimens measured P. gracilis 
had a height diameter index below 0.50 with an average of 0.45, 
while in P. mooreana the measured shells were all above 0.50 with 
an average of 0.57. The lip teeth are usually smaller and more 
widely spaced and are not as laterally compressed. The parietal 
tooth usually does not have the distinct lower angle found in P. 
mooreana, being more rounded. Despite its close relationship to 
P. mooreana, the two species could be readily separated in river 
drift material. 
Polygyra tamaulipasensis Lea. 

Polygyra texasiana texasensis Pilsbry, 1940, Land Moll. N. 
Amer. /, p. 619, fig. 394g, h. 

Polygyra texasensis Pilsbry, Pilsbry & Hubricht, 1956, Naut. 
69: 94. 

Polygyra texasiana tamaulipasensis Lea, Pilsbry & Hubricht, 
1956. Naut. 69: 95, PI. 5, figs., 1,1a. 

This species has been considered to be a subspecies of Polygyra 
texasiana (Moricand) , but is quite distinct. I have seen no inter- 
gradation in the beach drift material, nor in any material collec- 
ted in west Texas. It differs from P. texasiana in having a more 
depressed shell with narrower whorls and a larger umbilicus. The 
shell is never banded. The parietal tooth is longer, higher and 
straighter, the outer end extending a little beyond the lower end 
of the lip. The outer end of the upper branch is enlarged into a 
low tooth. The animal is pale gray rather than brownish-yellow. 
It lives in rock piles and under prostrate yuccas on the tops of 
the mesas, while P. texasiana in west Texas lives in low ground 
near streams and ponds. 

28 NAUTILUS Vol. 75 (1) 

Polygyra texasiana is quite variable in its sculpture. Those 
from near the coast are often strongly ribbed above and below. 
Away from the coast the ribbing on the base disappears and they 
are only ribbed above. Farther west the upper surface becomes 
smooth. Within the range of P. tamaulipasensis, P. texasiana can 
not be distinguished by the sculpture. The presence of the tooth 
on the end of the upper branch of the parietal tooth will always 
separate P. tamaulipasensis from P. texasiana. 

P. tamaulipasensis is found living from near Junction, Kimble 
Co., west to Alpine, Brewster Co. and north to Colorado City, 
Mitchell Co., Texas. The record from Lyford, Cameron Co., 
Texas is based on P. scintilla Pilsbry & Hubricht. The record 
from near Roswell, New Mexico is based on a smooth P. texasiana. 
Polygyra lithica, new species. PI. 4, figs. L & M 

Shell depressed, pale brown, of 5 to 6 whorls. Spire weakly con- 
vex, of slowly increasing whorls, periphery rounded. Embryonic 
whorls smooth, later whorls becoming increasingly striate, last 
two whorls weakly rib-striate, rib-striae becoming much weaker 
below the periphery, base with distinct spiral striae. Umbilicus 
deep, well-like, expanding in the last whorl to about one-third 
the diameter of the shell. Aperture oblique, deeply furrowed 
behind, lip, reflected and strongly thickened, with two teeth on 
the inner margin, the basal tooth laterally compressed, not im- 
mersed; the outer tooth only slightly immersed, of about the same 
size as the basal tooth. Ends of the lip joined by a heavy parietal 
callus bearing a heavy tooth which is rounded in front view. A 
short distance within there is a callous tubercle on the columellar 





3.2 mm. 

7.4 mm. 


0.43 Holotype. 

3.7 mm. 

8.1 mm. 


0.46 Paratype. 

3.3 mm. 

7.0 mm. 


0.49 Paratype. 

2.7 mm. 

7.9 mm. 


0.46 Rushing. 

Distribution. — Arkansas: Stone Co.: upland oak-hickory woods, 
6 miles east of Mountain View, holotype 205896 and paratypes 
205895 U.M.M.Z., other paratypes 15917, collection of the author: 
4 miles northwest of Allison; 2 miles northeast of Rushing. 

Polygyra lithica is related to P. dorfeuilliana Lea, differing in 
the teeth. The parietal tooth is lower and is rounded rather than 
squarish. The lip teeth are smaller and are not deeply immersed. 
It is apparently a species of quite limited range, being known only 
from a small area in north-central Arkansas. 
Stenotrema calvescens, new species. PI. 4, figs. P &: Q. 

Shell buffy brown to bister, globose-conic, imperforate, of about 



6.5 mm. 

9.7 mm. 

5.6 mm. 

8.1 mm. 

6.0 mm. 

9.3 mm. 

6.7 mm. 

10.2 mm. 

July, I9()l NAUTILUS 29 

5.5 whorls; periphery above the middle, rounded or slightly sub- 
angular in [rent of the aperture. Embryonic whorls sculptured 
with radially elongated granules, later whorls with fine radial 
lines, last three whorls with numerous soft deciduous liairs. 
Aperture naiTow, very pale brown, outer edge of the basal lip 
wholly adnate, inner edge with a distinct U-shaped central notch; 
interdenticular sinus rounded, moderately deep; there is no tooth 
within the outer arc of the lip. Parietal tooth rather low and 
slender, curved, leaning towards the basal lip. There is no buttress 
connecting the parietal tooth to the end of the outer lip. Edge of 
the parietal callus extending well beyond the parietal tooth. 
Fulcrum thin with slightly convex edge. 

Whorls H./D. 
5.7 0.67 Holotype. 

5.2 0.69 Paratype. 

5.6 0.65 Paratype. 

5.5 0.63 Hixon. 

Distribution. — Tetinessee: Marion Co.: Cumberland Mtn., 1 
mile east of Monteagle, holotype 205898 and paratype 205897 
U.M.M.Z., other paratypes A9690, collection of the author; Cimi- 
berland Mtn., 2.4 miles southeast of Monteagle; West Fork of 
Pr)'or Cove, 2 miles northeast of Jasper; under pine logs, summit 
of Suck Creek Mtn., 15 miles west of Chattanooga (W. F. Shay, 
coll.) Hamilton Co.: vacant lot, Thrasher Pike, Hixon (James 
R. Hood, coll.) ; near Silver Creek, 300 Signal Mtn. Road, North 

In the West Fork of Pryor Cove, Stenotrema calvescens was 
found associated with 5. stenotrema (Pfr.) from which it could 
be readily separated by its smaller size, less reddish color, finer, 
more numerous hairs, and by the absence of a buttress. The 
hairs are found only in young and recently matured shells. They 
are lost in old shells. Scrubbing of the shells in cleaning will 
remove most of the hairs. Stenotrema calvescens is apparently 
most closely related to S. florida Pilsbry, from which it differs in 
its smaller size, larger lip notch, and finer, more numerous hairs. 
Praticolella berlandieriana campi Clapp & Ferriss. 

Praticolella campi Clapp & Ferriss, 1919, Naut. 32: 78, PL 6, 
figs. 1-4. 

Praticolella berlandieriana taeniata Pilsbry, 1940, Land Moll. 
N. Amer. /, p. 696, figs. 427g, h. 

Praticolella campi is a winter resting stage of the form later 
described as P. berlandieriana taeniata Pilsbry. It is found only 
in the winter and is always found associated with adult shells. It 
always agrees in color pattern with the adult shells found with 
it. Adult shells sometimes show an iiTegularity in the spire corres- 

30 NAUTILUS Vol. 75 (1) 

ponding to this resting stage. Animals dissected by both Dr. 
Pilsbry and myself were found to be sexually undeveloped. It is 
a stage in which there is a high mortality, as they are common in 
river drift and fossil deposits while other immature stages are 
rare. The name campi will have to replace taeniata for the 
Praticolella berlandieriana X P. griseola hybrid populations of 
southern Texas and northeastern Mexico. 

Similar resting stages are also found in Praticolella griseola 
(Pfeiffer) and Praticolella pachyloma ('Menke' Pfeiffer). 
SucciNEA soLASTRA, new Species. PI. 4, figs. A-C; text-fig. 1, B. 

Shell pale buff, translucent, shining, with about 3.5 whorls, 
thin but firm, elongate-ovate in shape, sculpture of unevenly 
spaced growth wrinkles. Spire acute, moderately long, sutures 
well marked, periphery somewhat flattened. Aperture ovate, 
occupying about sixty percent of the length of the shell, outer 
and basal margins well rounded. Columella nearly straight. 

Animal pale, mantle pale with scattered brownish spots. Herma- 
phrodite duct well pigmented. Talon moderately pigmented, club- 
shaped with two terminal lobes and a longitudinal groove down 
the center, giving it the appearance of being bifurcate. Prostrate 
gland large, irregularly oval, compressed by surrounding organs, 
unpigmented. Vas deferens not very long, entering the penis a 
little below the apex. Vagina and atrium very short. Vaginal 
retractor muscle long and stout. Penis rather stout, with a con- 
striction about two-thirds the way up, above this it is somewhat 
inflated and free from the sheath. Penial retractor muscle thick 
and broad. Spermatheca globose, the duct slender throughout 
its length. 



Ap. H. 

Ap. W. 


15.3 mm. 

7.3 mm. 

9.7 mm. 

5.7 mm. 

3.5 Holotype. 

16,0 mm. 

7.3 mm. 

9.3 mm. 

5.7 mm. 

3.5. Paratype. 

16,0 mm. 

8.7 mm. 

9.7 mm. 

6.3 mm. 

3.5 Paratype. 

1 1 .7 mm. 

6.0 mm. 

7.0 mm. 

4.0 mm. 

3.5 Combes, Texas. 

9.5 mm. 

4.7 mm. 

5.7 mm. 

3.2 mm. 

3.5 Combes, Texas. 

Distribution. — Texas: Terrell Co.: Sanderson; 5 miles north of 
Sanderson; side of mesa, 5 miles southeast of Sanderson. Val- 
Verde Co.; Shumla. Uvalde Co.; Uvalde, Dimmit Co.; just 
north of Caterina; near culvert, 6.5 miles south of Caterina; 
near culvert, 2.5 miles northwest of Canrizo Springs. Frio Co.: 
culvert, 5.5 miles southwest of Moore, Bexar Co.: culvert, on 
US-281, 9.5 miles north of San Antonio. Live Oak Co.: near 
Ramirena Creek, 13 miles south of George West. Bee Co.: culvert, 
2 miles south of Pettus Jim Wells Co.: Premont. Starr Co.: cul- 
vert, 6 miles northwest of Roma; 1,5 miles southeast of Rio 
Grande City. Hidalgo Co.: near Bentzen Rio Grande Valley State 
Park, holotype 205900 and paratype 205899 U.M.M.Z., other para- 

July, 1961 



types 14322, collection ot the author. Willacy Co.: roadside, 0.7 
miles north of Santa Monica, Cameron Co.: 6.3 miles southwest of 
Boca Chica; 0.5 mile northwest of Combes; dump, 3.4 miles east 
of La Paloma. Mexico: Tamaulipas: 1 mile southeast of Ciudad 
Mier; San Fernando. Nuevo Leon: near arroyo, 4 miles north- 
east of Pesqueria Chica; 32.8 miles east-northeast of General 

Figure 1, genitalia: A & C, Succinea urbana Hubricht. B, S. solastra Hu- 
bricht. D & E, Catinella pugilator Hubricht. F-H, C. texana Hubricht. (ap., 
appendix; at, atrium; h.d., hermaphrodite duct; p., penis; p.g. prostate; v., 
vagina; s., spermatheca; v.d., vas deferens; t., talon.) 

Succinea solastra is frequently found associated with Succinea 
luteola Gould, from which it may be readily separated by the 
anatomical differences, its smaller size, more acute spire, and 
translucent shell. The shell is usually covered with dirt while 
that of S. luteola is usually clean. It is most closely related to 

32 NAUTILUS Vol. 75 (1) 

Succinea grosvenori which differs in its more globose, more 
opaque shell; and dark mantle. The genitalia are not distinguish- 
able from those of S. grosvenori Lea. 

An examination of many lots of anatomical material of suc- 
cineids, collected over the past several years by the author, has 
shown that the answer to the species problem in Succinea will 
not be found in the study of the genitalia. Species with readily 
distinguishable shells have very similar genitalia. Thus, Succinea 
pronophobus Pilsbry, S. wilsoni Lea, and S. urbana, new species, 
are not separable by the genitalia. Succinea campestris Say, S. 
luteola Gould, and 5. floridana Pilsbry, form another similar 
group, (The difference in the thickness of the penial sheath re- 
ported by Pilsbry does not exist in my material. Possibly the 
thickness may vary with the age of the animal.) The genitalia 
will be of value in the recognition of species groups, but shell 
characters will have to be used in many cases for species identifi- 
cation. On the other hand, in the genus Catinella very good spe- 
cific differences are to be found in the anatomy, especially in the 
shape of the penis. In this genus this will probably prove to be 
a more reliable character than the shape of the shell. 
Succinea urbana, new species. PL 4, figs. D & E; Text-fig. 1, A & C. 

Shell pale golden-brown in color, subtranslucent, dull, with 
about 3.5 whorls, elongate-ovate, rather thin, sculpture of un- 
evenly spaced growth wrinkles. Spire acute, long, sutures only 
moderately impressed; whorls moderately convex. Aperture ovate, 
rather small, occupying about 55% of the length of the shell, 
outer and basal margins well rounded. Columella nearly straight. 

Color pattern variable, some with mantle grayish, others with 
a bold pattern of black and pale grayish-brown, margin of foot 
gray. Hermaphrodite duct moderately pigmented. Talon mod- 
erately pigmented, club-shaped, two lobed and with a median 
groove. Prostate gland large, almost square due to pressure from 
the other organs, moderately pigmented. Vas deferens not very 
long, entering the penis below the terminal loop, somewhat 
expanded below where it joins the penis. Penis moderately 
slender, with an inflated loop at the upper end. Penial retractor 
connected to the penis below the terminal loop, Atrium and 
vagina very short. Spermatheca large, globose, duct slender. 

Length Diam. Apt. L. Ap. W 

11.1mm. 6.0 mm. 6.0 mm. 4.0 mm. Holotype. 

10.9 mm. 5.7 mm. 6.0 mm. 4.0 mm. Paratype. 

11.7 mm. 6.3 mm. 6.6 mm. 4.2 mm. Paratype. 

Type Locality. Alabama: Montgomery Co.: vacant lot, Dudley 
St., and Fairview Ave., Montgomery, holotype 205901 and para- 

July, 1961 NAUTILUS 33 

types 205902 U.M.M.Z., other paratypes 23392, collection oi the 

Succinea urbana appears to be most closely related to S. wilsoni 
Lea, which it resembles in its genitalia. It differs in its thicker 
shell with smaller aperture and more obese spire. 
(To be concluded) 



The symbols used for Puerto Rican localities were explained 
recently (1961) .^ Very few of the smallest species, especially the 
leaf arboreal ones, which may mature in a few months, were 
obtained, probably because these may be much reduced in num- 
bers or represented only by eggs and/or young during the dry 
periods, because cultivated areas were avoided, and since no 
humus was sifted or saved. All the pupillids are distributed 
widely, probably by human agencies. 

Pupoidcs (s.s.) nitidulus (Pfeiffer) . Enl; lowland species, 
mapped by van der Schalie, 1948:38, from Ps, Wn &: Ws, and 
reported by Dall & Simpson, 1901:372, from Fajardo (Ee) . 

Gastrocopta (s.s.) pellucida (Pfeiffer) . Lowlands, Es2, Ps2. 

G, (s.s.) servilis (Gould). Lowlands, Es2, Ps2, Ws. 

Pupisoma (Ptychopatula) dioscoricola (C. B. Adams). Pnl 
(young) . Two other leaf arboreal Nesopupinae, Bothriopupa 
tenuidens (C. B. Adams) and P. minns Pilsbry, reported by van 
der Schalie, 1948:36, 39. 

Vertigo (s.s.) ovata Say, subspecies? So reported by Pilsbry, 
1919 (3a): 87, pi. 13, fig. 16, from Humacao (Es) ; since he also 
stated "there is nothing to differentiate it," why not drop Pupa 
hexodon Adams, from Jamaica, as obsolete? 

Cerion (Strophiops) striatella (Guerin) + C. crassilabris (Sow- 
erby); see Pilsbry, 1943 (5) :34, but striatella retains its original 
ending. Psl, under bunch grass; Ps2, roosting on tree trunks; 
apparently limited to southwest limestone rim. Shells from 19.0 
mm. long with 814 whorls (Ps2) to 31.1 with 11 (Psl), so C. 
monaense Clench, 1951:274, not always smaller, but does have 
stronger and more widely spaced costae than in any P. R. shells 
seen. Color varying from whitish, through those with brown 
flammules, to almost uniformly brownish. 

Pallial complex of C. striatella (Ps2) and of C. uva (L) from 
just south of Willemstad, Curacao, fundamentally similar to that 
of C. incanum [Pilsbry, 1946(11) :160, fig. 76, G] but much more 

1 In the remarks about Cepolis boriquenae, p. 147, change Ps4 to Es4 
(Humacao) . 

34 NAUTILUS Vol. 75 (1) 

elongate (less contracted?) ; lung wall 5 times as long as kidney or 
7 times its apical width; sometimes with vague, whitish strip 
along hindgut (Cf. Nenin) especially towards pneumostome; 
kidney about 1 14 length of pericardium, with broad base (much 
as in Nenia) , without ureter but with similar subapical opening. 
Mantle collar with weak lappets (similar to those in Nenia) . 

Genitalia of C. strintella very similar to those of C. inconum 
(Pilsbry, figs. H to K); only differences, most of which might be 
due to contraction and/or sexual stage, will be noted. Ovotestis 
of 4 lobes of alveoli, imbedded in 3rd whorl of liver. Carrefour 
also without definite talon. Spermatheca similar but without 
diverticulum in 5 examples dissected; stalk more swollen near 
base; in 2 animals, containing spermatophores, each of which is 
a fragile, thin walled, very slender, horny tube (about as long as 
spermatheca and its stalk) and with, for at least part of its length, 
2 thickenings, that are separated by about \\ its circumference. 
Vagina similarly receiving branches from right ocular retractor, 
which does not pass through penis-vaginal angle. Vas epiphallus 
evidently involving most of free loop and thus about equalling 
spermatophore in length; beginning as gradual enlargement of 
vas lumen near penis-vaginal angle; mainly with thin wall which 
internally has irregularly transverse, lobulate thickenings; attain- 
ing its maximum diameter in descending limb; tapering and with 
wall becoming much thicker and more muscular (at expense of 
lumen) towards entrance into penis near base of last. Penis and 
largely atrial "stimulator" very similar; penial retractor arising 
low on diaphragm and inserting on apex and down along one 
side for about i/l length of caecum. 

Genitalia of C. nva (pi. 3, figs. 3, 4) and C. uva bonaircnsis also 
similar to preceding, but with salient differences as noted. Sperma- 
theca (S) extending to about 2/^ length of spermoviduct; di- 
verticulum (SD) present in those examined, recurved apically but 
not extending above aorta; common stalk (SS) longer and more 
slender but no spermatophores seen (resting?) , about 1 1/^ length 
of free oviduct (UV) . Long vas epiphallus (E) swollen (but with 
smaller lumen) near opening (EP) at bifurcation of large penial 
pilaster. Penis (P) relatively longer, with only I/3 its length above 
vas entrance. Penial apex internally (fig. 3) with heavy longi- 
tudinal folds but basal 2/5 with smooth wall except for pilaster 
and strictly penial "stimulator" (PP) which is similar in outline 
but is simply a discontinuous, elliptic thickening (i.e., without 
free tip). Atrium relatively shorter. 

The differences in at least the penis of C. xiva may represent 
subgeneric characteristics of Cerion s.s., of which it is the only 
species. On the other hand, the presence or absence of a sper- 
mathecal diverticuhmi may be of little significance; Pilsbry stated 

July, 1961 NAUTILUS 35 

it was variable in C. incanum and it evidently is a vestige of the 
long, more primitive kind of geophile spermatheca, in which the 
sac is imbedded above the aorta. Also, the swelling (Pilsbry: fig. I, 
epi) of the long vas epiphallus near its entrance into the penis 
apparently is caused by the contraction of this region with more 
muscular wall but smaller lumen. 

Nenia (s.s.) tridens (Schweigger) , Usually roosting on tree 
trunks or vines up to 8 ft., but rarely on fallen leaves or on rocks; 
Er-1, 2, 3, Es3, Jn I, 100 to 2400 ft., eastern Puerto Rico. Foot quite 
short but broad; sole with 2 or 3 coarse waves on middle zone; 
animal moves along some distance and then jerks shell up after 
it. Embryonic whorls of shell about 2i/^, teat-shaped, relatively 
smooth and without major costulae but assuming at about 14 
whorl close, microscopic, growth wrinkles, which become fine, 
punctulate threadlets by end; 2nd whorl much higher than first; 
later whorls assuming major costulae and crenulate suture very 
quickly; young shells without apertural teeth. 

Pallial complex (pi. 3, fig. 1) fundamentally similar to that of 
Cerion; lung wall about 5 times as long as its apical width or 3 
times length of kidney; similarly without much secondary vena- 
tion, although minute, pigmented tributaries are visible along 
sides of principal vein (HV) ; kidney (K) about twice as long as 
its base or length of pericardium (H) , with similarly subapical, 
external opening (KX) and without ureter. [Although, in some 
examples, discontinuous, whitish zones (mucus or thickening?) 
are visible along hindgut (HG) and kidney, these certainly are 
nothing like the definite thickening along the hindgut in those 
sigmurethrous geophiles (e.g., Oleacinidae) in which the external 
ureteric opening is in the apical corner of the lung. Possibly these 
vague zones were why Wiegmann, 1893:226, described Phaedusa 
as sigmurethrous; this apparently was copied by Thiele in his 
"Handbuch."] Mantle collar (MC) with moderate parietal (left) 
and small, separate palatal (right) lappets. 

Genitalia as represented by P. Hesse, 1925; only additions and 
differences will be noted. Ovotestis (not unlike that of Clausilia 
s. s.; Cf. Steenberg, 1914) consisting of 3-4 conical groups of simple 
and bifurcate alveoli; hermaphroditic duct much swollen and 
convoluted for y^ its length. Carrefour (Steenberg: pf) with very 
short talon (Steenberg: vs) . Sac of spermatheca extending almost 
to tip of diverticulum, which reaches to near apical end of sper- 
moviduct; spermathecal stalk relatively longer than in Hesse's fig. 
(apparent differences may be due to preservation and/or sexual 
stage) . Penis (P. fig. 2) thin walled, with several internal pilas- 
ters, passing gradually (EP) into vas "epiphallus" (Steenberg: 
canal deferent) which has thick, muscular walls and small lumen 
(Cf. Steenberg: fig. 20) and looks more like an ejaculatory organ 

36 NAUTILUS Vol. 75 (1) 

than an epiohallus; penial retractor (PR) arising low on diifram 
and inserting little below middle of "epiphallus" (DE to EP) ; 
base of penis and terminal end of vas proper (up to DE) bound 
by thin sheath. Left ocular retractor in penis-vaginal angle (as 
stated by Hesse) . 

Undoubtedly, the Clausiliidae (at least Nenia) ^ and the Cerii- 
dae (Cerionidae Pilsbry) are closely "related;" although the 
traditional family distinction should be retained, they belong in 
the same superfamily; its name (-oidea or -acea) would come from 
the oldest family, Clausiliidae Moerch, 1864. Both belong in the 
Mesurethra (1955) near the pupilloids (Orthurethra) , in which 
the pallial complex mainly differs by the proximity of the external 
renal opening to the pneumostome; even in this respect, the Par- 
tulidae somewhat approach the Mesurethra. 

Incidentally Thiele in his "Handbuch" retained this position 
for the Clausiliidae and it approximates the traditional location 
of the Ceriidae as well; Cerion uva (L.) is the type species of Pupa 
Lamarck, in which genus most of the older pupillid species were 
described, and the obsolete Pupidae Fleming, 1818, is the oldest 
name for the family. In this case, the resemblances in shell forms 
perhaps are not secondary convergences. As Pilsbry, 1904 (0) : 176, 
suggested, the Megaspiridae probably also belong near the Clau- 
siliidae (and the Ceriidae) . 

Added references (See 1961) 
1955, Naut. ^5:109. 1961, Naut. 74: 142-149. 
Clench, William J. 1951, J. de Conch. PO: 269-276 & pi. 
Hesse, P. 1925, Proc. Malac. Soc, London 7^:154-155. 
Steenberg, C. M. 1914, Mindeskrift, f. J. Steenstrup, no. 29:46 

pp. & pi. 
Wiegmann, F. 1893, ZooL Erg. Niederl. Ost-Indien 2:226. 

1889 to 1961 

Dr. Schwengel was born Jeanne Sanderson, in Stockton, Cali- 
fornia, July 12, 1889. She attended public schools at her birth- 
place, and also St. Xavier's Academy, Chicago, Illinois. To help 

2 Since the above was written, through the generosity of Dr. F. E. Loosjes, 
of Wageningen-Hoog, The Netherlands, animals of Clausilia bidentata 
(Stroem) and Lacinaria plicata (Draparnaiid) have been examined. In both 
of these species, the pallial complex is quite similar to that of Nenia. 

July, 1961 NAUTILUS 37 

her with her conchological studies, she also studied biology and 
Latin at Fordham University, New York City. On January 24, 
1918, she was married to Frank R. Schwengel, who subsequently 
retired from the U. S. Army as Brigadier General, and became 
Chairman of Joseph E. Seagram and Sons, Inc. On February 17, 
1961, while attending the annual meeting of the St. Petersburg 
Shell Club, at which she was scheduled to deliver an address, her 
over-willing heart failed her. She died suddenly, after her return 
to the hotel, where she was staying temporarily. She is buried in 
Scarsdale, where she resided for many years. Her husband survives 

She did many of her studies on mollusks at the Academy of 
Natural Sciences of Philadelphia, where she worked with Dr. 
Pilsbry, and was appointed Research Associate in 1935. In 1943, 
the University of Dayton conferred on her the honorary degree 
of Doctor of Science, in recognition of her contributions to the 
science of conchology. In 1950, the Board of Overseers of Harvard 
College appointed her a member of the visiting committee of the 
Museum of Comparative Zoology. 

In addition to her laboratory studies, she was an indefatiguable 
collector in the field, and helped in much dredging, especially off 
the coasts of Florida. She assisted Dr. Louise M. Perry in the 
compilation of molluscan records for the latter's "Marine shells 
of southwest Florida," 1940, helped revise it, and became co- 
author of "Marine shells of the western coast of Florida." 

Dr. Schwengel was elected Vice-President of the American 
Malacological Union in 1950; became its President in 1951; 
graciously presided over the eighteenth annual meeting at Boston 
in August, 1952; and helped its progress as a member of the 
council for the rest of her life. As many of the members will 
remember, she gave delightful evening parties at many of the 
meetings, before and after. 

She was a member of the following scientific associations: 
Academy of Natural Sciences of Philadelphia (since 1935), Ameri- 
can Malacological Union (1936) , American Association for the 
Advancement of Science (1940), Malacological Society of London 
(1947) , Paleontological Research Institute, Ithaca, N. Y. (1953) , 
Natural Science Foundation, Philadelphia, Pa. (Director from 
1955 to 1958), Philadelphia Shell Club (1955), and Hawaiian 

38 NAUTILUS Vol. 75 (1) 

Malacological Society (1955) . 

She also shared in many other activities. She was a member 
of the American Red Cross in Houston, Texas (1918) , in Chicago, 
Illinois (1920-1932) and in Scarsdale, N. Y. (1935-1945) ; and was 
Chairman of Production from 1943 to 1953. Since 1943, she also 
belonged to the Scarsdale Woman's Club and was Chairman of 
the Garden Section from 1954 to 1956. 

Jeanne, as she was known to her many friends, will be long 
remembered for her ready smile and happy presence. The name of 
Dr. Schwengel will be carried on by her publications about con- 
chology and by her large shell collection, from which she gave 
many lots to various institutions, and left the main part to the 
Museum of Comparative Zoology at Harvard College. — Editors. 

Contributions to conchology 
1938 (1) May 13, Zoological results of the George Vanderbilt 

South Pacific Expedition, 1937. Part I, — Galapagos 

Mollusca. Proc. Acad. Nat. Sci. Philadelphia 90:]-S, 3 figs. 

[Daphnella thalia, Marginella rosa, Tralia x)anderbilti]. 
1938 (2) July 22. Note on unreported marine molluscs from 

Sanibel, Florida, By Louise M. Perry, J. S. S. & Ted Dranga. 

Naut. 52(1) -.27-28. 
1940 (0) Jan. 28. [Cerithium auricoma, Latirus cymatiiis;. See 

1940 (0) ]. Naut. 55 (3) :pl. 12, figs. 6-8a. 
1940 (1) April 29. Two new Floridan marine shells. Naut. 53 

(4): 109-1 10. [See 1940(0).] 

1940 (2) Nov. 2. New Mollosca from Florida. Naut. 54 (2): 49-52, 
pi. 3, figs. 3, 3a, 6-9, 12. [Marginella jaspidea, Crassispira 
phasma, Fenimorea halidorema, Glyphostoma pilsbryi, Bella- 
spira (?) pentapleura.J 

1941 (1) Oct. 24. A genus and family of marine mollusks new to 
the United States. Naut. 55 (2) : 37-40, pi. 3, figs. 1-5. [Lobiger 

1941 (2) Oct. 24. Marginella hartleyana. Naut. 55 (2) :65, pi. 3, 
figs, 6, 7. 

1942 (1) May 7. Living Miira florida. Naut. 55 (4) :144. 

1942 (2) July 23. Some new and interesting marine shells from 
northwest Florida. By J. S. S. k Thomas L. McGinty. Naut. 
56 (1) : 13-18, pi. 3, figs. 2, 3, 5; pi. 4, f. d. [Calliostoma (Eutro- 
chus) faustum, C. fascinans, Douglassia benliana, Trivia 
maltbiana, Aclis hypergonia, Cyclostrema (Aorotrema) pon- 
togenes (n. subg.) ] 

1942 (2a) July 23. [Pteria xnnthia, Phos adelus, Tritiaria vir- 
giniae, Lamellaria leucosphaera.] Naut. 5<5 (l):pl. 3, figs. 1, 
la, 4, 6-9. See 1942(3). 

NAUTILUS 75 (1) 



July, 1961 NAUTILUS 39 

1942 (3) Oct. 14. New Floridan marine mollusks. Naut. 56(2) : 
62-66, pi. 6, figs. 1-3. [Terebra glossema; see also 1942 (2a).] 

1943 (1) Feb. 15. New marine shells from Florida. Naut. 5(5 (3) : 
75-78, pi. 7, figs, 1-7. [Marginella denticulata destina, M. 
idiochila, Eubela mcgintyi, Drupa didyma, Epitonium 
(Cirsostrema) Unteatum.] 

1943 (2) July 23. Diadora jaumei. Naut. 57 (1) :32. 

1944 (1) Feb. 9. Smaragdia viridis viridemaris. Naut. 37 (3) :106. 
1944 (2) Aug. 17. A new Modiolaria from Florida. By R. A. 

McLean and J. S. S. Naut. 58 (1) : 16-17, pi. 1, fig. 10. [Modio- 
laria skomma.] 

1944 (3) Aug. 17. A new Floridian Lamellaria. Naut. 55(1):17- 
18, pi. 1, figs. 3-6, 6a. [Lamellaria koto.] 

1949 (1) Mar. 18. A new Japanese limpet. Naut. 62 (1) :97-98, pi. 
6, figs. 1-lc. [Acmaca langfordi.] 

1949 (2) Mar. 18. Fusinus spectrum, a new record for the Gulf 
of Mexico, Naut. 62 (1): 101-102. 

1949 (3) Nov. 1. Liguus enlarges its menu. Naut. 63 (2) :72. 

1950 (1) Feb. 13. Two Pacific species of Phos. Naut. 63 (3) :80-82, 
pi. 5, figs, 3, 4. [Phos lannumi, P. amoenus.] 

1951 (1) Mar. 7. New marine mollusks from British West Indies 
and Florida keys. Naut. 64 (4) : 1 16-1 19, pi. 8, figs. 1-8. [Crassi- 
spira drangai, Oliva drangai, Pitaria cordata, Calliostoma 
jujubinum adelae.] 

1955 (1) Marine shells of the western coast of Florida. By Louise 
M. Perry and J. S. S. Paleont. Research Inst., 198 pp., 55 pis. 

1955 (2) Aug. 1. New Conns from Costa Rica. Naut. 69 (\) :13- 
15, pi. 2, figs. 1-15. [Conus drangai, C. andrangae, C. gradatus 
thaanumi, C. recurvus helenae.] 

1957 (1) April 29. Theodore Thomas Dranga, 1901-1956. Naut. 
7(? (4): 138- 140, 

1958 (1) Mar. 4. Dr. H. A. Pilsbry in marine malacology. Naut. 
71 (3) : 87-89, 

Dates of the Nautilus. — Vol. 74, no. I, pp. 1-40, pis. 1-4, was 
mailed July 1, 1960, No. 2, pp. 41-84, Oct. 5, 1960. No. 3, pp. 85- 
124, pis. 5-8, Jan 11, 1961. No. 4, pp. 125-166, pis. 9 & 10, April 6, 
1961. — H. B. B. 

First European Malacological Congress. — Under the 
auspices of a committee appointed by the Conchological Society 
of Great Britain and the Malacological Society of London, scien- 
tific and field meetings are planned in London, September 17 to 
21, 1962. All aspects of European malacology and the formation 

40 NAUTILUS Vol. 75 (1) 

of a European Malacological Union will be discussed. However, 
the Congress will be open to all interested in malacology from 
any part of the world. Inquiries should be addressed to the Honor- 
able Secretary: 

Rev. H. E. J. Briggs, 19 Si ward Road, Bromley, Kent, England. 

Directory of conchologists, 1962. — I wish to include all per- 
sons and institutions interested in the study or collection of 
mollusks in this new directory, which will be published next 
January. A copy need not be purchased, but those who remit $2.50 
now will receive the first copies printed. When known, the names 
will be followed by numbers, indicating interests from following 
key (or other special interests will be listed when mentioned): 
1, world wide shells, 2, land shells. 3, fresh water shells. 4, fossil 
shells. 5, exchange shells. Secretaries of shell clubs or institutions 
will please send me a list of their members and their specialties. 
— John Q. Burch, 4206 Halldale Ave., Los Angeles 62, California. 

On the delayed description of Xylophaga atlantica. — The 
wood-boring clam Xylophaga atlantica was described as a new 
species by Richards in 1942 (Nautilus 5(5:58) . Turner subse- 
quently remarked in her monograph on the Pholadidae that, 
"It is surprising that this western Atlantic species remained 
undescribed until 1942" (Johnsonia 5:153. 1955). 

Recently the writer discovered a letter in the file of E. S. 
Morse papers deposited at the Peabody Museum of Salem which 
indicates that this bivalve was recognized as a new species at a 
much earlier date. Following is the letter, published here with 
the permission of its writer. Dr. Henry Jackson, Jr., and the 
Director of the Peabody Museum, Ernest S. Dodge: 

"My dear Mr. Morse, 

I suppose you are very busy, but could you lend me the 
drawings you made of the Xylophaga I found at North Haven. 
It is, without much doubt, a new sp. This I will be able to 
ascertain very soon as Mr. Henshaw has sent to England for all 
the species of the genus. When I describe it I shall like to have 
all the drawings I can of it. I have prepared a very excellent one 
of Xylophaga dorsalis Turton and also one of my [sic] species 
which with your permission I will name Xylophaga [of Morse] 

July, 1961 NAUTILUS 41 

nov. sp. I will be very careful of your drawings if you condescend 
to have them published by me in the Proceedings of the Boston 
Society of Natural History." 

The letter was undated, but in the judgment of Dr. Jackson, 
it was written in 1907 or 1908. In Dr. Turner's monograph 
(ibid-p.l54) , specimens of X. atlantica are listed from "off 
North Haven (Maine) in ten fathoms" which are deposited 
in the U.S. National Museum. These specimens were the ones 
collected by Dr. Jackson in 1907. He does not now recall why 
his study was never published. Possibly because of his age (about 
15 years at that time), his paper was not accepted for publica- 
tion, although he did publish two short papers in the Nautilus 
in 1907 and 1908. Or, it might have been that interest in the 
matter was lost while preparing for a trip to Europe that year. 
Possibly he was discouraged by Dr. Morse from publishing a 
description at that time. Thirty-four years later the situation 
was corrected by Dr. Richards. — Ralph W. Dexter, Kent State 
University, Kent, Ohio. 


Pages in italics include new taxons. 


Adam, William. Les cephalopodes de la mer Rouge. Miss. R. P. 
Dollfus en Egypte. Res. Sci. 3 (28) : 125-193, pis. 1-9, 25 figs. 

Basch, Paul F. Two new molluscan intermediate hosts for Para- 
gonimus kellicotti. J. Parasit. 45:212). Studies on the develop- 
ment and reproduction of the fresh-water limpet, Ferrissia 
shimekii (Pilsbry) . Trans. Amer. Microsc. Soc. 75:260-276. 

Burch, John Bayard. Chromosomes of aquatic pulmonate snails 
(Basommatophora) . Dissertation abstracts 20 (4) : 2 pp. 

Emerson, William K, The gastropod genus Pterorytis. Amer. Mus. 
Novitat. no. 1974, 8 pp., 4 figs. 

Ghose, Krishna Chandra. Observations on the mating and ovi- 
position of two land pulmonates, Achatina fulica Bowdich 
and Macrochlamys indica Godwin-Austen. J. Bombay Nat. 
Hist. Soc. 55:183-187. 

Schalie, Henry van der ^ Dee S. Dundee. Transect distribution of 
eggs of Pomatiopsis lapidaria Say, an amphibious proso- 
branch snail. Trans. Microsc. Soc. 75:409-420, 5 figs. 


Adam, William. Cephalopoda from the Gulf of Aqaba. Israel 
Sea Fish. Res. Sta., Bui. 26:26 pp., pi. 1, 10 figs. Contribution 

42 NAUTILUS Vol. 75 (1) 

a la connaissance de I'hectocotyle chez les Ommastrephidae. 
Bui. Inst. roy. Sci. nat. Belgique 36 (\9) :10 pp. A propos de 
Chlamys (F.Acc. Hinnites) abscondita (P. Fischer, 1898) de 
la cote occidentale de I'Afrique. Ibid. (20) :10 pp., 2 pis. Les 
mollusques teiTestres et dulcicoles de la Belgique. Quelques 
additions et rectifications. Ibid. (22): 10 pp., 7 figs. Les 
c^phalopodes de I'lnstitut Francais d'Afrique Noire, 2. Bui. 
ibid, 22:465-511, 4 figs. 

Barbosa, Frederico S., Elizabeth Carneiro, Ivete Barbosa & Jose F. 
Magalhaes. Manual de malacologia medica (Trabalhos 
praticos) . Pp. 182, 110 figs. Fundacao Goncalo Moniz, Salva- 
dor, Bahia, Brasil. 

Branson, Bradley A. Gastropoda of the Rob and Bessie Welder 
Wildlife Foundation Refuge, San Patricio County, Texas. 
Southwest. Nat, 5:143-159. 

Burch, John B. Chromosomes of Gyraulus circumstriatus, a fresh- 
water snail. Nature 75<5:497-498, 1 fig. 

Cooke, C. Montague, Jr. & Yoshio Kondo. Revision of Tornatelli- 
nidae and Achatinellidae (Gastropoda, Pulmonata). Bui. 
Bishop Mus. 221:1-303, 123 figs, ($6.50). 

Emerson, William K. Remarks on some eastern Pacific muricid 
gastropods. Amer. Mus. Novitat. no. 2009:15 pp., 7 figs. 

Hubricht, JLeslie. The cave snail, Carycliium stys^ium CalL Trans. 
Ky. Acad. Sci. 27:35-38, 2 figs. 

Kuroda, Tokubei. A catalogue of molluscan fauna of the Okinawa 
Islands. Pp. 106, 3 pis. 

Leonard, A. Byron & John C, Frye. Wisconsinan molluscan 
faunas of the Illinois valley region. 111. State Geol. Surv., 
Circular 304: 32 pp., 4 pis., 3 figs. 

Morton, J. E. Molluscs: an introduction to their form and func- 
tions. Harper Torchbooks, The Science Library; 232 pp., 23 
figs. (11.40). 

Paraense, W. Lobato & Newton Deslandes. " Drepanotrema sur- 
inamense," with an addendum on "D. petricola" (Plan- 
orbidae) . Rev. Brasil. Biol. 20:257-263, 7 figs. 

Schalie, Henry van der. Egypt's new high dam — asiset or liability. 
Biologist ^2:63-70, 3 maps. 

Scott, M. I. Hylton. Nueva familia de pulmonado Basomatoforo, 
Neotropico 6:65-69, fig. 1, Sobre la presencia del genero 
Pupisoma en la Argentina. Ibid.: 25-29, figs. 1-9. 

Smith, Allyn G. A new species of Megomphix from California. 
Occ. Papers Calif. Acad. Sci. no. 28:5 pp., 3 figs. 

Soot-Ryen, T. Pelecypods from Tristan da Cunha. Results 
Norweg. Sci. Exp. to Tristan da Cunha, no. 49:'^7 pp. (3 
pis.) , 9 figs. 

Voss, Gilbert L, Bermudan cephalopods. Fieldiana, Zoology 
39:419-446, figs. 73-75. 


Vol. 75 October, 1961 No. 2 




In the summers of 1958 and 1959, studies on the Pleistocene 
geology and molluscan faunas of the White (Blanco) River area 
in northwestern Texas were carried out by us. In the course of 
these studies, numerous fossil shells of Strobilops of problematical 
taxonomic status were found associated with other pulmonate 
mollusks in fluviatile deposits ranging in age from Kansan to 
Wisconsinan. About 200 specimens of Strobilops have been ob- 
tained from the White River area. We were unable to assign 
these shells to any known species owing to their consistent char- 
acteristics intermediate between extinct S. sparsicostata F. C. 
Baker (1938:127) and living S. texasiana Pilsbry and F'erriss 
(Pilsbry, 1948:856) . For these reasons and because of their wide 
geographic distribution and stratigraphic range, it seems best to 
propose a new name for these fossil shells. Series of several 
Pleistocene Strobilops from the post-Nebraskan of Central High 
Plains were restudied. Those series of Strobilops, which had 
been confused with S. sparsicostata, appear to belong to the new 
species described here. 
Strobilops loxsdalei, new species. Plate 6, figs. 1-2; text figs. 1-2 

Diagnosis: Shell large for Strobilops, having trochiform out- 
line; decidedly angulate last whorl; SI/q to 6 convex whorls sculp- 
tured by 40 to 50 moderately spaced, moderately thickened, 
oblique ribs passing over base without diminishing; weak and 
nodose infraparietal lamella. 

Holotype: Catalogue number 12180, University of Kansas 
Museum of Natural History, obtained by A. B. Leonard and 
Tong-yun Ho, June 9, 1959. Original number TYH 34. 

Dcscriptioji of Jwlotype: Shell large, trochiform; whorls 6 in 
number, moderately convex, slowly increasing in size; last whorl 
decidedly angulate; base broadly rounded; first W/o whorls finely 
granulose, remaining whorls covered by moderately spaced, 
moderately thickened ribs passing over base Avithout reduction in 
size, except for those immediately in front of aperture; diameter 


44 NAUTILUS Vol. 75 (2) 

of umbilicus equivalent to i/g greater diameter of shell; aperture 
lunate, expanded, having moderately thick parietal callus and 
peristome; parietal lamella nodose, high, emerging to edge of 
parietal callus and penetrating about % of a whorl inward; inter- 
parietal lamella nodose, weak, shortly emerged and penetrating 
as deep as parietal lamella; infraparietal lamella nodose, weak, 
shortly emergent and penetrating as far as parietal lamella; col- 
umellar lamella weak; first basal fold moderate, triangular; sec- 
ond basal fold high, broadly triangular, nearly as long as first 
basal fold; third basal fold low, about i/^ length of second basal 
fold; fourth and fifth basal folds low, each approximately twice 
length of third basal fold; palatal fold low, longer than fifth basal 
fold (internal anatomy from paratypical shell) . 

Paratypes: Many variations occur among more than 200 para- 
types in the length, thickness, and number of baso-palatal folds, 
and in the development of ribs. The baso-palatal folds in most 
shells are 5 in number but in a few specimens a small fold is pres- 
ent between the third and fourth basal folds. The inter- and 
infra-parietal lamellae are extremely weak and may become dis- 
continuous in some examples. The number of ribs ranges from 
40 to 50. In some examples, a faint riblet is between major ribs. 
The ribs on the base may become weak on the last half of the 
base or near the umbilicus in some shells; as a rule they are 
conspicuous on last half of base. 

Comparisons: S. lonsdalei is clearly distinguished from S. 
sparsicostata (Plate 6, figs. 5-6; text figs. 4-5) by having ribbed 
base, weak nodose infraparietal lamella, and fine and narrowly 
spaced oblique ribs, always more than forty in number. The 
denticles as a group are always weaker than are those of S. 
sparsicostata. The parietal lamellae are nodose in lonsdalei, but 
most of them are always smooth in sparsicostata. The palatal 
fold is generally present in lonsdalei, but it is absent in some 
specimens of sparsicostata. 

The shell of S. lonsdalei differs from that of S. texasiana (see 
Pilsbry, 1948, fig. 464:5-11) in being large and trochiform instead 
of small and dome-shaped. The last whorl is subangulate or 
rounded in texasiana, whereas it is decidedly angulate in lons- 
dalei. The ribs, which are always more than 50 in number in 
texasiana, are always less than 50 in lonsdalei. The ribs are also 
more widely spaced and coarser in lonsdalei than in texasiana. 
The internal structure of the shell of lonsdalei is also different 

October, 1961 nautilus 45 

from that of texasiana. The latter has a tongue-shaped (blunt- 
topped) first basal fold, and high, rather flat-topped second 
basal fold, while in the former both the first and second basal 
folds are triangular in shape with subacuate tips. Unlike tex- 
asiana, lonsdalei has a low second basal fold and intraparietal 

The comparisons of dimensions in millimeters and numbers of 
whorls are summarized in the following table. In the table, speci- 
men HI is the holotype, and specimens PI and P3 are paratypes. 

Species S. sparslcostata S. lonsdalei S. texasiana 

Specimen 12 3 4 HI PI P2 P5 12 3 4 

Diameter 2.7 2.7 2.6 2.5 2.8 2.8 2.7 2.7 2.4 2.3 2.2 2.0 

Height 1.9 2.0 2.1 1.9 2.4 2.5 2.3 2.1 1.9 1.9 1.6 1.6 

No. of ribs 34 36 36 34 41 44 48 45 52 50 52 51 

No. of 5.5 5.5 5.5 5.5 6 6 5 .9 5 .9 5.5 5.5 5.3 5.5 


Type locality: Kansan terrace deposits in left bank of White 
River, near bridge on Texas Highway 261, 6.5 miles east of 
Kalgary, Crosby County, Texas. 

Areal and stratigraphic distributions: At present S. lonsdalei is 
known from the Pleistocene deposits of Kansan to Wisconsinan 
age at 13 localities distributed from western Oklahoma to 
northern Texas. Representative localities, age of deposits, and 
approximate number of specimens obtained at each locality are 
as follows: 

Kansan deposits, NW 14, sec. 23, T. 23N., R. 18W., 2.5 miles 
west and % mile north of Ouinlan, Woodward County, Okla- 
homa. No. 4337, KU, 70 specimens. 

Kansan deposits in road cut on Texas Highway 70, 1 mile 
north-northeast of Turkey, Hall County, Texas. No. 1 1818, KU, 
10 specimens. 

Kansan deposits in left bank of White River, near bridge on 
Texas Highway 261, 6.5 miles east of Kalgary, Crosby County, 
Texas. No. 11650, KU, 60 specimens. (Type locality.) 

Kansan deposits exposed in cut bank, 2.2 miles west of Post, 
Garza County, Texas. No. 10854, KU, 40 specimens. 

Wisconsinan terrace deposits on north side of U.S. Highway 83, 
4.1 miles east of Crosbyton, Crosby County, Texas. No. 10237, 
KU, 150 specimens. 

Wisconsinan terrace deposits, 5 miles northeast of bridge on 

46 NAUTILUS Vol. 75 (2) 

U.S. Highway 77 over Red River, northern central Cook County, 
Texas. No. 12123, KU, 10 specimens. 

Strobilops lonsdalei cansasiana, new subspecies. Plate 6, figs. 
3-4; text fig. 3. 

Diagnosis: Shell broadly conic, moderately elevated; whorls 
5i/2 in number sculptured by 43 to 52 moderately spaced ribs; 
base almost smooth or finely striate; parietal lamellae penetrating 
from 1/2 to Ys of last whorl. 

Holotype: Catalogue number 12181, KU, obtained by Dr. C. 
W. Hibbard, August 9, 1943. 

Description of Holotype: Shell moderate, broadly convex; 
whorls 51/2 in number, convex; last whorl decidedly angulate; 
base more or less narrowly rounded; first I14 whorls finely 
granulose, remaining whorls sculptured by moderately spaced, 
moderately thickened ribs passing over base in extremely fine 
striae; diameter of umbilicus contained about 8 times in diameter 
of shell; aperture lunate, expanded, having moderate peristome 
and parietal callus; parietal lamella nodose, penetrating one- 
half a whorl inward; interparietal lamella nodose, weak, shortly 
emerged and penetrating as deep as parietal lamella; infraparietal 
lamella shortly emerged, longer than interparietal lamella, pen- 
etrating as far as parietal lamella; baso-palatal folds similar to 
those of S. lonsdalei lonsdalei (internal structure of shell based 
on paratype) . 

Paratypes: Few variations are noted in more than 2000 para- 
types from 8 localities. The parietal lamellae pentrating 1/9 last 
whorl inward in most specimens, but about 5/^ last whorl in the 
specimens from Reno County, Kansas, and Greer County, Okla- 
homa. The base is smooth or finely striate in most examples, 
but a few ribs are present immediately behind the peristome 
of a few specimens. The baso-parietal folds are as variable as 
those of S. lonsdalei lonsdalei. Variations in size of shell, and 
number of ribs are exemplified in the following measurements: 

Holotype Paratypes 

Diameter (mm.) 3.1 2.6 2.7 3.1 

Height (mm.) 2.2 2.1 2.0 2.1 

No. of ribs 45 52 44 43 

No. of whorls 5.5 5.5 5.5 5.5 

Comparisons: This subspecies differs from Strobilops lonsdalei 
lonsdalei in having smaller shell with broad and smooth base, 
less elevated spire, more ribs, and shorter parietal lamellae. All 
other characteristics appear to be similar to S. lonsdalei lonsdalei. 
S. lonsdalei cansasiana resembles Strobilops texasiana in having 
the same number of ribs, but the former differs from the latter 

October, 19(51 
NAU riLUS 75 (2) 




Kigs. 1, 2. Strobilops lonsdalei lonsdalei Ho and Leonard. Lateral and basal 
\ iews of holot\ pica] shell. Figs. 3, 4. S. lonsdalei cansasiana Ho and Leonard. 
Lateral and basal views of holotypical shell. Figs. 5, 6. S. sparsicostata Baker, 
Lateral and basal views of topotypical shell from Rexroad Ranch, 9 miles 
south and 7 miles west of Meade, Meade C'oiintv, Kansas. All rigurcs enlarged 
approximately 15 times. 



Vol. 75 (2) 






Sections of last whorl of Sirobilops showing internal lamellae antl toicls ol 
ihe shell. All fie;iires enlarged approximately 14 times. Figs. 1, 2. Strobilops 
loiisdah'i lousdalci Ho and Leonard, paratvpes. Fig. 3. .S. lousdalei cansasiana 
Ho and Leonard, paratype. Figs. 4, .5. S. sparsicoslata Raker, lopotvpes. 

in bearing rather coarse and widely sj^ared ribs, smooth base, 
and a decidedly angulate last whorl. 

Type locality: Kansan deposits, SW 14, sec. 2, T. .81,S, R. 28W., 
6 miles north of Meade, Meade County, Kansas. 

Area! and strntigraphic distributions: S. lousdalei ca)isasiaua 
is now known only Irom Kansan deposits in Iowa, Kansas and 
Okhdioma. The southern range of S. lousdalei causasiaua over- 
laps witli the noiithern range of .S. lousdalei lousdalei in western 
Oklahoma. Representative localities and apj^roximate number 
of specimens in otir collections are: 

Kansan deposits, I mile east and 4 miles north ol Little Sioux, 
Harrison County, Iowa. No. 4517, KU, 1 1 specimens. 

Kansan deposits, SW 14, sec. 2, T. .8 IS., R. 28W., 6 miles north 
of Meade, Meade County, Kansas. No. ,8702, KV , 500 sjiecimens. 
(Tyj)e locality.) 

Kansan deposits, SW i/, , sec. 20, T. I IS., R. 2i:., I miles west 
of Navarre, Dickenson County, Kansas. No. l.^^iOl, Kl', 10 speci- 

October, 19(5 1 nautilus 49 

Kansan deposits, sec. 36, T. 24S., R. 7W., 7 miles east and 
1.1 miles north of Arlington, Reno County, Kansas. No. 9414, 
KU, 15 specimens. 

Kansan deposits, sec. 8, T. 5N., R. 28E, neaj- north border of 

Gate, Beaver County, Oklahoma. No. 4743, KU, 150 specimens. 

Kansan deposits in right bank of North Fork of Red River, 

north of Rock Island Railroad, 2 miles east of Granite, Greer 

County, Oklahoma. No. 10751, KU, 70 specimens. 

Ecology and relationships: The ecological requirements of this 
extinct species are not certainly known. The association of it 
with Retinella electrina, Pupilla muscorum, Stenotrema leai, and 
its occurrence in the alluvium of fine and well-sorted sands and 
silts seems to imply that the species thrived in rather humid and 
cool situations on the flood plains where vegetation was available. 
The exact cause of extinction is unknown, but the initiation of 
semiarid and otherwise severe climate in the High Plains at the 
time of the Bradyan interglacial interval may have been respon- 
sible. This view is strengthened by the fact that no single species 
of Strobilops and no other northern gastropod that lived in the 
area during pre-Wisconsinan time, is found alive in the White 
River region today. 

S. lonsdalei is morphologically and stratigraphically inter- 
mediate between 5. sparsicostata of the Nebraskan and existing 
S. texasiana, which extends to the Wisconsinan, but is more 
closely allied to 5. sparsicostata than to 5. texasiana on the basis 
of size and shape of shell. In the light of our present knowledge, 
5. lonsdalei seems to be on a side branch of the line leading from 
S. sparsicostata to 5. texasiana. S. sparsicostata may have given 
rise to S. lonsdalei by development of ribs on the base of the 
shell, by increase in the number of ribs, and by reduction in size 
of denticles. 

S. lonsdalei is named in honor of the late Dr. John T. Lonsdale, 
formerly Director, The University of Texas Bureau of Economic 
Geology, in recognition of his continued interest and support of 
our studies in Texas. 

Literature cited 
Baker, Frank C. 1938. New land and freshwater mollusca from 

the upper Pliocene of Kansas and new species of Gyraulus 

from early Pleistocene strata. Naut. 5/ (4) : 126-131. 
Pilsbry, Henry A. 1948. Land Mollusca of North America (north 

of Mexico) . Acad. Nat. Sci. Philadelphia Monographs no. 3, 

2(2):856-858, fig. 464:5-11. 

50 NAUTILUS Vol. 75 (2) 




Marine Biological Laboratory, Helsingor, Denmark 

While working up the marine lamelHbranchs of East Green- 
land (Ockelmann, 1959) a list was compiled of all the species 
known from the Atlantic sector of the Arctic and adjacent re- 
gions and, in many cases, I had to consult the original descrip- 
tions. It thereby appeared that the validity of some of the species 
of Thyasira, described by Verill & Bush (1898) from the east 
coast of North America, was rather doubtful and that they prob- 
ably were synonymous with certain earlier described species. In 
particular, this seemed to apply to Thyasira insignis, T. plana, 
and T. inaequalis. 

During a stay in the United States in 1959, I therefore took 
the opportunity to examine a major part of the collection of 
Thyasira kept in the U.S. Nat. Museum. This material includes 
the types as well as most of the specimens of Thyasira identified 
by Verrill & Bush. To facilitate a later thorough revision of the 
whole group, which is highly needed in spite of the useful papers 
by Dall (1901) , and Lamy (1920) , the results of my examination 
will be given here. 

Thyasira insignis: This form was described as Cryptodon 
insignis by Verrill & Bush (1898, p. 785, pi. 91, figs. 1, 2). Two 
specimens were figured and, apparently, the authors did not 
designate a holotype. However, one of these specimens is now 
in the type collection of the U.S. Nat. Museum, probably selected 
by Dall when he prepared his 'Synopsis of the Lucinacea and of 
the American species' (1901). It is an empty shell (U.S.N.M. 
52596) with the left valve drilled. This is depicted on fig. 2 in 
the paper by Verrill Sc Bush, while fig. 1, same plate, is drawn 
from a single left, drilled valve (U.S.N.M. 52733) . Both speci- 
mens are from U.S.F.C. station 2499, N.44°46'30'' W.59°55'45", 
depth 130 fath. ('Albatross', 1885). Further, 28 valves (U.S.N.M. 
52557 and 52733) from stations of the U.S.F.C. 2498 and 2499, 
were present. Of the whole material, consisting of 1 shell and 29 
valves, 11 still had remains of the ligament, but no trace of the 
soft parts whatever. Thus, tliey may vei7 well have rested in the 
bottom deposits for a considerable time after death. A single 

October, 1961 nautilus 51 

valve from off Cape Cod which at first was identified as Crypto- 
don sarsii by Verrill (1880, p. 399), but later (Verrill k Bush, 
1898) referred to C. insignis, seems not to be present in the 
material. According to the latest lists of the marine mollusks of 
the east coast of North America, T. insignis occurs from New- 
foundland to Cape Cod, Mass., 65-471 fms. (Johnson, 1934; La 
Rocque, 1953) . This distribution, however, is simply based on 
the data of Verrill & Bush (1898) whose material solely consists 
of empty shells and valves. From this, apparently T. insignis has 
never been' taken alive. 

My examination of the type as well as the other material 
strongly suggests that T. insignis is not specifically distinct from 
T. sarsi (Philippi) 1845, b, p. 91, as Axinus sarsii). The outline 
of the shell, its proportions, the appearance of its surface, the 
hinge margin, the size and the position of the ligament, as well 
as the scars and marks on the inner side do not differ in the two 
forms. The only observable differences are that, in some of the 
valves, the lower, posterior undulation from the umbo to the 
posterior-basal maigin ends a little more ventrally than in most 
of the specimens of T. sarsi, and that T. insignis may become 32 
mm. long, while the largest specimens of T. sai'si (valves from 
W. Norway) seen by me have a length of 23.5 mm. However, 
T. sarsi varies rather much in shape, as do several other species 
of Thyasira, and it also seems to attain fairly different maximum 
sizes in different localities. G. O. Sars, who gives good figures of 
the typical form (1878, pi. 19, fig. 5 a-b) , shortly describes a 
'monstr. oblonga' collected along with typical specimens at 
Lofoten. This form especially is virtually indistinguishable from 
T. insignis. 

According to my own studies, T. sarsi now occurs from the 
Sound (Denmark) through the Kattegat, along the Swedish 
west coast and the Norwegian coast northward to at least Lofoten, 
and it is also found in some fjords of northwest Icelantl. G. O. 
Sars (1878), however, states that it occurs as far northward as 
Vadso (E. Finmark) , and this may be correct, although I have 
not seen animals from that region. It has even been recorded 
farther north and east by other authors. Lamy (1920, p. 297), 
for instance, mentions it from the Kara Sea. However, since 
specimens of T. gouldi (Philippi) often have been identified 

52 NAUTILUS Vol. 75 (2) 

and listed as T. sarsi, the northern limit of distribution is not 
known with certainty. T. sarsi has also been recorded from W. 
Greenland (Posselt & Jensen, 1898, p. 80, as Axinus flexuosus var. 
sarsii). The Zool. Museum, Copenhagen, however, only possesses 
a single, empty shell from W, Greenland, viz. from Godthaab. 
The ligament of this shell is fairly well preserved, but soft parts 
are completely absent and the shell surface does not look 
'fresh'. There are thus three possibilities: 1) This specimen is a 
quaternary fossil, and T. sarsi is now extinct at W. Greenland, 
2) The specimen is wrongly labelled, and not collected at W. 
Greenland at all, or 3) T. sarsi occurs, although sparsely, in 
west Greenland water. At present, none of these interpretations 
can be excluded. 

From the preceding data, I conclude that T. insignis merely 
represents a large form of T. sarsi, and that the former should 
be regarded as a subspecies of the latter and referred to as 
Thyasira sarsi insignis (Verrill &: Bush) . Further, no proof 
exists that this is an extant member of the recent fauna of the 
east coast of N. America, since all the specimens referred to it 
may very well be Quaternary fossils, thus indicating a wider dis- 
tribution of T. sarsi in former times, viz. from the Scandinavian 
coasts to the region around Newfoundland and New England. 
This supposition also agrees with the record of an empty, pos- 
sibly fossil, shell of T. sarsi from W. Greenland, and with the 
existing isolated population of N.W. Iceland. 

Thyasira plana: This was described, with good figures, as 
Cryptodon planus by Verrill & Bush (1898, p. 788-9, pi. LXXX- 
VIII, figs. 3, 4). The type (U. S. N. M. 159893) is a specimen from 
U.S.F.C. sta. 254, Cape Cod Bay, Tishing ledge'. Wood End Light 
N. 50°E. 7 miles, 21 fath. ('Speedwell', 1879). A close examin- 
ation of it left no doubt that it is an about half-grown specimen if 
Thyasira gouldi (Philippi) (1845 a, p. 74-75). It has a well- 
rounded outline which is often exhibited by smaller specimens 
of T. gouldi. Its hinge-margin is comparatively strong and 
fairly well-rounded below the umbo, a feature also found in T. 
gouldi. Furthermore, Verrill & Bush's statement as to the ab- 
sence of a distinct tubercle or tooth-like projection is not quite 
correct. There is, just as in T. gouldi, a rather stout, though 
small, tooth in the right hinge close to, and below, the umbo, 

October, 19bl nautilus 53 

and — as also shown by Venill &: Bush, fig. 3 — a distinct pit or 
groove in the left hinge corresponding to it. A highly distinctive 
character of T. goiildi is the large, conspicuous prodissoconch 
wliich most often measures 210-250 microns across. This feature 
is also present in the type of T. plana, the prodissoconch of 
which measures about 215 microns across. Finally, all the other 
material investigated and originally labelled as Cryptodon 
planus also turned out to belong to T. gouldi, mostly being 
smaller specimens of it. This material originates from the follow- 
ing localities: Casco Bay, 1873. Seal Cove, 8-10 fms., Y. M., 1872, 
Chelsea Beach, Mass., Dall. Friars Head, Maine, Henderson Coll. 
U.S.F.C. stations 72+73, 42B+43B + 58B + 60B-f 66B-f 67B, 134, 
136, 140, IGO, 165, 181, 191, 199, 210, 283+285, 296, and 340 
(serial numbers U.S.N.M. 74281, 74283-74286, 74289, 108868, 
159848, 159849, 159876-159884, and 445797). There is also one 
sample from Maine, collected by Stearns, but without a serial 
nmnber. Living specimens, identified as Cryptodon planus, weie 
found at depth from 15-19m. to 188 m. and at temperatures 
between about 4.0° and 9.4°C. (These data are based upon 
Smith, 1887, according to the station numbers of the samples). 
The distribution agrees with that given by Johnson (1934) , and 
La Rocque (1953) , viz. from Halifax, N.S., to Cape Cod, Mass., 
and, according to these authors, the range of T. gouldi along the 
east coast of N. America is from Greenland to Connecticut, 5-400 
fath. Abbott (1954, p. 384) , however, gives the range as "Labra- 
dor to North Carolina," but without any references or details. 
T. plana thus occurs well within the range of T. gouldi, regard- 
less of the depth record of 400 fath. for T. gouldi which seems 
doubtful and may not apply to living specimens. Actually, T. 
gouldi is uncommon at depth exceeding 100 m. and even at 
depths of about 50 m. often only smallei" specimens are found. 
These are usually more rotund than are the larger ones; and 
apparently such specimens led Verrill &: Bush to describe Crypto- 
don planus. 

Thyasira inaequalis: This was described as Cryptodon (Ax- 
inulus) inequalis by Verrill & Bush (1898, p. 791, pi. 90, figs. 1, 
2). Dall (1901, p. 786) is correct in regarding it as a Thyasira 
S.S., not an Axinulus. The type (U.S.N.M. 159850) is an animal 
from U.S.F.C. sta. 98, 99, midway between Sandwich Point and 

54 NAUTILUS Vol. 75 (2) 

McNab's Island Light, Halifax Harbor, depths of 1 8 fms. (Speed- 
well, 1877). On a close inspection, it turned out to be nothing 
but an aberrant specimen of T. gouldi, aberrant in so far as the 
outline of the shell is concerned. Otherwise it shows all the 
features typical of T. gouldi; the prodissoconch, for instance, 
measures about 215 microns across. Also the other material in- 
vestigated (U.S.F.C. stations 72-f 73, 293, 296, and 340 (U.S.N. M. 
74287, 159847, 202836, and 202837) comprises only specimens of 
T. gouldi, but of a more normal outline and shape. This again 
points towards the true nature of the type: An individual variant 
of T. gouldi. Just as is the case with T. plana, T. inaequalis is 
sympatric with T. gouldi, as it occurs from Halifax, N.S., to 
Cape Cod, Mass., 14 to 49 fath. (Johnson, 1934; La Rocque, 

It is concluded, then, that both Thyasira plana (Verrill & 
Bush) , and T. inaequalis (Verrill & Bush) must be regarded as 
junior synonyms of T. gouldi (Philippi). The available data do 
not justify a claim for even their subspecific status. 

Most figures of T. gouldi in the literature are on a small scale 
and do not show the distinctive features satisfactorily. For con- 
venience, therefore, references to usable figures may be given 
here: G. O. Sars, 1878, pi. 19, fig. 6 a-b (as Axinus gouldii), 
and Ockelmann, 1959, pi. 2, figs. 4-5. 

My cordial thanks are due to Dr. Harold A. Rehder for 
kindly placing the collections and facilities of the U.S. National 
Museum at my disposal. 

A few living adult T. sarsi have been found by me since, in the Gotthab 
Fjord, W. Greenland, July, 1961. 

Literature cited 
Abbott, R. T. 1954. American seashells. Van Nostrand, New York. 
Ball, W. H. 1901. Proc. U.S. nat. Mus. 25(1237) :779-833. 
Johnson, C. W. 1934 List of Marine Mollusca of the Atlantic 

Coast from Labrador to Texas. Proc. Boston Soc. Nat. Hist. 

-/O(l): 1-204. 
Lamy, E. 1920. J. de Conchyliol. ^5:233-318 (3e Part.). 
La Rocque, A. 1953. Catalogue of the recent Mollusca of Canada. 

Bull. nat. Mus. Ottawa 129 (Biol. ser. no. 44) : 1-406. 
Ockelmann, K. W. 1959. Marine Lamellibranchiata. (In: The 

Zoology of East Greenland.) Medd. om Gronland 122 (4) : 

Philippi, R. A. 1845a. Zeitschr. f. Malakozool. 2:68-79. 
Philippi, R. A. 1845b. Zeitschr. L Malakozool. 2:87-91. 

October, 19()1 nautilus 55 

Possclt, H. J. S: Jensen, A. S, 1898. Gronlands Brachiopoder og 

Bloddyr. (In: Conspectus Faunae Groenlandicae) . Medd. cm 

Gronland 25; I-XIX, 1-298. 
Sars, G. (). 1878. Bidrag til Kundskaben om Norges Arktiske 

Fauna. I. Mollusca regionis arcticae Norvegiae. Christiania. 
Smith, S, 1889. Rep. U.S. Comm. Fish and' Fisheries for 1886 

(part XIV): 871-1017. 
Verrill, A. E. 1880. Proc. U. S. nat. Mus. 5:356-405. 
Verrill, A. E. R: Bush, K. J. 1898. Proc. U.S. nat. Mus. 20 (1 139) : 




Watei" Island is located about y^ of a mile east of St. Thomas. 
It consists mainly of volcanic rocks which rise to a height of 294 
feet at the highest point, leaving a very rough topography. The 
water aurrounding the island varies from shallow to a depth of 
50 to 85 feet in the channels. The bays, sand beaches, rock ledges 
and coral reefs furnish a great variety of ecological situations for 
mollusks. Collecting is very convenient, as the proprietor of the 
island, Mr. Walther H. Phillips, maintains tourist cottages, 
where wet clothes and specimens may be handled. The writer 
made two trips to the island, one in August, 1956, and the other 
in July, 1958, which included shallow-water collecting and some 
dredging. The species collected follow: 

Gnstropods. Arene cruentata 

Acmaea antillarum A. miniata 

A. cubensis A. riisei 

A. leucopleura Aspella elizabethae 

A. jamaicensis A. paupercula 

A. pustvdata Astraea americana cubana 

A. pustulata pulcherrima A, caelata 

Acteon punctostriata A. longispina 

Alaba incerta A. tuber 

Alabina adamsi Atys caribaea 

Alvania auberi A. sharpi 

Anachis albella Batillaria minima 

A. catenata Bittium cerithioides 

A. nitens B. varium 

A. obesa Bulla occidentalis 

A. subcostulata Bullata ovuliformis 

Antillophos candei Bursa thomae 

Architectonica nobilis Caecum cooperi 



Vol. 75 (2) 

C. decussatum 

C. regulare 

C. floridanvim 

C. pulchellum 

Calliostoma jujubinum 

C. pulcher 

C. zonamestum 

C. sarcodum 

Cantharus auritulus 

Cassis flammea 

C. madagascariensis 

C. tuberosa 

Cerithides costata 

Cerithiopsis iota 

C. flavum 

C. fusiforme 

C. greeni 

C. rugulosum 

Cerithium algicola 

C. eburneum 

C. literatum 

C. muscarum 

C. variabile 

Cerodrillia thea 

Charonia variegata 

Coliimbraria lanceolata 

Columbella mercatoria 

C. ovulata 

C. ovuloides 

Conus spurius 

C. jaspideus 

C. mus 

C. regius 

C. cardinalis 

C. dominicanus 

Coralliophila abbreviata 

C. bracteata 

C. caribaea 

Crassispira ebenia 

C. nigrescens 

C. ostrearum 

Crepidula aculeata 

C. maculosa 

C. plana 

Cyliiidrobulla beaui 

Cymatium gemmatum 

C. muricium 
C. nicobairium 
C. parthenopium 
C. pileare 

Cyphoma gibbosum 
C. macgintyi 
Cypraea cinerea 
C. spurca acicularis 

C. zebra 

Cypraecassis testiculus 
Cythara trilineata 
Daphnella lymneiformis 
Decipifus pulchellus 
Diodora arcuata 

D. cayenensis 
D. dysoni 

D. minuta 
Drillia elatoir 
Drupa nodulosus 
Echininus nodulosus 
Emarginula phrixodes 

E. pumila 
Engina turbinellus 
Engoniophos unicinctus 
Epitonium albidum 

E. candianum 
E. echinaticostum 
E, lamellosum 
E. occidentale 

E. unifasciatum 
Erato maugeiiae 
Euchelus guttarosea 
Fasciolaria tulipa 
Fenimorea fucata 

F. phasma 
Fissurella augusta 
F. barbadensis 

F. barbouri 
F. nimbosa 
F. fascicularis 
F. nodosa 
F. rosea 

Fossarus orbignyi 
Fusilatirus cayohuesonicus 
Glyphotunis diminuta 
Haliotinella patinaria 

October, 1961 



Haminoea antillarum 
H. elegans 

Hemitonia emarginata 
H. octo-radiata 
Hipponix antiquatiis 
H, submfus 
Hyalina albolineata 
H. avena 
H. avencia 
H. tenuilabra 
H. parkeri 

Ithycythara lanceolata 
Jaspidella jaspidea 
Kinziella (juadrilineata 
Latirus brevicaudatus 
Leucozonia nassa 
L. leucozonalis 
Litiopa melanostoma 
Littorina angulifera 
L. meleagiis 
L. mespillum 
L. ziczac 
Livona pica 
Lucapina aegis 
L. philippiana 
L. sowerbyi 
L. sufftisa 

Lucapinella limatula 
Mangelia biconica 
M. fusca 
M. melantica 
M. quadrilineata 
Marginella sulcata 
Meioceras nitidum 
Melampus coffeus 
Melanella jamaicensis 
Micromelo undata 
Mitra albicostata 
M. albocincta 
M. barbadensis 
M. cubana 
M. floridana 
M. hanleyi 
M. hanleyi gemmata 
M. moisei 
M. nodulosa 

M. straininea 

Mitrella duclosiana 

M, fusilormis 

Modulus carchedonius 

M. modulus 

Monilispira albocincta 

M. jayana 

M. leucocyma 

Morum oniscus 

Murex cabritti 

M. cailleti kugleri 

M. micromeris 

M. pomum 

Muricopsis oxytatus 

Nassarina glypta 

Nassarius albus 

N. vibex 

Natica canrena 

N. livida 

Neodrillia jamaicensis 

Nerita peloronta 

N. tessellata 

N. versicolor 

Neritina virginea 

Nitidella dichroa 

N. idalina 

N. laevigata 

N. nitidula 

N. ocellata 

Nodilittorina tuberculata 

Odostomia laevigata 

Oliva reticularis 

Olivella adelae 

O. dealbata 

O. nivea 

O. perplexa 

Opalia pumilo 

Parvi turbo weberi 

Pedipes mirabilis 

Persicula miniata 

Petaloconchus erectus 

Phalium cicatricosum 

Phenacolepas hamillei 

Pira monile 

Pisania pusio 

Planaxis lineatus 



Vol. 75 (2) 

P. nucleus 

Polinices lacteus 

P. uberinus 

Prunum virginianum 

Psarostula monilifera 

Puperita pupa 

P. tristis 

Pyramidella crenulata 

P. dolabrata 

Pyrunculus caelata 

Pyrgocythara coxi 

P. emeryi 

Retusa caniculata 

Rissoa gradata 

Rissoina bryeria 

R. cancellata 

R. chesneli 

R. decussata 

Rhizorus acutus 

Sigatica semisulcata 

Sinum perspectivum 

Siphonaria alternata 

S. pectinata 

Smaragdia viridis 

Spartophos floridanus 

Stigmaulax sulcata 

Synaptocochlea picta 

S. coccina 

Tectarius muricatus 

Tegula excavata 

T. fasciata 

T. hotessieriana 

T. lividomaculata 

T. substriata 

T. viridula 

Terebra dislocata 

T. glossima 

T. hastata 

Thais deltoidea 

T. haemastoma floridana 

T. rustica 

Tonna maculosa 

Torinia infundibulum 

T. cyclostoma 

Trailia ovula 

Tiicola adamsi 

T. bella 

T. tessellata 

Trigonostoma rugosum 

Trimusculus carinata 

Triplioia decorata 

T. mirabilis 

T. nigrocincta 

T. pulchella 

Tritonalia intermedia 

Trivia pediculus 

T. quadripunctata 

Truncatella caribaeensis 

Turbo canaliculatus 

T. castanea 

Turbonilla curta 

T. interrupta 

Vasum muricatum 

Vermicularia spirata 

Xenophora conchyliophora 


Abra aequalis 

Aequipecten acanthodes 

A. muscosus 

A. gibbus 

Americardia guppyi 

A. medium 

Anadara notabilis 

Anadontia alba 

Anomalocardia brasiliana 

Antigona listeri 

A. rugatina 

Area umbonata 

A. zebra 

Arcopagia fausta 
Arcopsis adamsi 
Asaphis deflorata 
Barbatia cancellaria 

B. Candida 

B. domingensis 
B. tenera 

Basterotia newtoniana 
B. quadrata 
Botula fusra 
Brachidontes citrinus 
B. exustus 
Cuspidaria costellata 

October, 1961 



Chama congregata 

C. florida 

C. macerophylla 

C. sinuosa 

Chione cancellata 

C. grus 

C. mazycki 

C. pygmaea 

Chlamys benedicti 

C. imbricata 

C. multisguamata 

C. ornata 

Codakia costata 

C. orbicularis 

C. orbiculata 

C. pectinella 

C. portoricana 

Congeria leucophaeta 

Coralloiphaga coralloiphaga 

Corbula operculata 

Crassinella guadeloupensis 

C. lunulata 
Cumingia coarctata 
Cyathodonta cruziana 
Diplodonta punctata 
Divaricella dentata 

D. quadrisulcata 
Donax denticulata 
Echinochama archinella 
Ervilia nitens 
Glycymeris decussata 
G, pectinata 

G. undata 

Gouldia cerina 

G. insularis 

Isognomon alatus 

I. bicolor 

I. radiatus 

Laevicardium laevigatum 

L. 1. sybariticum 

L. mortoni 

Lima pellucida 

L. lima 

L. scabra 

L. tenera 

Limea bronniana 

Lioberus castaneus 

Lithophaga antillarum 

L. bisulcata 

L. nigra 

Lucina multilineata 

L. pensylvanica 

L. radians 

L. trisidcata blandus 

Lysonsia beana 

Lyropecten antillarum 

L. nodosus 

Macoma brevifrons 

M. constricta 

M. tenta 

Macrocallista maculata 

Mactra fragilis 

Modiolus americanus 

Mulina lateralis 

Muscukis lateralis 

Ostraea frons 

Papyridea semisulcata 

P. soleniformis 

Pecten ziczac 

Petricola lapicida 

Phacoides pectinatus 

Pholadomya Candida 

Pinctata radiata 

Pinna rudis 

Pi tar albida 

P. aresta 

P. circinnata 

P. fulminata 

Plicatula gibbosa 

P. sp? 

Pododesmus rudis 

Pseudochama radians variegata 

Pteiia colymbus 

Quadrans lintea 

Rocellaria cuneiformis 

R, ovata 

Semele bellastriata 

S. nuculoides 

S. proficua 

S. proficua radiata 

Solemya occidentalis 

Solecurtus cummingia 



Vol. 75 (2) 

S. sancta-marthae 
Spengleria rostrata 
Spondylus aniericanus 
Strigilla carnaria 
S. mirabilis 
S. piciformis 
Tagelus plebeius 
T. divisus 
Tellina alternata 
T. candeana 
T. crystallina 
T. cuneata 
T. interrupta 
T. guildingii 

T. laevigata 
T. promera 
T. similis 
T. sybaritica 
T. versicolor 
Thyasira trisinuata 
Tivela abaconis 
Trachycardium isocardia 
T. magnum 
Transenella cubaniana 
T. culebrana 
Trigonicardia antillarum 
Venericardia tridentata 




(Concluded from July no.) 

Succinea Indiana lives in places which are so dry that no other 
species of native land snail can survive there. Not only can it 
withstand sun and drought, but the ravages of roadgraders and 
bulldozers as well. Below the Clark Hill Dam, it is found abun- 
dant on land from which some twenty feet of earth had been 
removed during the construction of the dam. Land that even 
the weeds did not find entirely hospitable. At this locality the 
snails were heavily infested with leucochloridium. Every speci- 
men opened contained larvae or sporocysts. 
Succinea campestris Say 

Succinea campestris Say, 1817 Jour. Acad. Nat. Sci. Phila /.• 281. 

Succinea campestris vagans Pilsbry, 1900, Naut. 14: 74. 

Qiiickella vagans (Pilsbry) , Pilsbry, 1948; Land Moll. N. Amer. 
//, p. 848, figs. 443d. 

An examination of the shells of the type lot of Succinea cam- 
pestris vagans Pilsbry, disclosed that it was a mixture of two 
species of shells. One species, of which there were 5 or 6 spe- 
cimens, had a very short spire, large aperture and almost hyaline 
shell, and were not distinguishable from shells of Succinea 
campestris Say of the same size. The remaining shells, of which 
there were 25 to 30, were smaller, with longer spire and smaller 
aperture. These were not distinguishable from Catinella vermeta 
(Say) of the same size. Pilsbry's original description was clearly 

October, 1961 nautilus 61 

based on the shells of S. campestris. Later, when he described the 
animal, he used the animals of C. vermeta. Since the original 
description was based on shells of C. campestris, the name x/agans 
must be placed in the synonomy of that species. 

Succineids become sexually mature when very small. After 
they have mated they will add at least a full whorl to the shell. 
Dr. Pilsbry, apparently, was not aware of this, as vagens was dis- 
tinguished from campestris only by its smaller size and fewer 

Catinella texana, new species. PI. 4, figs. F to H; text-fig. 1, F 
to H. 

Shell pale golden-brown in color, translucent, rather dull, with 
about four whorls, elongate-ovate, sculpture of unevenly spaced 
growth wrinkles. Spire acute, long, sutures well marked, whorls 
well rounded. Aperture ovate, occupying about 58% of the length 
of the shell, outer and basal margins well rounded. Columella 
nearly straight. 

Mantle and sides of foot covered with a dark gray flecking. 
Hermaphrodite duct well pigmented. Talon strongly pigmented, 
almost black; club-shaped and undivided but with a terminal 
notch. Prostate gland large, oval and unpigmented. Vas deferens 
not very long, entering the penis at the apex. Penis rather broad 
unpigmented, with a large pouch, connected near the base on 
the anterior side. There is no trace of a sheath. Penial retractor 
connected to the side of the penis near the middle. Spermatheca 
large, globose, duct slender. 

Height Diameter Ap. H. Ap. W. 

13.3 mm. 6.7 mm. 7.7 mm. 5.0 mm. Holotype. 

13.3 mm. 7.3 mm. 8.3 mm. 5.7 mm. Paratype. 

10.8 mm. 6.0 mm. 6.3 mm. 4.0 mm. Paratype. 

Type Locality. — Texas: Milan Co.: near small spring, near 
Brazos River, 4,7 miles northeast of Cause, holotype 205892 and 
para types 205891 U.M.M.Z., other paratypes 15427, collection of 
the author. 

Catinella texana is most closely related to Catinella vermeta 
(Say) , differing in having a more slender shell, and in having 
the appendix basal rather than in the middle of the penis. 
Catinella pugilator, new species. PI. 4, figs. I to K; text-figs. 1, 
D & E. 

Shell straw-yellow, translucent, shining, oblong-ovate, with 
about three whorls, sculpture of fine growth wrinkles. Spire 
rather short, sutures moderately impressed, whorls convex. Aper- 
ture ovate, occupying about 68% of the length of the shell, outer 

62 NAUTILUS Vol. 75 (2) 

and basal margins well rounded. Columella nearly straight. 

Mantle gray, overlayed with black blotches. Hermaphrodite 
duct well pigmented. Talon moderately pigmented, club-shaped, 
with a terminal notch. Prostate gland rather large, somewhat 
rectangular in shape, lightly pigmented. Vas deferens rather 
long, slender, abruptly enlarging before it enters the penis, 
entering the penis at the apex. Penis broad, unpigmented, with 
a small thumb-like appendix near the center of the posterior 
side. There is no trace of a sheath. Penial retractor connected to 
the anterior side of the penis near the middle and to the thick- 
ened portion of the vas deferens. Spermatheca rather large, 
globose, duct rather short, thick. Atrium and vagina very short. 
Height Diameter Ap. H. Ap. W. 

13.2 mm. 7.1mm. 9 3 mm. 5.1mm. Holotype. 
11.1mm. 6.0 mm. 7.8 mm. 4.7 mm. Paratype. 

10.3 mm. 5.3 mm. 7.0 mm. 4.0 mm. Paratype. 
Type Locality. — North Carolina: Currituck Co.: salt marsh, 0.5 

mile south of Maple, holotype 205889 and paratypes 20589D 
U.M.M.Z., other paratypes 20980, collection of the author. 

Catinella pugilator resembles C. hubrichti Grimm in its Oxy- 
Zoma-like shell, but differs in its boxing-glove shaped penis. The 
very small penial appendix plus the thick spermatheca duct will 
readily separate this species from all other described species. 

Vertigo teskeyae, new species. Text-fig. 2. 

Shell ovate, dark chestnut, with about five whorls. Spire con- 
vexly conic, with obtuse summit; whorls increasing rapidly, the 
last much the largest, its height equal to almost two-thirds of the 
shell, sculpture of fine irregular growth lines. Lip thin and 
expanded, with a distinct crest behind it. Aperture with a dis- 
tinct sinulus. Parietal lamella rather strong, about twice as long 
as high; angular lamella small; an infraparietal tubercle is never 
present. Columellar lamella strong, horizontal, located in about 
the center of the columella. Basal fold variable, rarely very 
large, often nearly obsolete, located on the end of a callous ridge 
which extends along the basal and lower outer walls just inside 
of the lip. Infrapalatal tubercle often present, although usually 
very small. Lower palatal fold moderately well developed, upper 
palatal fold somewhat smaller. Suprapalatal fold never present. 




2.6 mm. 

1.6 mm. 

5 Holotype. 

2.1 mm. 

1.5 mm. 

4 Chocowinity. (smallest) 

2.9 mm. 

1.8 mm. 

5 Lightwoodknot Creek, (largest) 

Distribution — Maryland: Dorchester Co.: subfossil, near beach, 
0.5 mile south of Elliott. North Carolina: Camden Co.: swamp, 1 

October, 1961 



Figure 2. Vertigo teskeyae: A, holotype. B, paratype. C, from Light woodkiiol 

mile south of South Mills. Gates Co.: near Bennetts Creek, Gates- 
ville. Chowan Co.: swamp, 1.2 miles south of St. Johns. Beaufort 
Co.: swamp, 4.8 miles north of Pantego; near Pamlico River, 
opposite, VVashington; swamp, 1.4 miles northeast of Chocowinity. 
Craven Co.: near Little Creek, 1 mile north of Askin; swamp, 
1.8 miles south of Vanceboro. Columbus Co.: bank of canal, west 
side of Lake Waccamaw, holotype 205903 and paratypes 205904 
U.M.M.Z., other paratypes 20870, collection of the author. South 
Carolina: Williamsburg Co.: near Black Mingo Creek, north of 
Rhems. Georgia: Screven Co,: near Brier Creek, 10 miles east- 
northeast of Sylvania. Toombs Co.: swamp, 6.5 miles south of 
Parkers. Florida: Flagler Co.: near Pellicer Creek, 13.5 miles 
north of Brunnell. Holmes Co.: low woods, 2.5 miles west-north- 
west of Sweet Gum Head. Alabama: Montgomery Co.: low woods, 
0.5 mile north of Mt. Meigs. Covington Co.: near Lightwoodknot 
Creek, 3.7 miles west of Opp. Mississippi: Wanen Co.: low woods, 
near Big Black River, 6.2 miles east-northeast of Bovina. 

Vertigo teskeyae is most closely related to V. ovata Say, from 
which it may be readily distinguished by its distinctly darker 
color (when fresh) , larger size, and fewer teeth. The infraparietal 
and suprapalatal tubercles are never present. The columellar 
lamella is horizontal, not sloping upward, and is lower in posi- 
tion on the columella. The basal fold is smaller and is lower 
down. The aperture is much more open than is usual in V. ovata. 
V. teskeyae is the largest Vertigo known from the southeastern 
United States. It is a species of the southern Atlantic and Gulf 
Coastal Plains. In North Carolina, it is found cra^vling on the 

64 NAUTILUS Vol. 75 (2) 

wet mud and debris of swamps. Farther south it may be found 
on the undersides of palmetto leaves in company with Pupisoma. 
It is frequently found associated with V. ovata. 

Vertigo teskeyae is named in honor of Mrs. Margaret C. 
Teskey, in recognition of her services in behalf of the American 
Malacological Union. 



The symbols used for Puerto Rican localities were explained 
recently (I96I) . 

Caracolus carocoUa (Linne). Under leaves on ground when 
dry, but fair climber on tree trunks, up to 10 ft. around El 
Yunque; En, Er, En3,4, Jn, Pn, Pr, Wn, Wr, Ws (dead) ; every- 
where except in driest places; not even empty shells recorded 
from Ps or Ww; 0-4000 ft. Animal (added to Wurtz, 1955:112) 
sluggish; foot large, with sides varying from slate color to dark 
chestnut with buff spots; all tentacles dark; sole trizonal, with 
coarse waves. 

The obsolete name Helix arecibensis Pfeiffer, 1856, evidently 
was based on a juvenile shell of Caracolus, probably this species. 

Caracohis marginella (Gmelin) . Very good climber on tree 
trunks, nearly subarboreal, often seen 15-20 ft. above ground; 
Ee, En, Es, Jn, Pn, Wn; mainly on lowlands, especially near 
cultivation; not seen above 1500 ft. in Luquillo Mts. or in the 
eastern Cordillera Central (Pr) . Animal dorsally almost black, 
but shading into whitish near sole; ommatophores dark but 
eyes darker. Uniformly light greenish shells with whitish animals 
(Jnl, near which "albinos" were found in 1864, v. Bland, 
Caracolus marginella mayaguezi, new subspecies. 

Pleurodonte bornii (Pfr.) Pilsbry, 1889 (6b) : 127, in part, pi. 
56, fig. 6 type (ANSP. 20861) from Swift Collection, labeled 
"Mayaguez." Type locality (the older collectors often gave the 
nearest port): top of Pico Montoso, near Maricao, elevation 2300 
ft. (Wr2) , where shells match Pilsbry's figs, very closely. Shell 
differs from typical C. marginella by its broader dark zones, 
which on the apical side restrict paler to narrow, sutural and 
carinal zones. Also found: Maricao Forest (Wr3) , near Cabo Rojo 
(Ws) and near Guanica Bay (Psl) , 0-3000 ft. Also noted by 
Bland, 1875:80. 

All shells collected further east, including those around mouth 
of Rio Guajataca (Wn), which slightly approach mayaguezi, 

October, 1961 nautilus 65 

are quite typical marginella. Probably the 2 forms intergrade, 
since the typical one thrives in lowland, cultivated places; in 
fact, from field notes, I saw living "marginella" (which form?) 
on trees in the garden of the Agricultural Station in Mayaguez. 

Polydontes (Luquillia) luquillensis (Shuttleworth) . Adults on 
trunks of trees, 5-12 ft. up, but young (looking like those of P. 
lima) usually on ground; Luquillo Mts., above 2000 ft., where it 
replaces the "closely allied" P. lima. Animal bright ochraceous 
(but one almost white); eye and inferior tentacles almost black; 
sole blackish slate. 

Although these lots show considerable variation in banding 
and height (Cf. Pilsbry, 1889 (2) :pl. 7, figs. 63, 64) none matches 
that of van der Schalie, 1948: pi. 6, fig. 9, which seems immature 
and weather worn. Despite considerable differences in its later 
whorls, Granodomiis Pilsbry, 1931, seems only a section of the 
subgenus Luquillia Crosse, 1892 (Cf. Wurtz, 1955:132). 

Polydontes (Granodomus) lima (Ferussac, 1821) Fair climber 
on tree trunks and brush, but less so than C. marginella; espe- 
cially common near cultivation; Ee2, Es, Jn, Js, Ps2 (dead) , Wn, 
Ws (1 shell form castrensis) , Ww; lowlands. Secretes quantities 
of disagreeable mucus; seen in reciprocal copulation. ANSP. 
1125 (from Bland, without exact locality) contains 2 shells of 
form castrensis and one of subspecies maricao; see Pilsbry, 
1889 (6b) :pl. 1, figs. 3 (castrensis) and 4 (maricao). 

P. (G.) lima maricao Clench. Apparently climbs less than 
typical lima; Prl-3, Wr2,3; Cordillera Central, 2500-4000 ft. 

Dr. van der Schalie's (1948:79) nice "curves" show long and 
careful study, but is a "random sample" from a large area 
actually possible? Might not the "normal distribution" of his 
composite one be due to his larger series, as shown by his maps, 
of shells from lower altitudes? If based on enough animals, that 
from over 1500 ft. may be significantly bimodal, and the habitat 
of typical maricao seems considerably higher. 

My lots of P. lima maricao often show less prominently the 
peripheral brown band and sometimes additional, finer ones 
like the larger shells from the Virgin and Leeward Islands. For 
this reason, in part, the very involved nomenclature of the latter 
is outlined. Perhaps, rnaricao and asperula may be "microspecies," 
and "typical" lima, which thrives in cultivated areas, might 
consist of very unstable hybrids between them? 
P. (G.) LIMA, VAR. (?) iNCERTA ("Fer." Bcck) or (Ferussac) . 

66 NAUTILUS Vol. 75 (2) 

Helix incerta Ferussac, 1823, livT. 18: pi. 105, figs. 2 (named on 
cover?) . (Helix) Otala incerta Beck, 1837:36, from Lesser Antilles 
(sic) , founded on figs, cited. Not Helix incerta Draparnaud, 1805. 
(?) H. incerta Pfeiffer, 1841, Symb. Hist. Helic.:37, from St. 

Dir. Pilsbry and I always disagreed on the delicate question as 
to whether Beck's name should be preoccupied in his genus 
Helix, or only in his subgenus Otala, but Beck himself, 1837a, 
when he added description to a few of his nude names, regularly 
used his genus or its initial; e.g., Nanina (Microcystis) ornatella 
(p. 2) . In any case, "incerta" should be returned to the synonymic 
oblivion, from which Pfr. snatched it. The Philadelphia Acad. 
Nat. Sci. has 3 lots of the fossils from St. Thomas (ANSP. 1033 
from Bland) and I agree with R. Swift (Mss. in ANSP. 30634 
from St. John I., and in 30636 from St. Thomas) that Ferussac's 
figures look most like one, worn, deformed shell of P. lima. The 
apparently dark band of his left fig. is absent from his right one 
and Deshayes made no mention of it. 

P. (G.) LIMA ASPERULA (Beck) . 

Helix lima, var. notab. (sic) Ferussac, 1832, livr. 22, Explic. 
pis. livr. 22-27: p. ij; livr. 24, pi. 46a, figs. 4 & 5 [copied by Pils- 
bry, 1889 (2) :pl. 4, figs. 36 & 37]; certainly not a valid name. 
(Helix) Otala asperula Beck, 1837:36, founded solely on figs, 
cited above, from Curacao (sic) , but with H. Hgen. lima var. 
notabilior Fer., in synonymy. Helix notabilis Shuttleworth, 
1854:40, partly founded on figs, cited above, from Is. San Juan 
(St. John) and Anegada, and with Helix lima var. notabilis 
Fer. cited. Thelidomus notabilis (Sh.) Pilsbry, 1889 (6b) & 1890 
(5b) : pi. 1, fig. 1 (ANSP. 1047, Tortola, from Thos. Bland) and 
var., pi. 1, fig. 1 [ANSP. 30634, labeled as from San Jan (St. 
John I.) by Krebs (from Swift Collection) ]. 

Since they all are founded on exactly the same figures, I now 
select Ferussac's fig. 4 (the adult, cited above) as representing 
the type shell, and Tortola, Virgin Islands, as the type locality 
of notab. Fer., asperula Beck and notabilior Beck. The ANSP. 
lots from St. John Island (now designated as type locality of 
H. notabilis Sh.) seem usually a little thinner and more com- 
monly have additional bands above and below periphery, but 

Incidentally, all the records of "incerta" (actually asperula) 
from Puerto Rico are due to Kreb's label of "San Jan," which 

October, 1961 nautilus 67 

I certainly (and probably Pilsbry) took to be a misspelling 
of San Juan, Puerto Rico, where asperula does not occur. To 
add to the confusion, the principal town of Antigua apparently 
was named after the same saint, and shells similar to the 
Tortola form in banding, but thinner (like those from St. 
John I.) lived on St. Bartholomew (ANSP. 1051 from Bland), 
which also is in the Leeward Islands. Anegada (ANSP. 1049 
from Bland) is another Virgin I. Finally, one shell (ANSP. 
30628 from W. G. Binney) is labeled Martinique, and looks 
like the Tortola form. 

Polydontes (Parthena) acutangula (Burrows) . Adults truly 
arboreal, but young animals often found under leaves on 
ground; Enl, Er, Es4, Janl, Pn2 (long dead), Prl; 0-3400 ft. 
east of San Juan, but westward only found living at 4000 ft. 
Green mantle of adults showing through shell, but young ones 
ochraceous brown. Animal (added to Wurtz, 1955:112) in- 
capable of withdrawal into shell, extending out to 8 inches 
(20 cm.) long, including about I1/2 inches of tentacles, but 
mantle only extending slightly over columella; tail long, angu- 
late dorsally with median groove; sides of foot becoming brown- 
ish near sole: ommatophores darkish slate; inferior tentacles 
lighter; mantle collar orange-brown; mucus yellow; seen in 
reciprocal copulation on trees. 

Van der Schalie, 1948:82, obtained another camaenid, Zachry- 
sia auricoma havanensis Pilsbry, a Cuban species. 
Additional references (See 1961) 
Baker, H. Burrington. 1961 (1), Naut. 7^:142. 1961 (3) „ Naut. 

Beck, H. 1837, Index molluscorum, etc.: 1-124. 

1837a, Specierum novarum, etc.: 1-8. 

Wurtz, Charles B. 1955, Proc. Acad. Nat. Sci. Philadelphia 
707:99-143, pis. 1-19. 



Oegstgeest, The Netherlands 

To identify Papuina ferussaci (Lesson) , I looked up the history 
of the species and its whereabouts. The description, published 
in 1831, is rather short but sufficient; moreover a figure was 
given by him. Unfortunately his type specimen is probably 
lost, Tapparone Canefii tried in vain to locate it in the museum 
at Paris. Pfeiffer, who had in 1848 to deal with the species, 

68 NAUTILUS Vol. 75 (2) 

could only copy the description in his Mon.Hel.Viv., p.231, but 
in the year 1868, in Vol. 5. of his Monograph, he identified 
material from Cumniing's collection as being Lesson's species 
and gave a new description of his own. This new description, 
however, was in many important characters, very different from 
the original one. Tapparone recognized the error and demon- 
strated convincingly in the year 1883, that Pfeiffer's so-called 
Helix ferussaci, both the description and the colored plate in 
the Novitates Conch, tab. 56 fig. 7-9, was a characteristic new 
species and named it Helix exsultans. In the well-known Manual 
of Conch. Vol. 7, p.30, f.51 the true species of Lesson was placed 
by Pilsbry in the genus Papiiina; he gave a good description, 
taken from the original one and from the plate. This description 
runs as follows: "Shell imperforate, trochiform, deep-chestnut, 
with a golden-yellow line at the carina, and ascending the spire 
to the apex, whorls 6, completely flat, the apex is enlarged, 
convex; the last whorl has a salient acute carina. Aperture 
much depressed, triangular, wider than high, very acute at the 
junction of the appressed basal lips; peristome reflexed, sinuous 
at the angle." 

In the year 1933, Lesson's species was mentioned by Madame 
V. Benthem Jutting on the list of all the species of the genus 
Papuina, known at that time. This important and useful list 
was published in the periodical Nova Guinea, Vol. 17 (Zoology) 
Livr. 1, p. 411. She placed in it var. exsultans (Tapp. Canefri) , 
the color being much diluted to cinnamon. Because this color 
is the typical one of Tapparone's species, the identification, in 
my opinion is probably correct. 

The most recent mention of Papuina ferussaci is given by Adam 
and Leloup in Vol. 2, p.29 (Mus. Roy. Sc. Hist. Nat.) in the 
year 1939, together with fig. 16 on Plate 2. The only specimen 
they had, was found in 1929 near Manokwari, which is the 
modern name of the type locality. Port Dorey of Lesson. Their 
specimen was in bad condition, worn and nearly totally bleached, 
but with the yellow keel still present at the base of the shell 
and with the typical brown color. I agree with both authors 
that their specimen belongs to the true P. ferussaci of Lesson, 
after comparison with their figure 16. Since that time, no further 
data are known to me. 

In the year 1949, Father Rombouts, a missionary at Sorong, 

October, 1961 nautilus 69 

told me that he would penetrate what he called the Aifat Anoek 
Tifa Region, in the northeast part of "The Vogelkop," to visit 
some papua-settlements. Because he was much interested in 
natural history and the region was never visited by collectors, 
I asked him if he could give some attention to living land-mol- 
lusks. He promised to do that. Although he was obliged to 
return earlier, on account of the unkind and hostile natives, 
he collected a very interesting lot. Upon his arrival home, he 
handed the specimens to me and we examined them together. 
Among them there were species I had already found myself, 
but others were quite new to me; among them was a very beau- 
tiful species of Papuina, the subject of this paper. 

Pointing to this Papuina Father Rombouts said: "This one 
must be very rare and I found it so beautiful, that I tried to 
find some more, but could not obtain another one." 

All the characters, given by Lesson and from the translation 
by Pilsbry, given above, agree perfectly with this specimen. 
Lesson mentioned, however, only the general and easily rec- 
ognizable ones, so that the more subtle ones (mostly visible 
under a strong lens) are wanting in his description. I give here 
an additional description of my specimen. 

High trochiform shell, somewhat higher in proportion to 
the width, dark castaneous, the faintly convex base shining, 
^vhereas the whorls above the very sharp keel are of a dull 
color. Whorls nearly totally flat, only very slightly concave 
to their lower sutures, the profile very different from those of 
the allied species P. exsultans (Tapp.) and P. pythonissa (Tapp.). 
Keel with a golden-yellow, 1/2 mm broad band, visible also along 
the sutures and below the keel. The common oblique growth- 
lines very distinct, under a strong lens; the whorls, especially 
the lower one, with about 5 regular spiral-lines, base radiately 
striated, towards the umbilical region somewhat deeper and 
wider grooved. Whorls just below the suture slightly darker 
colored, forming on the ultimate one a faint, not sharply de- 
fined zone. Apex somewhat darker colored. Umbilicus a faint 
perforation, obtected by the columellar margin. Peristome 
darker, more blackish-brown and the aperture provided within 
with a grayish-white, rather sharply defined margin, the inner 
part of the aperture brown. Columellar margin rather broadly 
reflexed and provided at its base with a distinct, although 
small tooth-like pale projection, the columella passing rather 
bluntly to the nearly straight lower margin of the peristome; 
outer margin of the peristome beaked, with a sharp keel. 

70 NAUTILUS Vol. 75 (2) 

The small columellar tooth, mentioned in my description, was 
not indicated by Lesson and, in the lithographic picture by 
the artist, this character did not show fully. Moreover, the 
dimensions are not indicated on Lesson's plate. In the descrip- 
tion the diameter is given as 11 lines; accepted as French lines, 
this becomes 24.8 mm, and my specimen measures 28.2 mm. 
My shell is therefore somewhat larger. Because, however, in 
many species of the genus Papuina, there occur fluctuations in 
the dimensions, there is, for the time being, no reason to accept 
a special variety for my larger one. 

Pilsbry placed P. ferussaci in his 6th group, the "Group of 
the P. pileolus." He characterized this group as "subperforate 
or imperforate, pyramidal, sharply keeled and flat at the 
base." Unfortunately, many species, mentioned by him, were 
not sufficiently known to him, especially if there were no 
figures, and this group is very heterogeneous. P. leonardi (Tapp.) 
and P. turris (Adams) are not sharply keeled at the base; they 
do not belong to this group. The former certainly belongs to 
the 5th group of Pilsbry, called: "the group of the P. antiqua." 
This also was recognized by Madame v. Benthem Jutting, who 
accepted P. leoyiardi simply as a synonym of P. antiqiin (Adams 
& Reeves) , in the list of the species of Papuinn in Nova Guinea 
Vol. 17, cited above p. 121. Papina turris (H. Adams) has only 
a compressed last whorl with a very blunt keel and must be 
eliminated also. The remaining species of the group are, as 
to form and outline, very different and it is thus very heterogen- 
eous. We can, however, divide the group easily into two homo- 
geneous ones, if we accept for the true "group of the 
P. pileolus" only the low-spired species without a subcolumellar 
tooth, viz. P. pileolus (Ferussac), P. rhynchostoma (Pfeiffer) and 
P. fergusoni (H. Adams). The other group, containing high-spired 
species with a very distinct subcolumeller tooth is to accept as the 
"group of the P. ferussaci." 

The new group is not only very homogeneous but, moreover, 
zoogeographically sharply limited, because all the species, be- 
longing to this group, inhabit the northwest part of Dutch New 
Guinea. All its members are represented in my collection. I 
shall summarize them afterwards more in detail in another 

October, 1961 nautilus 71 


Lake Miragoane in the southwest peninsula of Haiti is a 
small muddy lake which is about three square miles in area 
and lies near the coast about half-way between the towns of 
Petit Goave and Miragoane, approximately 60 miles on the 
road west of Port-au-Prince. It is the largest fresh- water lake 
in Haiti and its vicinty is sparsely populated. There are two 
other large lakes on Hispaniola which are several hundred 
square miles in area which are saline and were originally inlets 
of the Gulf of Haiti in geological times. These are the Etang 
Saumatre and the Lac Enriquilla (150 ft. below sea level) , both 
of which lie due east of Port-au-Prince about 50 miles. The 
latter lake is all within the Dominican border. 

In 1927 while collecting plants and land snails for the 
Smithsonian institution and reptiles for Dr. Thomas Barbour 
in the southwestern peninsula of Haiti and the islands of Gonave 
and Grande Cayemite, I also made a special effort to collect 
all the freshwater shells that could be found. More than 1000 
lots of land shells were collected and sent to Dr. Bartsch. Most 
of them were taken alive or in a good condition. For one thing 
I found that Haiti compares in no manner with Cuba in rich- 
ness of its land shell fauna. There is only one species of Liguiis, 
and only a very few species of Cerion and Urocoptis, of which 
there are many hundreds of species in Cuba. The only family 
of land snails that seems fairly well represented is the 

There are many areas, some of which are of considerable 
extent where snails are exceedingly scarce. The topography 
of Haiti is very rough and mountainous with more than 
3,000,000 people of which over 99 percent are negro. Most of 
the popvilation are peasants. 

It was my good fortune to team up with the great Swedish 
botanist, and naturalist. Dr. Erik Ekman, the foremost authority 
on Caribbean plants who knew intimately the flora of Cuba 
and Haiti. Before he died in the Dominican Republic in 1930, 
he had discovered over 2,000 new species of plants in Cuba 
and Haiti during his 20 years of botanical rambles. 

I spent nearly 5 months in hard marches and almost con- 

72 NAUTILUS Vol. 75 (2) 

tinuous biological collecting with Ekman, who would quickly 
drop a companion if he couldn't keep up with his pace and 
live the primitive life to which he had adapted himself. We 
never hired natives or pack animals and we carried all our 
supplies on our own backs; we lived on native food, drank water 
wherever it could be found, often from stinking pools; we 
shot birds, stuffed the skins, and ate the meat. Guinea fowl are 
abundant in some areas and the natives seldom molest them. 
Haitian peasants are usually very poor, but invariably hospitable, 
so generally we slept in native huts while collecting in the 
mountains and islands. 

At that time the American Marines still occupied Haiti be- 
cause of the Caco rebellion a few years before. Ekman, a very 
eccentric man who cared nothing about formalities and hated 
shams or snobbishness was, nevertheless, a very learned man 
and a delightful companion. He was well posted on a wide 
variety of subjects, was conversant in all the native dialects and 
could speak at least 8 European languages. He was known per- 
sonally throughout Haiti by much of the peasantry, the military 
officers and by the president. The American Marine officers 
called him the "Iron Man of Haiti" or the "Crazy Swede," be- 
cause very few men had the stamina and drive to keep up a 
continual pace of hard marches into the rough jungles and 
live on native food and sleep wherever he happened to be 
when darkness fell. 

Our regular routine was to be up at 4 A.M., start off imme- 
diately, but without a breakfast, eat a few mangoes or wild 
fruits if we found them, or purchase a couple of pennies' worth 
from peasants during the day, and at sundown we usually had 
a dish of rice and a boiled fowl and Haitian hard tack with 
plenty of tea and sugar, around a small campfire. 

Lake Miragoane is quite round and has a very muddy bottom 
of unknown depth. It is completely surrounded by reeds and 
dangerously deep mud. The flora is particularly interesting 
around the outlet of the lake. A large waterlily grows here 
with stems 12 to 15 feet long. Ekman always knew right away 
when he found a new species of plant and was almost always 
correct. Here I found a new fern and a duckweed. Numerous 
species of ferns a nd several beautiful species of Canna grew 


October, 19(il nautilus 73 

in the swamp. While Ekman and I gazed longingly at the lake 
and contemplated its floristic possibilities for our botanical col- 
lection, wondering how to find an area where we would not 
disappear into the treacherous mud, we finally found a small 
boy, who had a hollowed out log boat, and who agreed to take 
us wheie we wanted to go. He paddled and poled the boat 
along the margin of the lake until an ample series of desired 
plants was made. The marsh around the lake, and the surface 
of tiie water, was a regular aviary of interesting birds. Glossy 
ibis, two kinds of egrets, several species of ducks, four kinds 
of herons, the blue, the green, the Louisiana and night 
herons, were there. Curlew, yellowlegs, grebes, coots and sand- 
pipers and the curious jacanas, skipping and running about 
on the lily pads after insects. To a real ornithologist or a 
naturalist with proper equipment and plenty of time, this lake 
has a great deal of interest. 

On this special excursion we asked the little boy to put us 
ashore on the margin of the lake of the opposite side. There 
the mud appeared firm enough to walk on through the marsh 
to the land. The objective was some rare species of ferns and 
grasses that could be seen from the lake. The boy left us, and 
we went our way to the shore. Soon we started to break through 
the mud crust, and were floundering around in the deep ooze 
for some time, often falling through the upper crust only to 
become more deeply mired. We began to grow quite anxious 
as to how we should get back to safer ground, and out of the 
swamp. Suddenly we came upon a black woman intently engaged 
in setting snares for wild fowl. She hadn't noticed us at all 
until we were almost upon her when she became terribly 
frightened at our appearance out of the mud. She screamed 
and tried to run away, but, after a small gift of a few cigarettes, 
she calmed down and showed us a sort of path out of the big 
swamp. It was really a lifesaver for us. 

Upon gaining safer ground on the lake shore, I found a few 
spots in shallow water inhabited by moUusks. During about 
one half hour, spent searching for freshwater shells, I collected 
the following species, from which Dr. Clench (1936) and 
Clench Sc Aguayo (1937) named a subgenus and 4 forms. 
Physa (Haitia) elegans Dr. Clench. Made type of a subgenus. 

74 NAUTILUS Vol. 75 (2) 

because not closely related to the other forms in the West 

Helisoma eyerdnmi Clench and Agiiayo. 
Helisoma caribaea (Orbigny) 
Tropicorbis pallida (C. B. Adams) 
Tropicorbis albicans (Pfeiffer) 
Drepanotrema cimex (Moricand) 
Drcpanolrcrna anatiniim (Orbigny) 
Potamopyrgus coronatus ariomus Clench & Aguayo. 
Ferrissia (Laevapex) haitiana Clench &: Aguayo. 

Up to that time, less than 50 species of freshwater shells were 
known from the West Indies and Bahama islands. 

Later in the afternoon, we found a beautiful waterfall where 
we had a delightful bath, collected some Tare plants and snails, 
and I also collected a rare frog, Eleotherodactylus inoptatus 
(Barbour) and a rare snake Uromacer oxyrhyricliys (Dumeril 
and Bibron) . 

Several times before I had come alone to the south side of the 
lake to collect specimens. There in the jungles many fine land 
snails were taken, including Biilimulus, Cepolis and Caracoliis, 
and a colony of Liguiis xiirgirieiis on the lignum vitae and 
logwood trees yielded several hundred fine specimens. On the 
ground, amongst these trees, I also collected scorpions, taran- 
tulas, centipedes and several specimens of the giant Haitian 
armoured millepede, over 8 inches long and about one inch 
thick. When disturbed it sprays acid at its enemies a distance 
of more than a foot. The substance is exuded from pores along 
the sides. It smells and burns like sulphuric and prussic acid. I 
did not realize how dangerous these creatures could be with 
careless handling until I met Dr. Cook, American specialist of 
palm trees, centipedes and millipedes a few days afterward in 
Port-au-Prince. Most of the skin of one side of his face was 
burned off, and his helper had one of his hands badly burned 
as a result of heedless collecting of these large millipedes. 

No serious work has ever been done in collecting the fauna 
of lake Miragoane except a casual excursion made by Dr. Wm. 
Beebe in 1927 a lew weeks before I came to Haiti when he 
discovered two new endemic fishes. One of them, a tiny guppy 
about 14 inch long, fully grown, was rather common in a 
ditch near the lake. With its high degree of endemism, a careful 
collecting survey of its limnological fauna by a couple of 

October, 1961 nautilus 75 

competent biologists should bring to light a surprising number 
of new and interesting species of animals. 


Clench. William J. 1936. Mem. Soc. Cuba Hist. Nat. /0:335-342, 
pi. 25. 

& C. C. Aguayo. 1937. Mem. cit. //:61-76, pi. 7. 

Cochran, Doris M. 1928. Biol. Soc. Washington 41:53-60. 



University of Wisconsin-Milwaukee 

This note describes an experiment in which snails of the genus 
Pliysa, identified as P. Integra Haldeman^ were tested for geotac- 
tic behavior in a mud medium. 

Method. Six populations of P. Integra, collected from the 
locations given in Table 1, were used in the study. Experiments 
were conducted between September 25 and Octobei- 8, 1960. Each 
population was collected early on a given day, and tested in 
the afternoon or evening of the same day. The medium in which 
the snails were tested was a silty mud obtained from the banks 
of the Milwaukee River at Estabrook Park, on the north side 
of the city of Milwaukee. The populations designated as "2", 
"4" and "5" in Table 1 were collected in the same area as the 
mud. The mud was washed in tap water, sieved through a fine 
mesh screen, and allowed to dry to a tacky consistency. Some 
oligochaetes of the family Tubificidae were present in the mud 
after sieving; no other forms of life were obvious. Before 
testing a given population, tap water was added to a portion of 
the mud, which was then used in the test as described below. 
In some cases, the mud utilized had been employed in previous 
tests, in other cases, fresh mud was used. 

Twenty snails were selected at random from the population 
to be tested, each snail being measured at the time of selec- 
tion. The average shell length of the animals used was about 
7mm. Each snail was placed alone in a 150 ml. glass beaker 
filled to a level of 4mm. with mud. Although the orientation 
of the snails was not controlled, most animals appeared to land 
in an upright postion. Care was taken to place the snails in the 

1 The writer wishes to thank Dr. H. van der Schalie, Museum of Zoology, 
University of Michigan, for kindly identifying the specimens. 

76 NAUTILUS Vol. 75 (2) 

middle of the mud surface. Mud was then added over the 
snail in each beaker to form a mud column totaling 50mm. in 
height. The mud column was typically of a consistency such that 
after a half hour approximately 5mm. of water had collected 
above its surafce. At the start of a test, each snail was therefore 
immersed in a mud medium at an approximate distance of 
4mm. from the beaker floor, 25mm. from the beaker wall, and 
45mm. from the mud surface. The 20 beakers were then placed 
on a table 4 feet below a 100 watt lamp. Room temperature 
during the tests ranged from 19-22°C. Beakers were inspected 
at 5 minute intervals, mud surface, beaker wall, and beaker floor 
being watched for the appearance of snails. As the mud medium 
was homogeneous in texture, snails appearing could readily be 
identified. Each beaker was observed for a period of two hours. 
If a snail had not appeared by the end of this period, its 
position in the mud was ascertained by probing. 

Results. Of the total of 120 snails used in the experiment, 
61 surfaced on the mud. Surfacing usually occurred at a point 
roughly between the center of the beaker and the beaker wall 
so that, assuming travel in a straight line, the 61 snails moved 
towards the surface along a path that was at an average angle 
of 70° from the horizontal. The average time lapse between 
start of a run and surface appearance was 42 minutes. Eight of 
the 120 snails were recorded as appearing at beaker walls at 
points ranging from half way up the beaker wall to just below 
the mud surface. Again assuming travel along a straight line, 
snails moving towards a beaker wall did so along a path 
that was at an average angle of 50° from the horizontal. Average 
time lapse between start of a run and appearance at a beaker 
wall was 6.^ minutes. Fifty-one of the 102 test snails did not 
appear at either the surface or the wall of a beaker. Probing 
indicated that 15 of these had moved up through the mud 
medium, having traveled from I/3 to 14 the distance to the sur- 
face. The remaining 36 snails were found to be in approximately 
the location where they had been placed at the start of the 
two-hour period. 

Snail size did not appear to be a significant factor in the 
behavior shown. The consistency of the mud medium differed 
slightly between tests, and this may have been a partial cause 

October, 1961 



of the behavior variations found between populations. Possibly 
also more exact tests would reveal actual behavior differences 
between snails of different populations. 
A summary of the results is given in Table 1. 

Table 1 

Behavior < 

3f Physa 

in a Mud Medium 



of Snails Recovered in Each 
Four Locations 




Mud Part Way 
Surface to Surface 



















































Collection sites of popiilations were as follows. 1: Aquarluw in Botany Department 
at University of Wisconsin — Milwaukee. 2, h, $: Milwaukee River bank at Estabrook 
Park, Milwaukee County. 3: Stone fountain at Whltnall Park, Milwaukee Coonty. 
6? Stream at Kletzsch Park, Milwaukee County. 

Discussion. Possibly vertical gradients involving physical- 
chemical properties exist in a mud medium and serve to guide 
an immersed snail to the surface of the mud. Gradients of this 
type might conceivably involve differences in temperature, light, 
oxygen, or in the concentration of various organic or inorganic 
materials in the mud. 

An alternate, and perhaps more likely, explanation of the 
upward movement of the snails in the present experiment is 
that the mud itself provided no cue as to direction, but that 
the movement was a true geotactic behavior, or response to 

Snails are usually considered to orient to gravity either by 
statocysts, organs designed to ascertain position in space; or by 
proprioceptors, sensitive to the differential stresses in muscle 
which occur in an organism subjected to the pull of gravity 
(Carthy, 1958). 

While statocysts are present in snails (Bouvier, 1887; Lacaze- 
Duthiers, 1872) , these organs have not been clearly shown to 

78 NAUTILUS Vol. 75 (2) 

I'unction in gcotaciic behavior, for the experimental work which 
has been done in this respect is open to criticism, having proved 
difficult to repeat (Pieron, 1928; Crozier and Navez, 1930). On 
the other hand, a strong case has been made for the existence 
of a proprioceptive control of snail geotactic behavior. Experi- 
ments have shown that snails which normally show negative 
geotactic movement will change their direction of travel if 
muscle stress becomes greater on one side of the body than 
the other. This latter condition can be brought about by 
attaching a thread to the shell apex and gently twisting the 
shell out of line with the snail's body. Snails so manipulated 
turn so that their body lies along the same axis as the shell, 
and travel in the new direction determined by this axis, even 
if the new direction is one that is positively geotactic. It has 
been concluded from experiments of this sort that snails show- 
ing negative geotactic behavior have oriented themselves so 
that there is an equal muscle tension on both sides of the body, 
as would presumably be true when the shell hangs ventrally. 
A snail traveling in any direction but upward woidd presumably 
experience unequal muscle tension due to the weight of the 
shell (Crozier and Navez, op. cit.) . 

While possibly snails may orient to gravity by this method 
in air or water, orientation of this kind would seem less likely 
to occur in a medium such as mud. In a mud medium, there is 
probably very little tendency for a snail's shell to sink faster 
than its body and so create muscle tension, forcing the body to 
orient upward. This is almost certainly true when a snail is at 
rest in a mud medium, and is probably true to a large extent 
when a snail is moving through the medium, although this 
remains to be verified. In any case, the results of the present 
experiment indicate that a re-examination of the possible role 
of statocysts in snail orientation seems desirable. 

The utility of negative geotactic behavior in some species 
of snails, such as the tree forms, may be that of enabling an ani- 
mal to regain its normal habitat after accidental dislodgement 
(Crozier and Navez, op. cit.) . 

In aquatic pulmonates, negative geotactic behavior may be a 
mechanism which sends an animal to the surface to gain 
oxygen (Cheatum, 1934) , although other factors may also be 

October, 1901 nautilus 79 

involved, as many pulmonates appear never to surface (Russell- 
Hunter, 1953). The negative geotactic behavior (if such it 
should prove to be) described here may have been a response 
to oxygen loss, the latter perhaps resulting from sudden immer- 
sion in the mud medium. Possibly also a medium such as 
mud will stimulate receptors in all or a part of the snail's body 
and that negative geotactic movement results from the stimula- 
tion. This latter possibility is strengthened by the fact that, 
wliile no quantitative records were kept, snails often ceased 
their upward movement as soon as they had surfaced on the 
mud, although they still had no direct access to air, being 
below^ the surface of the water which was above the mud. 

Bouvier, E. L. 1887. Systems nerveux, morphologie generale et 

classification des gastropodes prosobranches. Ann. Sc. Nat. 

Zool. (7) iii: 1-570. 
Carthy, J. D. An introduction to the behavior of invertebrates. 

New York. 1958. 
Cheatum, E. P. 1934. Trans. Amer. Microsc. Soc. 55:318-407. 
Crozier, W. J. and Navez, A.E. 1930. Jour. Gen. Psychol. 5:3-37. 
Lacaze-Duthiers, H. 1872, Arch. Zool. Exp. Gen. 7:97-166. 
Pieron, H. 1928. Jour. Genet. Psychol. 55:3-17. 
Russell-Hunter, W. 1953. Proc. Roy. Soc, Edinburgh. ^^5 (ii) :- 



Louisiana State University in New Orleans 

Collections made during the past year have revealed three 
mollusks not previously reported in Louisiana. 

Gulella bicolor (Hutton) , known previously from this area 
by Dr. Harold Harry but not reported by him (personal com- 
munication, 1959), has been taken from 20 localities in and 
around New Orleans. Nowhere is it abundant; colonies appear 
to consist of few individuals which are found only after rains 
or in very damp habitats. 

A small colony of Praticolella griseola (Pfeiffer) was dis- 
covered in the yard of the Laclede Steel Corporation on France 

1 This paper is an outgrowth of a current investigation supported by a 
research grant, RG 7194, from the National Institutes of Health, Public 
Health Service. 

80 NAUTILUS Vol. 75 (2) 

Road in New Orleans. We have found it nowhere else even 
though extensive searches have been made. At this locality it is 
living in and around a pile of steel girders. The pile shows signs 
of having been there undisturbed for some time. It is part of 
a barge which used to make trips to Texas. Possibly this is how 
this colony came to be here. It is an interesting find in view of 
Pilsbry's statement (1940, p. 689) concerning Praticolella dis- 
tribution: "The herd of eastern Mexico and Texas is separated 
from that of the southeastern states by the Mississippi River and 
State of Louisiana, 300 miles more or less." 

Vaginula sp. (?) first came to our attention while collecting 
in Mobile, Alabama. There it is found in large localized colonies. 
Up to that time we had not discovered it in Louisiana (probably 
because of a dry period), but shortly thereafter we began finding 
it in New Orleans. To date it is known from 4 localities in the 

Since little is known of the mollusks of Louisiana, we consider 
it useful at this time to publish a list of the terrestrial mollusks 
which have been reported. 

In this list letters following localities indicate the source of 
the information as follows: (G) = Goodrich, (Ha) = Haas, 
(HH) = Harry, (H) = Hubricht, (?) = Pilsbry, (T) = Taylor, 
(V) = Vanatta, (Vi) = Viosca. Localities are parishes unless 
otherwise noted. 

Helix (ispefsa (Miiller) — Orleans, East Baton Rouge (HH, P, 

H. aperta Born— Orleans (HH, Vi).^ 

Otala vermiculata (Miiller) — Orleans. (P) ^ 

O. lactea Miiller— Orleans (HH) .2 

Polygyra septemvola febigeri (Bland) — Orleans, Vermillion (P) ; 
Plaquemines (HH). 

P. auriformis (Bland) — Orleans, Calcasieu (P) . 

P. leporina (Gould)— DeSoto (P) ; Grant (H) ; Catahoula (V) . 

P. triodontoides (Bland) — Calcasieu (P) . 

P. texasiana (Moricand) — DeSoto (P, HH) ; Grant, Rapides, 
Caddo, Orleans, Calcasieu, Nachitoches, Tensas (HH) .-^ 

P. dorjeuilliana Lea — Nachitoches, DeSoto, Bienville (P) ; Clai- 
borne (H) . 

2 These were no longer found in 1948 (Harry) and we have found none 
of them in Orleans Parish. 

3 Specimens may be P. triodontoides (Harry, 1951) . 

Odobci. HXil NAUTILUS 81 

Stcnoticnut lahio.stnii (lihmcl) — -Rapides, I'nioii (P). 

S. sIcnotreiiKi (Plcillcr) — Madison, Rapides (P) . 

S. nionodoii (illciac (Pilsbry) — DeSoio, Bienville, (Calcasieu 

(lype loealiiy (P): Evangeline, Allen, Vernon, Rapides (H) . 
S. iiionodon loiiu f) icrsoni Pilsbry — -Catahoula (V) . 
Mcsodon tliyroidus (Say) — Morehouse, I'laukiin. Calahoida 

(\'): Caleasieu, Grant, Ouaehita (H) . 
M. nijlfitus (S'ly) — Morehouse, Franklin (V) ; (Caleasieu (H, P) ; 

Lxangeline, Vernon, Grant, LaSalle (H) . 
Triodopsis vulluosa (Goidd) — C^alcasieu (P) . 
T. oagini Call — Naehitoches (P) ; Grant, Claiborne (H) . 
T. carolifiioisis (Lea) — -DeSoto, Caddo (P) . 
T. jostcri (F. C^. Baker) — Lake Pontchaitraiu (no parish given) 

/. (ilhohihris (Say) — included in the range given by Pilsbry — 
no dehnite localities listed (P): Oua( iiiia (H) . 

T. dn'csta (Gould) —DeSoto (P) . 

I Idplotrciiui concavum (Say) — LaSalle (H) . 

Hiilniiiiliis dealbatiis (Say) — DeSoto (P) . 

liiDiniiti decollata (Linnaeus) — Orleans (Ci, P, Vi) ; East Baton 
Rouge (HH). 

I junellaxis gracilis (Button) — Orleans (HH, P) ; East Baton 
Rouge (HH). 

I:iigl(indinti rosea (Feussac) — Orleans, Franklin, Iberia, Iber- 
ville. \V. Carroll (P) : Plaquemines (HH) . 

J., rosea biillata (Gould) — Orleans, St. Landry, Lake Pontchar- 
train (no parish given) , Iberia (P); Franklin (P, V) . 

liuconulus chersinus (Say) — State of Louisiana (P) ; Catahoula 
(V); Plaquemines (HH) . 

/:'. cliersi)ius trocJndus (Reinhardt) — Caddo (P) . 

E. chersinus dentatus (Sterki) — Catahcjula (\'); Rapides (P) . 

Guppya sterkii (Dall) —Bienville (?) . 

Retinella indentata (Say) — LaSalle (H); Morehouse, Catahoula 
(V) ; Plaquemines (HH). 

''. indentata paucilirata (Morelet) — State of Louisiana (P) ; 
C:iaiborne (P, H) ; Morehouse (V). 

Mesonipliix friabilis (W. G. Binney) — Morehouse (P, V). 

M. inilgatus H. B. Baker — Calcasieu, Iberia, Morehouse, Franklin 
(P) ; Vernon, LaSalle (FI); Morehouse, Franklin (\') . 

Haivaiia rniniscula (Binney) — included in the range given by 
Pilsbry — no definite localities given (P) ; Plaquemines (HH); 
.Morehouse, Catahoula (V) . 

I'entridens demissus (Binney) — Vernon, Grant, LaSalle (H) ■ 

/'. demissus brittsi (Pilsbry) — DeSoto, Bienville, Calcasieu (P). 

/'. intertextus (Binney) — Calcasieu, Claiborne. Evangeline, La- 
Salle (H) ; Bienville (P). 

82 NAUTILUS Vol. 75 (2) 

Zonitoidcs arboreus (Say) — State of Louisiana (P) ; Morehouse, 
Richland, Ouachita, Franklin, Catahoula (V) : Plaquemines 
(HH) . 

Limax fiaviis Linnaeus — Lincoln (T). 

Limax marginatiis Miiller — Caddo (HH). 

Deroceras sp. — Plaquemines (HH) . 

Deroceras Uicvc Miiller — East Baton Rouge. Ciaddo, DeSoto 

Angiaspira (iltern/iln (Say) — Catahoula (P, V): Morehouse, 
Franklin (V). 

A. alternata crassa Walker — Caddo, Franklin, Morehouse, De- 
Soto (P) ; Claiborne, Ouachita, Grant, Calcasieu, Evangeline, 
Vernon (H) . 

HeUcodiscus singlcyanus (Pilsbiy) — Morehouse (P, \^) ; Oua- 
chita (V). 

H. singleyaniis inermis H. B. Baker — Morehouse, Ouachita (P). 

Pliilomycus carolinianus (Bosc) — Calcasieu, Evangeline, Rich- 
land, Ouachita (H) . 

P. carolinianus flexuolaris Rafinescjue — Grant (P) . 

Fallifera marmorea (Pilsbry) — Vernon, Grant (H) . 

Oxylonia sallaena (Pteifter) — Orleans, DeSoto (P). 

Succinca nnicolor Tryon — Orleans (type locality) (P). 

.S'. grusx'etwri Lea — Rapides (P) . 

S'. lutcula Gould — Vermillion, Plaquemines (HH) . 

S. concordialis Gould — Concordia (type locality) (P) . 

Strobilops tcxasiana (Pilsbry and Ferriss) — Morehouse, Rapides 
(P. V). 

S. labyrinthica (Say) — Morehouse, Catahoula (V). 

S. aenea Pilsbry — State of Louisiana (P) ; Morehouse, Richland, 
Catahoula, Franklin; form micrompliala: Rapides, Morehouse, 
DeSoto, Richland, Franklin, Catahoula (P) . 

Gastrocopta contracta clnneana (Vanatta) — Orleans, Franklin 
(V) ; Morehouse, (P, V) ; Plaquemines (HH) . 

G. corticaria (Say) — Rapides (P). 

G. pentodon Say — Plaquemines (HH) . 

G. tappaniana (C. B. Adams) — INIorehouse, Franklin (V) . 

G. rupicola (Say) — included in the range given by Pilsbry — no 
definite localities given (P) : Plaquemines (HH) . 

G. pelliicida hnrdaccUa (Pilsbry) —Gulf States (P). 

Pupoides albilabris (C. B. Adams)— Gulf States (P) . 

P. modicns Gould — Placpiemines (HH) . 

Vertigo inilium Gould — Plaquemines (HH) . 

V. oscariana Sterki — Madison (P) . 

V. rugosula Sterki — State of Louisiana (P) ; Morehouse (V) . 

IJcUcina orbiculata (Say) — DeSoto, Jefferson, Orleans (P) . 

Bradybaena similaris — (Ferussac) — Orleans (Ha, HH, G). 

October, 1961 nautilus 83 

Literature cited 
Goodrich, C. 1940. Naut. 53 (3) : 105. 
Haas, F. 1945. Fieldiana. Zoology 31:2, 3-14. 
Harry, H. VV. 1942. Occ. Papers of La. State Univ. Marine Lab., 

No. 1. 

1948. Naut. 62 {\) : 20-24. 

1951. Naut. 64{?,): 96-99. 

Hubricht, L. 1956. Naut. 69(4) : 124-26. 

Pilsbry, H. A. 1939-41. Land Mollusca of North America (North 

of Mexico) Acad. Nat. Sci. Monographs 3, vols. 1 & 2. 
Taylor, W. E. 1899. Gulf Fauna and Flora Bull., 1 (3):69-73. 
Vanatta, E. G. 191 1. Proc. Acad. Nat. Sci. Phil.: 525-531. 
Viosca, P. 1928. Naut 41 (4): 139-40. 


Twenty-eighth Annual Meeting, June 20-23, 1961 

For the third time in its 30 year history, the American Mala- 
cological Union convened at the U. S. National Museum in 
VV^ashington, D. C. Attendance broke all records (150) while the 
four-day program rolled smoothly along thanks to careful plan- 
ning and the dedicated services of volunteer members of the 
National City Shell Club. Behind the scenes was Dr. Harald 
A. Rehder together with his henchmen Drs. Morrison and Rose- 
water, all veterans of past A.M.U. meetings, therefore anticipating 
the requirements of their guests well in advance. 

President Thomas E. Pulley presided over the academic ses- 
sions; it was due to his vigilance in enforcing time limits that 
every paper on an unusually full program was heard. 

An evening garden party at the beautiful home of Mr. and 
Mrs. A. Lothrop Lutrell in Rockville, Maryland was the high- 
light of opening day, and on Thursday the annual dinner was 
held at the Cumberland Country Club where food, service and 
sumptuous surroundings left nothing to be desired. 

The Executive Council met on Wednesday evening and at the 
annual business meeting on the following afternoon it was an- 
nounced that it had been found necessary to raise the annual 
dues one dollar per year with joint and life memberships to be 
advanced accordingly. (This advance will not become effective 
until 1962.) 

No site had been selected for the 1962 meeting, but a com- 
mittee has been appointed to consider the matter and it is ex- 

84 NAUTILUS Vol. 75 (2) 

pected that the time and place will be announced in the 1961 
report bulletin. 

The following slate of officers for the coming year was pre- 
sented by the nominating committee and elected by unanimous 
vote of the members present: 

President, William K. Emerson. 

Vice-president, Albert R. Mead. 

2nd Vice-president, Robert W. Talmadge. 

Secretary-treasurer, Margaret C. Teskey. 

Publications Editor, Morris K. Jacobson. 

Councillors-at-Large, John B. Burch, H. B. 

Herrington, Edward H. Michelson, Virginia Orr. 

The field trip on Friday was a 50 mile excursion to Scientists 
Cliffs on Chesapeake Bay where the clay banks of the Calvert 
formation yield a seemingly endless series of Miocene fossils. 
The trek back to the city was the final featine of the 1961 meet- 
ing, another to be added to the long list of memorable reunions 
of the AMU. — Margaret C. Teskey 


Beckianum — New genus (or subgenus of Leptinaria Beck, 
1837?) in Achatinidae (Subulininae) . Type species is Synopeas 
beckianum (Pfeiffer) H. B. Baker, 1945, Naut. 58:9\; 1947, Naut. 
61 (1) : plate 1, from Cordoba, Mexico. As stated by Zilch, 1959, 
Handb. Palazool. <5(2):351, Synopeas Jousseaume, 1889, is a 
homonym of Foerster, 1856. 

As pointed out in 1945:88, this unique species is very distinct 
from either Opeas (Ferussaciinae) or Lamellaxis and Leptinaria, 
sensu stricto (Subulininae) but, from its shell sculpture, it may 
belong near Pelatrinia Pilsbry, 1907 (1):324, which was proposed 
as a subgenus of Leptinaria. Pelatrinia has a larger, more conoid 
and imperforate shell, which carries its columellar cord into the 
aperture. — H. Burrington Baker. 

Maxwell Smith — Dr. Joshua L. Baily, Jr., writes from Ashe- 
ville, N. C, that Maxwell died there Sept. 12, and that a 
biographic sketch will be prepared for a future number. — H.B.B. 

Southern California slugs, additional locality records. — The 
following records are here listed because of their importance in 
establishing knowledge of distribution. 

Lehrnannia poiricri (Mabille) . Santa Cruz Island, Stanton 

October, 1961 nautilus 85 

Ranch headquarters (introduced) , 23-1-1960. Dr. Carey Stanton! 
Prol)abIy all records of Limax marginatvs iMiill. from North 
America, and certainly all California records of same should be 
referred to this species^. 

Anadenulus cockerelli (Hemphill). Kern County, soutli of 
sawmill at head of Tejon Canyon, Tehachapi Mts., altitude 
about 5,300 feet, 3-V-1958. W. O. Gregg, W. B. Miller! 

Ariolimax coliimbianiis stromineus Hemphill. Ventura County, 
Santa Paula Canyon, altitude about 2,700 feet, 29-V-1955. Ed 

Hesperarion niger (J. G. Cooper) . Kern County, west of Poso 
Creek and about one half mile north of Glenville — Woody road, 
8-VI-1945. Under oak log in damp meadow. W. O. Gregg, M. L. 
Walton! This is the record which Dr. Pilsbry- refers merely to 
"northern Kern Co." Tulare County, along Arrastre Creek, about 
3 miles east of White River, altitude about 2,200 feet, 26-V-1957. 
W. O. Gregg, W. B. Miller, W. B. Miller Jr.! 

Hesperarion hemphilli (W. G. Binney) . Santa Cruz Island, 
Canada del Puerto, 1.3 mile from Prisoners Harbor, 17-1-1960. 
Found under old log. W. B. M., W. O. G.! Santa Barbara County, 
along Salsipuedes Creek near Calif. Hwy. 1, about 3.4 miles 
south of junction with Hwy, 150, 10-III-1951. W. O. G., W. B. M., 
W. B. M. Jr.! San Luis Obispo County, near tributary of San 
Luis Obispo Creek at old ranch road .8 mile north of U. S. Hwy. 
101, south side Cuesta Pass, Santa Lucia Mts., 13-n-1960. W. O. 
G., W. B. M.! 

Binneya notnbilis J. G. Cooper. Santa Barbara Island, El 
Primero Canyon, 19-IV-1959. W. O. G., W. B. M.! Shells only, 
but with epidermis remaining on some shells. Very dry at time 
of collecting. — Wendell O. Gregg. 

Mass movement of a colony of the mud snail Ilyanassa. — In 
a survey of the marine life at Cape Ann, Massachusetts, a dense 
colony of Ilyanassa obsoleta was discovered on a mud flat border- 
ing a salt marsh about midway up Little River in 1933. Little 
River is a side channel to the Annisquam Tidal River, a marine 
inlet which divides the Cape Ann Promontory. The habitat was 
very soft, black, sticky mud. The colony in question occupied 
an area about 85 feet wide and extended outward from the edge 
of the salt marsh for some 80 feet downshore. In 1935 a series of 
quadrat counts gave a range of 25-61 with an average of 43 snails 

1 Quick, H. E., Proc. Mai. Soc. London, vol. 29, pt. 5, pp. 181-189; British 
Slugs'. Bull. British Museum, Zool., vol. 6, no. 3, p. 197-200. 

- I'ilsbi7, H. A., Land MoUusca of North .America, vol. 2, pt. 2, p. 724. 

86 NAUTILUS Vol. 75 (2) 

per quarter square meter. Measurements the next year indi- 
cated the same abundance. In 1956, quadrat counts ranged from 
7-36 with an average of 21 snails. Counts made the next two years 
gave a range of 9-53 and an average of 26 in 1957; a range of 
8-64 with an average of 26 in 1958. I observed, however, that a 
band of snails, usually not far from the marsh, had a density of 
about 3 snails per square inch. This band of maximum density 
seemed to shift with fluctuations of tidal levels between spring 
tides and neap tides. The 5 quadrat levels at which the counts 
were made and are quoted above, never fell on this narrow band 
of maximum density. In 1960 this colony was first visited on 
August 6. The water level at the time of observation did not 
permit an extensive sample, but abundance seemed to be about 
the same as in the past 4 years. On September 6, however, a 
single snail was found in the area formerly occupied by the 
colony described above. A search disclosed that the entire colony 
had moved to a new area some 150 feet northeastward, to the 
entrance of a ditch which cuts back into the marsh. No explan- 
ation seems apparent for this mass movement. — Ralph W. 
Dexter, Department of Biology, Kent State University, Kent, 


Pages in italics include new taxons 

Benthem Jutting, W. S. S. van. Non-marine Mollusca of the lime- 
stone hills in Malaya. Some zoogeographical remarks on the 
non-marine Mollusca of the Moluccas. Proc. Cent. & Bicent. 
Congress of Biology, Singapore, 1958: 63-68, 4 figs.; 164-169, 
1 fig. 

Chernin, Eli, Edward H. Michelson & Donald L. Augustine. 
Daubaylia potomaca, a nematode parasite of Helisoma tri- 
volvis, transmissible to Australorbis glabratus. J. Parasitology 
-^^.•599-607 (2 pis.). 

Michelson, Edward H. Chemoreception in the snail Australorbis 
glabratus. Amer. J. Tropic. Med. & Hygiene 9:480-487, 3 figs. 


Abbott, R. Tucker. The genus Lnmbis in the Indo-Pacific. Indo- 
Pacific Mollusca 1 (3) '.28 pp., 18 pis. (1 colored) , maps. 

Abbott, R. Tucker. How to know the American marine shells. 
Pp. 222, 12 col. pis., many text figs. Signet Key Book, P. O. Box 
2310, Grand Central Station, New York 17, N. Y. {Ibi +5«s 
postage) . 


Vol. 75 January, 1962 No. 3 


Deparimeni of Biological Sciences, Florida State University, Tallahassee 

Abbott (1954) has suggested that the opisthobranch gastropods 
Acteocina candei (Orbigny) and Retusa canaliculata (Say) may 
belong to the same species and may be merely geographic races or 
subspecies. The recovery of a large number of Acteocina candei 
from offshore waters in the vicinity of Cape Hatteras has per- 
mitted us to compare this form with an equally large number of 
superficially similar specimens of Retusa canaliculata collected in 
Pamlico Sound. Our observations and conclusions are presented 
in this report. Our comparisons led to the discovery of certain as- 
pects of the biology of Retusa canaliculata not previously re- 
corded, including the presence of a radula and the occurrence of 
non-pelagic reproduction in this species. 

Material. More than 300 specimens of Acteocina candei were 
recovered from the stomachs of seastars (Astropecten articulatus) 
collected from a sand bottom off Ocracoke Inlet, North Carolina, 
at a depth of 4 to 7 fathoms (Wells et al 1961) . Most of these 
were fresh specimens, from the soft parts of which radulae could 
be recovered. A similar number of specimens of Retusa canalicu- 
lata were collected from sand flats 2 to 3 inches below mean low 
tide along the eastern shore of Pamlico Sound 1 mile north of 
Avon, N. C. These two collecting stations are separated by a 
distance of less than 30 nautical miles. While one habitat is 
oceanic, the other is estuarine, with a greater range in tempera- 
tures, more rapid temperature changes, and reduced salinities. 

In addition to these specimens from North Carolina, shells of 
both species from different parts of their ranges have been ex- 
amined at the U. S. National Museum. 

Observations. These "forms" differ in several consistent charac- 
ters: Oxierall shell shape. Variations in the shell shape of Acteo- 
cina candei and Retusa canaliculata are shown in Figures 8-10 
and 1-3, respectively. Specimens from the oceanic collection {A. 



Vol. 75 (3) 

Figs. 1-3. Retusa canaliculata, showing variation in shell shape. Fig. 3. R. 
canaliculata with extreme development of the spire (protoconch eroded) . 
Fig. 4. Acteocina candei, protoconch. Fig. 5. R. canaliculata, protoconch. 
Fig. 6. R. canaliculata, apical view. Fig. 7. A. candei, apical view. Figs. 8-10. 
A. candei, showing variation in shell shape. (Fig. 10 shows juvenile condition.) 
(Figures were prepared with aid of a camera lucida.) 

January, 1962 



Fig. 11. Egg mass of Retusa ca?ialiciilala. Fig. 12- Radiilar teeth of 
Acleoci)ia candei. Fig. 13. Radiiiar teeth of R. canaliculata. Fig. 14. R. canali- 
culata, egg membrane containing veliger. Figs. 15-16. Shell of newly hatched 
R. canaliculata. (Figures were prepared with the aid of a camera lucida.) 

candei) are generally spindle-shaped, with an elevated spire and 
a tapered base, so that the shell is widest in the middle (Figs. 8 
and 9) . However, juvenile specimens of A. candei may appear 
more or less truncate (Fig. 10) , because of the lesser development 
of the spire. The aperture tapers gradually at its apical end. 
Specimens from the estuarine collection [R. canaliculata) are 
more cylindrical in shape, usually with lower spire, with a more 
rounded base, and with nearly parallel sides, so that the shell is 
shouldered and nearly as wide near its apical end as it is in the 
middle (Figs. 1 and 2) . However, there is some variation in the 
elevation of the spire in R. canaliculata, occasionally approaching 
(as in Fig. 3) the development characteristic of A. candei. The 
existence of individuals with such "intermediate" shell characters 
has no doubt led to confusion in identification. Specimens of both 
species that show a broad shallow depression in the middle of the 

90 NAUTILUS Vol. 75 (3) 

last whorl (as in Fig. 2) , producing a sinuous outer lip such as 
that figured by Marcus (1958, fig. 26) for Acteocina {:=Torna- 
tina) candei have been examined. A similar depression is shown 
in the shells of Retusa obtusa figured by Lemche (1948, figs. 43- 
53) , but in that species also this character is variable and not 
always present. Because of its occurrence in several similar species 
and the variability of its expression, such a shallow depression 
cannot be used as a reliable character for the separation of species. 

Protoconch. Whereas in A. candei the protoconch is obvious 
and projecting, in R. canaliculata only a small portion is visible 
as a small protuberance, and this is often eroded. The hetero- 
strophic protoconch of A. candei is composed of about one and 
one-half whorls (Fig. 4) set at an angle of 80° from the axis of 
the adult shell. It is about i/4 submerged in the first adult whorl 
(Fig. 7) . In contrast, the heterostrophic protoconch of R. canali- 
culata contains only about 3/^ of a whorl (Fig. 5) set at an angle 
of 80° from the adult shell axis. It is about % hidden by the first 
adult whorl (Fig. 6) . This difference in protoconchs would indi- 
cate that the planktonic stage is of moderate length in the larva of 
A. candei, while the planktonic stage is suppressed in the larva of 
R. canaliculata. 

Radular teeth. Using the technique described by Turner 
(1960) , the radulae were mounted for examination. The radular 
teeth of a specimen of A. candei 2.5 mm. in length are shown in 
Figure 12. The median tooth is 18 microns wide, 10 microns 
high, and bears 4 or 5 forward projecting denticles on each of two 
arches. The lateral tooth, with, an overall length of 45 microns, 
has a base 33 microns long and a moderately curved cusp 30 
microns long; it bears 5 to 7 sharp denticles on an expansion at 
the angle formed at the base of the cusp. By the same technique, 
a radula was discovered in R. canaliculata, the teeth of which are 
shown in Figure 13. For a specimen 3 mm. in length, the median 
tooth is 30 microns wide, 19 microns high, and is comprised of 
two arches, each bearing ten or eleven denticles. The lateral tooth 
is approximately 65 microns long, with a base 46 microns long 
and a strongly curved cusp 41 microns long that bears 16 to 20 
small denticles on a curved ridge. For each species, the radulae 
of at least 10 specimens were examined. The shape of the teeth 
and the distribution and relative size of the denticles clearly dis- 

January, 1962 nautilus 91 

tinguish the radulae of the two species. 

Because of the minute size of the radulae in these species, ade- 
quate radular preparations are difficult. Evidently, the radulae 
of both species escaped detection by early workers. Pilsbry (1893) 
treated both species as having no radula; however, later workers 
recognized the presence of a radula in Acteocina (Thiele 1931; 
Marcus 1958, as Tornatina) . Thiele (1925) could find no radula 
in Retusa canaliculata, and many workers (Marcus 1958, Zilch 
1959, and others) have utilized the absence of a radula to charac- 
teiize the Retusidae, and to place R. canaliculata in this family. 
The possession of a radula is used for the placement of Acteocina 
candei in the Scaphandridae. Since R. obtusa (Montagu) is the 
type species of Retusa, the discovery of a radula in canaliculata 
makes a re-examination of R. obtusa for the presence of a radula 
necessary' in order to clarify the position of canaliculata in 
opisthobranch classification. Until the presence or absence of a 
radula in R. obtusa is ascertained, we prefer to retain canaliculata 
in the genus Retusa. 

Although the possession of a slightly modified shell may reflect 
environmental differences between two habitats occupied by the 
same species, the differences considered above can scarcely be 
attributed to the environment. On the basis of this comparison, 
these forms should be regarded as biologically distinct species. 
This group of differentiating characters, particularly the differ- 
ences in protoconchs and radular teeth, provide unquestionable 
grounds for the recognition of two species. 

Acteocina candei occurs from North Carolina south to Argen- 
tina (Carcelles & Parodiz 1938), principally in offshore or oceanic 
habitats. In contrast, Retusa canaliculata occurs from Cape Cod 
to the West Indies, primarily in estuarine habitats. As Parker 
(1959) noted for the Texas coast, R. canaliculata is a character- 
istic inhabitant of enclosed bays of variable, low to intermediate 

Reproduction in Retusa canaliculata. Egg masses produced by 
Retusa canaliculata (Fig. 11) were collected on sand flats in Pam- 
lico Sound, where they were found to be relatively common from 
May to October. The egg masses of this species bear a resem- 
blance to one figured by Thorson (1946, fig. 151) as that of 
Philine scabra. They are composed of a spherical jelly mass about 

92 NAUTILUS Vol. 75 (3) 

2.0 to 2:5 mm in diameter, attached by a strand about 5 mm long 
to stems and blades of marine grasses, worm tubes, shell frag- 
ments, and other objects. Each mass contained a number of large 
eggs, each within its own egg membrane (Fig. 14) . The envelop- 
ing egg membranes are ovoid and 0.30 to 0.35 mm in length. 
Although eggs were usually found only in the enlarged spherical 
part of the mass, in some cases they were also contained in the 
attachment strand, as figured. A number of egg masses were re- 
moved to the laboratory for observation. Development progressed 
within the egg capsule, through a shelled veliger stage, to a min- 
iature crawling juvenile. These young snails were observed crawl- 
ing within their individual egg capsules, from which they event- 
ually emerged to crawl about the aquarium. These observations 
constitute the first record of non-pelagic development in Retusa 

The retention of the veliger stage within the egg capsule un- 
doubtedly serves to maintain this species in estuarine areas wdiere 
the net flow of water seaward could carry planktonic larvae away 
from an otherwise favorable environment. The evolution of 
such a mechanism would have considerable survival value for 
the larvae of species that inhabit estuaries. Indeed, non-pelagic 
development insures a more stable population size (Thorson, 
1950) . Such a non-pelagic development has been demonstrated 
for some species of echinoderms, polychaete annelids, prosobranch 
gatropods, and pelecypods (Thorson, 1950) . Generally, tecti- 
branchs have been considered to produce planktonic larvae. 

Lemche (1948) indicated that studies of the apical whorls of 
tectibranch gastropods may give reliable evidence about their 
larval development, and Thorson (1950) successfully applied 
similar correlations of apical shell form to the development of 
prosobranch gastropods. According to Lemche's standards, the 
blunt, relatively coarse protuberance formed by the protoconch 
of Retusa canaliculata fits the general protoconch morphology 
that is correlated with a reduced or suppressed planktonic devel- 
opment. Significantly, Lemche found the protoconch of Retusa 
obtusa to be wanting, and suggested that it probably exhibits a 
non-pelagic development. In his svnvey of Danish planktonic 
larvae, Thorson (1946) had not found larvae that he could assign 
to R. obtusa, although adults of that species had been known 

January, 1962 nautilus 93 

from the area studied. Evidently, R. obtusa shares with R. canali- 
culdta a non-pelagic type of reproduction. 

The relatively large size of the eggs and the prolonged repro- 
ductive period of R. canaliculata are features that have been 
correlated with non-pelagic development in other marine inverte- 
brates (Thorson 1950) . By increasing the chances of successful 
reproduction, these characters contribute to the effectiveness of 
non-pelagic development as an effective mechanism for the pro- 
duction of a new generation without the waste usually associated 
with plauktonic reproduction. 

Acknowledgments. These specimens were obtained and studied 
during the course of research supported by a grant (G-5838) from 
the National Science Foundation to Dr. I. E. Gray of Duke Uni- 
versity and aided by the Cape Hatteras National Seashore of the 
National Park Service. The authors wish to express their appreci- 
ation to Dr. Harald A. Rehder and Dr. J. P. E. Morrison of the 
Division of Mollusks, United States National Museum, for making 
available specimens and literature. 


Abbott, R. T. 1954. American Seashells. D. Van Nostrand Co., 
New York. 541 pp. 

Carcelles, A., and J. J. Parodiz. 1938. Physis, 12: 251-266. 

Lemche, H. 1948. K. Danske Vidensk. Selsk. Skr. (Biol.) , 5: 1-136. 

Marcus, E. 1958. Bol. Oceanogi-. S. Paulo, 7 (1-2) (1956) : 31-80. 

Parker, R. H. 1959. Bull. Amer. Assoc. Petrol. Geol., 43: 2100- 

Pilsbry, H. A. 1893. George W. Tryon, Jr., Manual of Conchology; 
structural and systematic. Order Opisthobranchia, vol. 15: 134- 
436. Philadelphia. 

Ihielc, J. 1925. Gastropoden der Deutschen Tiefsee-Expedition, 
Teil 2. Wiss. Ergebn. Deutschen Tiefsee-Exped., 17 {2) Opistho- 
branchia: 257-288, 348-352. Jena. 

1 93 1. Handbuch der systematischen Weichtierkunde, vol. /. 

G. Fischer, Jena. 778 pp. 

Thorson, G. 1946. Meddel. Komm. Danmarks Fiskeri- og Havun- 
dersogelser, Scr. Plankton •/(!): 1-523. 

1950. Biol. Rev., 25: 1-45. 

Turner, Ruth. 1960. Nautilus, 75(4): 135-137. 

Wells, H. W., M. J. Wells, and I. E. Gray. 1961. Biol. Bull., 

120 (2) . In press. 
Zilch, A. 1959. Gastropoda von Wilhelm Wenz; Euthyneura. 

Handbuch der Palaozoologie, 6(2): 1-200. Berlin. 

94 NAUTILUS Vol. 75 (3) 



U. S. Department of the Interior, Fish and Wildlife Service 

Bureau of Commercial Fisheries Biological Laboratory 

Woods Hole, Massachusetts 

A specimen of Cymatium caribbaeum CI. Sc T. was recently sent 
to me by Mr. Richard Spencer of Charleston, South Carolina. He 
had collected the example alive from a navigation buoy after it 
was brought in from its station off Port Royal, South Carolina, to 
the U. S. Coast Guard Base at Charleston for cleaning and servic- 
ing. Clench and Turner (Monograph of the family Cymatiidae 
in the western Atlantic. Johnsonia: 3 (36) : 206, 1957.) record 
the northernmost range for this species as Lake Worth, Florida. 
The buoy record thus extends the range northward along the 
Atlantic coast about 350 miles. 

The specimen is a thin-lipped juvenile, height 31.7 mm., width 
16.1 mm. This is the second species of this genus whose range has 
been considerably extended northward as a result of buoy sam- 
pling. The first was a juvenile Cymatium labiosum (Wood) 
which I collected in 1948 from a buoy brought in from off Cape 
Romain, South Carolina. This record appears in the monograph 
by Clench and Turner (1957) on page 202. 

The Cymatiidae are stenothermic and range widely in all tropi- 
cal waters. A probable long larval stage aids in their wide dispersal 
according to Clench and Turner. The larvae of the Cymatium 
species found on the South Carolina buoys probably chanced to 
be dispersed northward from Florida by way of rapid transport 
in the Gulf Stream. Caught in the coastal waters below Cape 
Hatteras, they finally settled on the convenient buoy surfaces. 

The normal longitudinal distribution of a species with pelagic 
larvae is controlled for the most part by the temperature ex- 
tremes that it can tolerate as an adult. The chance dispersal 
beyond the consistent range of such a species is probably common. 
When this occurs and when the larvae happen to settle in a favor- 
able habitat during a season in which the temperatures are also 
favorable, the organisms may survive and grow until such time as 

* Wells, W. Harry and \. E. Gray. The seasonal occurrence of Mytilus 
ediilis on the Carolina coast as a result of transport around Cape Hatteras. 
Biol. Bull.: 119 {3): 550-559, 1960. 

January, 1962 nautilus 95 

the temperature becomes lethal. The tact that an adult of the 
genus Cymatium has never been recorded from the bottom in off- 
shore Carolinian waters suggests that juveniles are winter-killed. 

Wells and Gray* recently reported on a cold water species, 
Mytilus ediilis, which commonly spreads beyond its normal south- 
ern limit at Cape Hatteras. The species is unable to survive sum- 
mer temperatures in the northern part of the Carolinian sub- 
province. However, the larvae of the fall spawning colonize this 
area after the water temperatures fall below lethal values. 

There is no need to stress the importance of accurate range 
records for ecological purposes. In order to increase the accuracy 
of range records, one must distinguish between that part of the 
range within which a species is able to maintain itself and propa- 
gate, and that portion, usually at the extremes, where it is unable 
to complete its life history. It is also important to record whether 
the specimen was alive or dead when collected; especially near the 
borders of its geogiaphic or bathymetric range. Knowledge of 
extreme range is particularly useful to indicate possible modifica- 
tions in the usual biota of an aiea should long-range environ- 
mental changes alter, even slightly, in any direction. 



Department of Zoology, The University of Kansas 

The extended ranges of the two species of mussels herein re- 
ported are the result of studies of the unionid fauna of Kansas 
from the years 1956 to 1959. Call (1885, 1886 and 1887) listed 
numerous species of unionids in Kansas without descriptions or 
illustrations, and Scammon (1906) described and illustrated the 
species of fresh-water mussels occuiTing in Kansas. Neither Call 
nor Scammon reported these two species as occurring in Kansas. 
A search of the literature indicates that neither has been recorded 
for Kansas. Catalogue numbers refer to the mollusk collection of 
the Museum of Natural History, The University of Kansas, 
Lawrence, Kansas. 

Arcidens confragosiis (Say) , KUMNH. 11083. This species was 
not collected by the author but was uncovered, quite by accident, 
in the museum collection. R. W. Reese obtained a single gravid 

96 NAUTILUS Vol. 75 (3) 

specimen of Arcidens confragosus from the Marais des Cygnes 
River, 3 miles east of Ottawa, Franklin County, Kansas, on Sep- 
tember 25, 1949. Unfortunately, only a half shell of this specimen 
remains in the museum collection. An empty shell of A. con- 
fragosus was found at a later date at the same locality, but has 
subsequently been lost. The half shell remaining in our collection 
measures 60 mm. in length and 42 mm. in height. A. confragosus 
was taken from a gravel shoal where the water ranged in depth 
from one to five feet. Subsequent attempts to collect additional 
examples have been unsuccessful. 

Because Utterback (1916:103) reported A. confragosus as oc- 
curring in northern and central Missouri and because the Osage 
River (Marais des Cygnes River in Kansas) flows through central 
Missouri, the presence of A. confragosus in Kansas is not surpris- 
ing. Possibly A. confragosus may occur in isolated areas in eastern 
Kansas not yet adequately sampled. 

Proptera capax (Green) , KUMNH. 10486. A gravid specimen 
oi Proptera capax was recovered by the author on August 27, 1956 
from the Neosho River, 7i/2 miles east and 1 mile south of Em- 
poria, Lyon County, Kansas. Simpson (1914:47) reported the 
westernmost range of P. capax as the St. Frances River in eastern 
Arkansas and two almost certainly invalid records from the Elk- 
horn and Blue Rivers, Nebraska. 

Inasmuch as the St. Frances River in Arkansas empties into the 
Mississippi River a short distance from the Arkansas River and 
inasmuch as the Neosho River is a tributary of the Arkansas 
River, it is not surprising that P. capax should occur in the 
Neosho River in Kansas. Although Utterback (1916:163) reported 
P. capax in Missouri, his records, as best as can be determined, are 
from the extreme eastern portion of Missouri. P. capax has not 
been reported to this date in those streams flowing from Kansas 
through Missouri to the Mississippi River, which is somewhat 
surprising considering the appearance of P. capax in the Neosho 

The single example of P. capax thus far known for Kansas is 
typical in most respects for the species. The nacre is a somewhat 
darker purple, and the shell is somewhat more elongate than 
specimens from the Mississippi River. 

The Kansas specimen measures 155 mm. in length and 101 mm. 

January, 19(32 nautilus 97 

in height. In length, this specimen exceeds the previously re- 
ported maximum length of P. capax by 12 mm. (Haas, 1941 : 261) . 
P. capax ^vas recovered from slowly moving water 3i/^ feet deep 
in a substrate of small rocks and sand having very little silt. 

Because A. confragosus is a species with unusual morphological 
features and not likely to be confused with other species, and 
because Call (1885, 1886 and 1887) and Scammon (1906) did not 
report this species in Kansas, it seems most likely that A. con- 
fragosus has made its appearance in Kansas after 1906. It is my 
opinion that P. capax may have occurred sparsely in Kansas for 
many years and may possibly have been confused by Call and by 
Scammon with Lampsilis ovata ventricosa (Barnes) . 

Literature Cited 
Call, R. E. 1885. Bull. Washburn College Lab. Nat. Hist., i.-48- 

1886. Bull. Washburn College Lab. Nat. Hist., 7.177-183. 

1887. Bull. Washburn College Lab. Nat. Hist., 2.11-25. 

Haas, F. 1941. Zool. Series, Field Mus. Nat. Hist. Chicago, 2^.-259- 

Scammon, R. E. 1906. Univ. Kansas Sci. Bull., 5.-279-373, pis. 

Simpson, C. T. 1914. A descriptive catalogue of the naiades of 

pearly fresh-water mussels. Detroit, 1540 pp. 
Utterback, W. L 1916. Amer. Midi. Nat. 4: (1-10) : 200 pp., 29 pis. 


Department of Biology, Kansas State Teachers College, Emporia 

freshwater limpet snails, constituting a distinct subfamily of the 
family Ancylidae, and limited in distribution to the southeastern 
United States. Walker (1917) recognized two "sections" or sub- 
genera of Rhodacmea based partly upon conchological characters, 
but principally upon features of the radula, to be discussed below. 
In connection with a continuing study of North American fresh- 
water limpets, I have reported briefly on the anatomy of one 
species of Rhodacmea (Basch, 1960) , and more recently have ex- 
amined the radulae of other species in an attempt to understand 
better this unusual genus of mollusks. I am indebted to Dr. Henry 

1 Supported bv grant G-14125 from the Xaiional Science Fouiidaiion. 
The genus Rhodacmea consists of about half a dozen species of 

98 NAUTILUS Vol. 75 (3) 

van der Schalie of the University of Michigan Museum of Zoology 
(UMMZ) for kindly making available to me the material used 
in this study. 

The radulae upon which Walker based his systematic con- 
clusions were prepared for him by Rev. H. M. Gwatkin of Cam- 
bridge, England. The specimens from which those slides were 
made were collected in the Tennessee and Coosa River drainages 
in Alabama, presumably by A. A. Hinkley or H. H. Smith, in the 
first decade of this century. In Walker's 1917 paper, and again 
the following year (Walker, 1918) , two figures illustrate the radu- 
lae of Rhodacmea filosa (Conrad) and R. rhodacme Walker. 
These drawings were the work of Mrs. Lydia M. H. Green. Al- 
though probably Mr. Walker examined the preparations himself, 
there is to my knowledge no direct evidence that he did so. 

I have examined 7 slides of radulae prepared by Rev. Gwatkin, 
including those from which Mrs. Green made her original illus- 
trations. The fact that his preparations have remained in ex- 
cellent condition for over half a century is a tribute to the skill of 
Rev. Gwatkin, who died in November, 1916. In addition, I have 
prepared 24 slides of radulae extracted from dried animals found 
within their shells in the UMMZ collection. The wide open aper- 
ture of the limpets and the lack of spiral coiling allowed easy re- 
moval of the dry animals without damage to the shells. After 
removal, the animals were soaked for several hours in a 0.5% 
solution of trisodium phosphate to soften the tissues, and the 
radulae then dissected out. Although large portions of these frail 
ribbons could be removed intact, I could not obtain an unbroken 
radula in this manner. The number of transverse rows of teeth 
per ribbon could therefore not be determined. In some cases, 
where bits of tissue adhered to the radula, the entire piece of 
ribbon was immersed briefly in a dilute solution of sodium hypo- 
chlorite (clorox) to clean it. Such treatment does not harm the 
teeth if performed judiciously, and is far more convenient than 
the traditional sodium hydroxide bath. All ribbons were rinsed 
in 70% ethyl alcohol and mounted in polyvinyl lactophenol, a 
mounting medium which dries fairly rapidly and does not require 
previous dehydration and clearing of the specimen. 

The following specimens were examined, all UMMZ numbers: 
A. Prepared radulae from the Gwatkin collection 

January, 1962 nautilus 99 

#946, 947, 948 — 7^. rhodacme Walker, Coosa River. 

•^9A9 — R. gwatkiniana Walker, Coosa River. 

:jijt950, 951 — R. clatior (Anthony), Tennessee drainage, Flor- 
ence, Alabama. 

#976 — R. filosa (Conrad) , Coosa River 
B, Radulae extracted and mounted from dry specimens 

#65998 — R. cahawbensis Walker, Little Cahaba River, Bibb 
County, Alabama. 2 specimens. 

#69213 — R. filosa, Coosa River, Talladega County, Alabama. 
8 specimens. 

#69221 — R. giuatkiniana, Coosa River, Coosa County, Ala- 
bama. 3 specimens. 

#69223 — R. giuatkiniana, Coosa River, Chilton County, Ala- 
bama. 4 specimens. 

#69237 — R. rhodacme, Coosa River, St. Clair County, Ala- 
bama. 7 specimens. 

All the identifications of species are those of Walker. A sixth 
species, Rhodacmea hinkleyi (Walker) was unavailable. 

The specific features utilized in establishing the two subgenera 
of Rfiodacmea are as follows (Walker, 1917) : 

"Section Rhodacmea, s.s. Shell elevated. Radula with a uni- 
cuspid central, which has the base triangularly expanded; laterals 
with the cusp of the mcsocone extending bvit little beyond the 
base and not overlapping the base of the central tooth." Included 
in this section were R. filosa, R. cahawbensis, R. elatior, and R. 

"Section Rhodocephala, n. sect. Shell depressed. Radula with a 
faintly bicuspid central which has the sides of the base straight 
and not expanded; laterals with the cusp of the mesocone extend- 
ing far beyond the base and overlapping the base of the central 
tooth." In this section were placed R. rhodacme and R. gxvat- 

From my study of these species I have come to the conclusion 
that there is no constant character by which it is possible to dis- 
tinguish species or species groupings within Rhodacmea. I submit 
that the characters utilized by Walker are largely artifacts, which 
are based upon two conditions — the amount of wear on the teeth 
examined, and variations in techniques of mounting the ribbon 
for study. In figs. A to C are shown 3 rows of teeth from a single 
radula of R. cahawbensis. Changes in the conformation of indi- 
vidual teeth as a consequence of the amount of wear may be easily 
seen. The central tooth, originally bicuspid, becomes worn down 
to a single cusp; the length of the mesocone may vary by 30% 



Vol. 75 (3) 


3 4 5 6 7 8 


r- ,.' "t 

Representative teeth from one radula of Rhodacmea cahaxubensis, UMMZ. 
#65998, prepared and mounted January, 1961, from a dried specimen. A, teeth 
near the front of ribbon, showing severe wear; B, teeth from an intermediate 
area, showing some wear; C, unworn teeth from a region near end of radular 
sac. Only central and 8 of the 11 teeth on one side are shown. Features marked 
by small letters in C ilhistrate variable characters: a, central tooth bicuspid 
or unicuspid; b, degree of overlapping between rows; c, amoimt of "shoulder- 
ing" on medial surface of first tooth; d, configuration of finer denticles on first 

January, 1962 nautilus 101 

within the same ribbon, and the amount to wliich it overlaps the 
base and the base of the central tooth are both dependent partly 
upon wear and partly upon the amount of pressure applied to 
the coverslip in making the slide. Depending upon the manipu- 
lation of the preparation, teeth may be rotated into various posi- 
tions, and may be separated from each other to a greater or lesser 
degree. The specific areas of greatest apparent variation in the 
sample studied are pointed out on figure C. 

Previous studies on the reliability of the radula as a taxonomic 
tool (Howe, 1930; Van Cleave and Richey, 1936) suggest that this 
structure alone is not a dependable criterion of specific differences 
within a genus (in those cases, Vivipariis) . In the present genus, 
although the radula is of the greatest importance in separating 
Rlwdacmca, sensu lato from other patelliform fresh-water mol- 
lusks, I feel that no reliance can be placed upon it in making 
critical taxonomic decisions within the genus. 

With regard to the elevated versus depressed shell, this may be 
a character useful for species differentiation, but I do not believe 
it to be of sufficient importance to separate subgenera, particularly 
when the radulae are indistinguishable in all species studied. I 
must conclude that the subgenera (or sections) Rhodacmea, s.s. 
and Rhodocephala are based largely upon artifact rather than 
biological differences, and therefore invalid. In a later paper I 
hope to discuss the relationships of Rhodacmea within the Ancy- 
lidae, and of the various species to one another. 

Literature cited 

Basch, Paul F. 1960. Naut. 73 (3) : 89-95. 

Howe, Sam W. 1930. Naut. •/-/ (2) : 53-63. 

Van Cleave, Harley J., and Emily M. Richey. 1936. Trans. Amer. 

Micros. Soc. 55(2): 223-229. 
Walker, Bryant. 1917. Naut. 31 (1) : 1-10. 
1918. A synopsis of the classification of the fresh-water Mol- 

lusca of North America, north of Mexico, and a catalogue of 

the more recently described species, with notes. Misc. Publ. 

Mus. Zool. Univ. Mich. 6: 1-213. 

row of teeth; e, lateral distance between teeth, and basal characters; f, pro- 
nounced notch present or absent on 4th tooth; g, ninnber of teeth per row 
(10, 11, or 12, with 1 ribbon of 13). All figures drawn with the aid of a 
camera lucida. 

102 NAUTILUS Vol. 75 (3) 



HELICODISCUS MULTiDENS, new species. Plate 7, D-F; Text fig. 1 
Shell discoidal, spire flat or nearly so; whorls 4i/2 to 5; pale 
greenish-yellow, dull, opaque. Umbilicus wide and shallow, show- 
ing all the whorls, occupying from 45 to 50% of the diameter of 
the shell. Whorls well roundfed, slowly increasing, the last slowly 
descending; sculptured with numeious, fine, spiral threads. Aper- 
ture lunate, the peristome somewhat thickened within. Within 
the last quarter whorl there are 3 pairs of teeth on the outer and 
basal walls. These teeth are radially elongate, raised on a heavy 
callous ridge, and separated by a rounded sinus. Alternating with 
these are 3 teeth on the parietal wall. These teeth extend out to 
about the center of the whorl, are about twice as broad as high, 
the ends are turned forward, the upper end more so than the 
lower. Of the 3 sets of teeth the center set is usually more fully 
developed than the others. As the shell grows, the teeth farthest 
within are absorbed and a new set added near the aperture. 

Height, 1.88 mm. Diameter, 4.75 mm. Umbilicus diameter, 2.22 
mm. Aperture height, 1.55 mm. 4.5 whorls. Holotype. 

Distribution: — Tennessee: Putnam Co.: in Jared Hollow Cave, 
3 miles northeast of Chestnut Mound (Thomas C. Ban", Jr., 
coll.) . DeKalb Co.; in Jim Cave, 1.5 miles southeast of Dowell- 
town, holotype 207798 U.M.M.Z., paratypes 17063, collection of 
the author; in Avant Cave, 1 mile east of Dowelltown. 

Helicodiscus multidens Hubricht; A, view of central pair of teeth. B, 
parietal tooth from above. C, diagram of tooth arrangement. 

Helicodiscus multidens is most closely related to H. triodus 
Hubricht. In H. triodus the teeth are smaller, and the sets of 3 
teeth are placed at irregular intervals in the last whorl, not 
crowded near the aperture. H. multidens is a rare snail, found 


Map 1. Distribution of Helicodiscus shimeki Hubricht as represented by 
specimens in the collection of the author and in the collection of the Uni- 
versity of Michigan. 

only in caves. Of the nine specimens so far collected only one was 
found alive. 

Helicodiscus shimeki, new species. Plate 7, A-C; Map 1. 

Shell discoidal, pale yellow, somewhat shining, translucent, 
spire flat or slightly convex. Umbilicus wide, shallow, showing all 
the whorls, occupying about 50% of the diameter of the shell. 
Whorls 5 to 6, well rounded, very narrow and slowly increasing; 
nuclear whorls with faint spiral striae; later whorls with numer- 
ous spiral threads. Aperture lunate, peristome thin. Within the 
last whorl there are usually three pairs of small conical teeth; on 
the outer and basal walls, the earlier teeth are absorbed. 

Height, 1.7 mm. Diameter, 4.2 mm. Umbilicus diameter, 2.1 
mm. Aperture height, 1.7 mm. 5.2 whorls. Holotype. 

Type locality: loioa: Delaware Co.: Backbone State Park, holo- 
type 207796 and paratypes 207797 U.M.M.Z., other paratypes 
13807, collection of the author. 

Helicodiscus shimeki is a species of the northern United States 
and probably southern Canada, although the author has seen no 
specimens from there. It ranges from Iowa eastward to northern 
New York. Map no. 1. 

Helicodiscus shimeki may be readily distinguished from H. 
parallelus (Say) by its more slender whorls and its broader, shal- 
lower umbilicus. It resembles H. salmonaceus W. G. Binney in its 
proportions, but the thread striae are coarser, and the umbilicus 
is not quite so broad. It stands somewhat intermediate between 
these two species. It is named in honor of the late Bohumel 
Shimek of Iowa City, Iowa. 



Vol. 75 (3) 

Map 2. Distribution of Helicodiscus notius Hubrifcht as represented by 
specimens in the collection of the author. 

Helicodiscus notius, new species. Plate 9, N-P; Map 2. 

Shell discoidal, the spire flat or nearly so; whorls 5 to 5i/^; pale 
yellowish, dull, translucent when young, becoming opaque with 
age. Umbilicus wide and shallow, showing all the whorls, occupy- 
ing from 40 to 45% of the diameter of the shell. Whorls well 
rounded, slowly increasing, sutures well impressed; sculptured 
with numerous spiral threads. Aperture lunate, peristome thin. 
Within the last whorl there are usually 2 or 3 pairs of conical 
teeth, one on the basal wall and one above it on the outer wall. 

Height, 1.66 mm. Diameter, 3.66 mm. Umbilicus diameter, 1.62 
mm. Aperture height, 1.04 mm. 5 whorls. Holotype. 

Type locality: Alabama: Jackson Co.: side of Keel Mtn., Paint 
Rock, holotype 207792 and paratypes 207793 U.M.M.Z., other 
paratypes 17588, collection of the author. 

Helicodiscus notius is found over most of the southeastern 
United States, ranging from the Gulf north to Virginia and west 
to Missouri and Oklahoma. See map no. 2. 

Helicodiscus notius differs from H. parallelus (Say) in being a 
little larger due to the additional whorl; in having a broader, shal- 
lower umbilicus; and in having the thread striae on the embry- 


NAUTILLfS 75 (.H) 




HoldlNlK's. \-( . Hclirodiscus shhnrhi Huhiidii. HI'. H. innllidrii.s Huhrichl. 

NAUTILUS 75 (3) 






II()h)l\|)LS. (,-1. flrli((>ilis(iis noliiis spcciis Hiihiitlil. | .<: R. //. UadciuH'ciis 
Ihihiiilu. L c<: M, //. jucksoiii Hubricht. 

January, 1962 nautilus 105 

onic whorls more strongly developed. It is most closely related to 
H. eigenmanni Pilsbry, from which it differs in being smaller and 
in having a larger umbilicus. 
Helicodi,sc;u.s notius specus, new subspecies. Plate 8, G-1. 

Differing from the typical form in being without the internal 
teeth in shells laiger than two and one-half whorls. The last half 
whorl slowly descends to the aperture, and the peristome is often 
much thickened. 

Type locality: Kentucky: Barren Co.: in Burnette Cave, 0.6 
mile west of Park City, holotype 207794 and paratypes 207795 
U.M.M.Z., other paratypes 17027, collection of the author. 

Helicodiscus notius specus is known only from the type locality. 
In this cave, it was feeding on the guano of the cave-cricket, 
Hadenoecus subterraneus (Scudder) , in company with Carychium 
stygium Call. Typical H. notius is occasionally found in caves 
also, but only where leaves have been washed or blown in. It 
never has become a guano feeder. 

Helicodiscus saludensis (Morrison) 

Gastrodonta (Clappiella) saludensis Morrison, 1937, Proc. Biol. 
Soc. Wash. 50-58, PI. 4, figs. 1-4. 

Clappiella saludensis (Morrison) , Pilsbry, Land Moll. N. 
Amer. II, p. 433, fig. 233. 

I believe that this species should be placed in the genus 
Helicodiscus. It is so like H. parallelus (Say) in its general ap- 
pearance that one needs a lens to distinguish it. H. fimbriatus 
Wetherby has a similar pattern of internal teeth, although they 
are not as regularly alternating. 

Troglodiscus, new subgenus. 

Shell with numerous, fine, revolving, epidermal fringes; but 
without the revolving ridges of Helicodiscus s.s. There are no 
internal teeth at any stage of growth. 

Type species: Helicodiscus barri. 
Helicodiscus barri, new species. Plate 9, R-T. 

Shell small, pale gieenish-yellow, subtranslucent, thin, de- 
pressed; spire low, convex; whorls 4 to 41/2, well rounded, sutures 
well impressed. Umbilicus moderately large and deep, contained 
about 3 times in the diameter of the shell. Sculpture of numerous, 
fine, revolving epidermal fringes. There being from 40 to 50 of 
these fringes on the last whorl. Aperture lunate, peristome thin. 
There are no internal teeth at any stage of growth. 

Height, 1.8 mm. Diameter, 3.9 mm. Umbilicus diameter, 1.4 
mm. Aperture height, 1.3 mm. 4.5 whorls. Holotype. 

Distribution: Tennessee: Dickson Co.: in Columbia Caverns, 
2 miles southwest of Van Leer, holotype 207799 and paratypes 

106 NAUTILUS Vol. 75 (3) 

207800 U.M.M.Z., other paratypes 17446, collection of the 
author. Davidson Co.: in Bull Run Cave, 2.5 miles northwest 
of Scottsboro. 

Helicodiscus barri is known only from the total darkness of 
caves. It was found feeding on raccoon dung. It is named for its 
discoverer, Thomas C. Barr, Jr. 

Helicodiscus hadenoecus, new species. Plate 8, J-K; PI. 9, U. 

Shell very small, yellowish, opaque, subdiscoidal; whorls 4, 
well rounded, sutures well impressed. Umbilicus moderately 
large and deep, contained from 21/2 to 3 times in the diameter of 
the shell. Embryonic whorl smooth, later whorls, with fine, re- 
volving, epidermal fringes, which are found on both the upper 
and lower surfaces and in the umbilicus. There are from 30 to 40 
of these fringes on the last whorl. Aperture lunate, peristome 
sinuous, not thickened. There are no internal teeth at any stage 
of growth. 

Height, 1.2 mm. Diameter, 2.7 mm. Umbilicus diameter, 1 mm. 
Aperture height, 1 mm. 4 whorls. Holotype. 

Distribution: Kentucky: Barren Co.: in Beckton Cave, 0.5 mile 
northwest of Beckton. Tennessee: DeKalb Co.: in Avant Cave, 
2 miles east of Dowelltown. Van Buren Co.: in McElroy Cave, 
1.5 miles northeast of Bone Cave P. O., holotype 207801 and 
paratypes 207802 U.M.M.Z., other paratypes 17444, collection 
of the author. White Co.: in Indian Cave, 2.5 miles southeast 
of Quebeck. Jackson Co.: in Hargis Cave, 1 mile north of Gran- 
ville (Thomas C. Barr, Jr., coll.) Alabama: Madison Co.: in 
Aladdin Cave, 7 miles northeast of Maysville. 

Helicodiscus hadenoecus is known only from the total dark- 
ness of caves. It feeds on the guano of the cave-cricket, Hade- 
noecus subterraneus (Scudder) . It is related to Helicodiscus 
barri, differing in its smaller size. Dead shells from which the 
epidermis has been removed show no sculpture other than weak 
growth lines. They show no ridges like those in H. parallelus. 
Dead shells resemble those of Hawaiia minuscula (Binney) . 
Helicodiscus punctatellus Morrison. 

The type locality for this species is in, not near. Whites Cave, 
near Mammoth Cave. The author found dead shells fairly com- 
mon in Whites Cave, but no living specimens. The author visited 
nearly every accessible cave in Mammoth Cave National Park 
without finding this species in any other cave. It apparently 
occurred only in Whites Cave and is now extinct. 
Helicodiscus jacksoni, new species. Plate 8, L-M; PI. 9, Q. 

Shell very small, depressed, thin; whorls 4 to 414, well rounded, 

January, 1962 nautilus 107 

sutures well impressed, spire low but convex. Umbilicus wide, 
occupying about 40% of the diameter of the shell. Embryonic 
whorl smooth, later whorls with irregularly spaced growth wrin- 
kles, lower surface of last whorl nearly smooth. Aperture lunate, 
peristome simple. 

Height, 1.11 mm. Diameter, 2.44 mm. Umbilicus diameter, 
0.96 mm. Aperture height, 0.77 mm. 4.5 whorls. Holotype. 

Type locality: Maryland: Dorchester Co.: subfossil, in shell 
mound near beach, 0.5 mile west of Elliott, holotype 207803 and 
paratypes 207804 U.M.M.Z., other paratypes 22476, collection 
of the author. 

Helicodiscus jacksoni is most closely related to Helicodiscus 
singleyanus inermis H, B. Baker, with which it was found. It 
differs in its distinctly larger umbilicus and in the more prom- 
inent growth wrinkles on the upper surface of the last whorl. The 
whorls are a little more slender and not so tightly coiled. The 
aperture is smaller. Only dead shells were found, and most of 
these were chalk white. Some were fresh, these were translucent, 
with a very pale yellow shell. 

It is named in honor of Ralph W. Jackson, who discovered this 
interesting shell locality. 
Paravitrea roundyi Morrison. 


These two species seem out of place in the genera to which 
they have been assigned. Possibly when they are better known, 
they will prove to belong to the genus Helicodiscus. 



The type species of the genus Engina Gray, 1839, is Engina 
zonata Gray, 1839, by subsequent designation (Gray, 1847) . This 
species has not been correctly identified by authors because it 
was not figured; the color pattern is uncommon for the species 
but principally because, in error. Gray described it as II/2 inches 
long (misprint?) . 

Mr. S. P. Dance, of the British Museum, recently showed me 
a lot of two shells, BV. 113, marked Enzina (sic) zonata Gray in 
Gray's handwriting. They came from Mr. Guilding and the 
locality was "West Indies". Neither the adult nor the juvenile 
shell shows white knobs around the base, the common color-form 



Vol. 75 (3) 

Radula of Engina turbinella (Kiener) , ANSP. 195841, Buccoo Reef, Tobago 
Island, B. W. I. app. 970 X- 

of the species, but both are easily identified as the common west- 
ern Atlantic species, Engina turbinella (Kiener, 1836) . The adult 
shell measures 11.5 mm. in length (approximately 14 inch) and 
is here designated lectotype of Engina zonata Gray. I figure it 
(Plate 10, fig. A) with a similarly colored specimen from St. 
Thomas, Virgin Islands, ANSP 34755 (fig. B) . 

The lot from which the lectotype is taken is the one referred 
to by Tomlin (1928) as "a possible type lot" and misidentified 
as Engina leucozona (Philippi, 1844). My figure A is from the 
photograph published by Bartsch (1931) . However, I have in- 
cluded a scale showing in the original photograph but not previ- 
ously published because it explains why Bartsch did not recognize 
the species. 

Bartsch saw only the photograph of the specimen and had no 
notes on the size of the shell except the scale photographed with 
it. Reading, "Axis: II/2 inches" in Gray's description he pre- 
sumed the scale was an inch rule and translated the size to 27 mm. 
He made no attempt to identify this outsized Engina. 

Although, from the misspelling of the generic name noted 
earlier, this lot of Engina zonata evidently was not labeled by 
Gray in 1839 (he misspelled Enzina in 1842 and 1847), it prob- 
ably was in his hands when he designated the type of the genus. 
It is also the only Gray material of that name which has come to 
light. Except for the discrepancy in size, the specimens agree well 
with Gray's description. 

Fortunately, the better-known name for the species, Engina 

NAUTILUS 75 (3) 



Holotypcs. N!'. Ilclicodiscus noliiis Ilubviclit. (). H. jacksoiii Hubrichi. 
R- 1 , H. harri Hubrichi. U, H. hadrnocrtis Hiibiichl. I'holographs bv John V,. 
iV.iuh. I'liiversity of Michigan. 

NAUTILUS 75 (3) 


4 i: n 

:- 'f^y. 

/ . '» 

3 4 

Suci'lla lupillii.s: Figs. I -L'. spec inuii lioni liaiut- (I'.S.X.M. (at. No. 12{i,'i(j) . 
Figs. 3-4, original figures in .\Iaiiini. ITSO. pi. 124. ligs. I IPS, 111). (All ligines 
approximateh natural size.) 

I'ig. .V. LctlotNpc oi luio/iui zoitdla (.ia\. H\ . Ii;f. liiitisli Miisciiin. West 
Indies. .Scale in nun. Fig. M. l-:>ioi)in linb'niclla (Kiencr), AN.SP, 'H7."i,"). St. 
Ihomas, Virgin Islands. 

January, 1962 nautilus 109 

turbinella ( Kiener) precedes E. zonata by 3 years. Therefore, the 
type species of the genus Engina Gray is Engina zonata Gray, 
1839 (non Reeve 1846) = Purpura turbinella Kiener, 1836. 

References cited 
Bartsch, Paul. 1931. Proc. U. S. Nat. Mus., 79 (15) : 1-10, pi. 1, 

fig. 6. 
Gray, J. E. 1839. Zoology of the Blossom, pp. 112-113. 

1847. Proc. Zool. Soc. London, p. 133. 

Tomlin, J. R. leB. 1928. Nautilus, -/2 (2) : 40. 



U. S. National Museum, Smithsonian Institution 

The genus Nucella was instituted by Roding (Musuem Bol- 
tenianum, 1798, p. 130-131) for the 5 species listed below. Fol- 
lowing each name I have given what I believe to be the present 
correct or proper assignment of the species in question: 

reticulata Cancellaria reticulata (Linnaeus) . 

moschatellina Species dubia. 

macina Nomen nudum. 

lapillus Paralagena smaragdula (Linnaeus) 

theobroma Nucella lapillus (Linnaeus) . 

It will be noted that I have been unable to identify one of the 
species: N. moschatellina. Roding rechristened with this name 
Gmelin's Buccinum laeve, repeating the latter's reference to Mar- 
tini's figures in volume 4 of the Conchylien-Cabinet (1780, pi. 
124, fig. 1150). Martini's description of this species in the ac- 
companying text (Conchylien-Cabinet, vol. 4, 1780, pp. 36-37) 
does not quite agree with the figures, and later authors have 
been unable satisfactorily to identify the species. Gray (Zoology 
Beechey's Voyage, 1839, p. 126) placed it in his new genus 
Bullia and gave as a synonym sqiialida King. However, the finely 
striated sculpture and grooved interior of the aperture mentioned 
by Martini speak against this assignment. The same criticism can 
be leveled against Pfeiffer's (Krit. Register Martini & Chemnitz 
Konch. — Kat., 1840, p. 33) suggestion that the species may repre- 
sent the juvenile stage of Buccinum undatum Linnaeus. 

Clench, in his treatment of the western Atlantic species of 
Purpura and Thais (Johnsonia, vol. 2, no. 23, 1947), has dis- 
cussed on pages 85-86 the problem of the type of Nucella, but 

110 NAUTILUS Vol. 75 (3) 

because 1 have come to a conclusion differing from his I am 
placing my views on record, as I feel that the nomenclature of 
this common boreal group should be settled. 

Nucella was first actively revived by Dall in 1909 (Prof. Paper 
U. S. Geol. Survey, no. 59, 1909, pp. 46-48, 50) as a subgenus of 
Thais Roding. Clench (I.e.) is coiTect in stating that Dall did 
not specifically designate a type, although he undoubtedly in- 
tended to do so in his outline of the classification on page 50, 
where he cites T. [hais] lapillus after the name Nucella s.s. Later 
on, in his paper on the members of this group from northwestern 
America (Proc. U. S. Nat. Mus. 49, no. 2124, 1915, p. 558) he 
did definitely designate N. lapillus (Linnaeus) as type. As Clench 
has pointed out, this is an invalid designation since Linnaeus' 
species was not included by Roding in his genus. Therefore, 
Suter (Manual of New Zealand Mollusks, 1913, p. 425), Iredale 
(Trans. Proc. New Zealand Inst. 47, 1915, p. 472), and Grant 
and Gale (Mem. San Diego Soc. Nat. Hist. 1, 1931, p. 716) were 
wrong in accepting Dall's type designation. Wenz (Handbuch 
der Palaozool., vol. 6, Pt. 1, Lief, 6, 1941, p. 1123) credits Dall 
with citing filosa Gmelin (lapillus L.) as type, while Winkworth 
(Proc. Malac. Soc. London 26, 1945, p. 141), credits him with 
giving N. theohroma as the type species. Both these statements 
are also erroneous, since Dall, on the one hand, merely listed 
filosa Menke (not filosa Gmelin) as an example ( 1909: 46) , and, 
on the other hand, as Clench states, makes no mention at all of 
Roding's theobroma. 

Clench, however, has overlooked Winkworth's earlier type 
designation (Journ. of Conch. 19, 1932, p. 229) , where he un- 
equivocally gave N. theobroma Roding as type, without crediting 
it to Dall. This is the earliest valid type designation for Nucella 
that I have been able to discover. 

The principal problem seems to be the identity of Nucella 
theohroma which Clench concludes must be considered a species 
dubia. Roding refers his species to Bucciniim filosum Gmelin 
(Syst. Naturae, Ed. 13, vol. 1, Pt. 6, p. 3486), and follows the 
latter author in citing two figures in Martini (Conch.-Cabinet, 
vol. 3, 1777, p. 433, pi. 121, fig. 1113-1114) as illustrating the 

Clench feels that these figures are unidentifiable, but to me it 

January, 1962 nautilus 111 

seems to be evident on a number of counts that they represent a 
form of the common Nucella lapillus (Linnaeus) . First of all. 
Martini himself, in the accompanying text, considers the shell a 
variety of lapillus, stating that it differs only in the color pattern; 
the locality he doubtfully gives as East Indies. Secondly, Roding 
has renamed the same shell again on page 132 of the "Museum 
Boltenianum" as Nossa ligata, again referring to Buccinum 
filosum Gmelin, and to the same figures in Martini. Here it is 
placed immediately next to Nassa rudis Roding, which as Clench 
points out, is merely a new name for lapillus. Thus it seems cer- 
tain that Roding regarded Buccinum filosum as being close to 
lapillus Linnaeus. Finally, an examination of the U.S.N.M. col- 
lection of Nucella lapillus revealed a specimen, collected in 
France, that very closely resembles the figure given by Martini. 
On plate 10 I give a photograph of this specimen together with 
a reproduction of the figures of Martini. Considering the varia- 
bility of the species, I am sure that specimens could be found 
that match the figure of Martini even more closely. 

Polytropa Swianson, 1840 (Treatise Malacology, pp. 80, 305) 
is the next available name, and the one that Clench (op. cit.) 
used instead of Nucella. Swainson included in this genus Purpura 
lapillus, which Gray in 1847 (Proc. Zool. Soc. London, pt. 15, 
p. 138) designated as type. Thus Polytropa, it is true, is the first 
taxon that was fixed unequivocally on lapillus, but is a name 
that has been little used, and Nucella, as explained above, is 
clearly available for this group, and is earlier by 42 years. 

A survey of the literature over the past 15 years, using the 
Zoological Record as a source, reveals that 18 authors used the 
name Thais, without subgeneric assignment, for the species 
lapillus, 4 (all American) , used Polytropa (as a subgenus of 
Thais) , and 14 authors used Nucella, all but two as a distinct 
genus, a procedure that Clench (op. cit.) suggested probably 
should be followed for this group. 


Chicago Natural History Museum 

Widely distributed in Europe, Asia, and north Africa, Cionella 
luhrica (Miiller) is generally spread over the North American 

112 NAUTILUS Vol. 75 (3) 

continent from northern Alaska to northern Mexico (Pilsbry, 
1948) . The species is quite ubiquitous within its range. Although 
there has been little work done on population density, collectors 
generally report that C. Inbrica is difficult to obtain in t|uantity. 
AVhenever large aggregations have been noticed, the observation 
has been thought of sufficient importance to place on record. 
Such recorded observations are summarized in the accompanying 
table, to which the following may now be added. 

On October 27, 1960 Miss Maidi Wiebe, Staff Artist, Geology 
Department, Chicago Natural History Museum, observed a large 
aggregation of C. lubrica, probably totaling several hundred in- 
dividuals, on the sidewalk in front of her apartment in Oak 
Park, Illinois, at about 7:30 A.M. The pavement was moist from 
a brief shower the preceding evening, although the adjacent 
ground was not particularly wet, the moisture having permeated 
into the soil. Air temperature was about 47° F. No other organ- 
isms were observed in association with the snails. At 5:00 P.M. 
Miss Wiebe noticed that about a fourth of the number of snails 
observed that morning were still present on the pavement. At 
7:30 A.M. the following morning the snails had almost vanished. 
The temperature was somewhat higher, about 54° F., and the 
pavement was dry. Heavy rains fell on each of the following two 
days. A few snails were observed on the walk for a period of 
about 14 days, none being noticed thereafter. 

The available data are insufficient to warrant drawing final 
conclusions. Some tentative suggestions do seem in order, and 
the summation will at least serve to call attention to the need 
for additional observations and to indicate the type of informa- 
tion required. 

Aggregations of C. lubrica have been observed in widely scat- 
tered localities in North America, from Massachusetts and Penn- 
sylvania to British Cohnnbia. I have been unable to find refer- 
ences to any observed aggregations in the Old World. Quick 
(1954) states that C. lubrica is much more abundant that its 
close relative C. lubricella Porro in England. 

The aggregations have occurred from late May or early June 
to late October. The majority of observations have been made in 
late summer or fall. Krull and Mapes (1952) have noted that 
these snails became increasingly numerous after September 1 in 

January, 1962 nautilus 113 

central New York. They ascribed the greater numbers of snails 
observed in association with rocks at this time to the increased 
warmth and protection afforded by the stones. 

1 he observed aggregations have generally occurred either dur- 
ing or preceding a storm. This would tend to support Hender- 
son's (1905) suggestion that they are a response to physical fac- 
tors. However, Mapes and KruU (1951) found collecting difficult 
even after a rain. Additional data on response to various environ- 
mental conditions is given in Krull and Mapes (1952) . That such 
aggregations are for the purpose of reproduction has been sug- 
gested (Anon., 1946; Pilsbry, 1948) . Possibly both physical and 
biological factors are involved, perhaps depending upon the 

A hint of a cyclic phenomenon is contained in Clapp's (1914) 
statement that "in 25 years collecting in the Sewickley Valley 
[Pennsylvania] I have found this species decidedly rare." Long- 
time observations on populations of C. lubrica are necessary. I 
plan to keep tab on the Oak Park population during succeeding 

Stephens (1918) found large numbers of these snails fastened 
by their secretions to the walls of a building, all with the apex 
of the shell directed downwards. This suggests some sort of 
oriented reaction (oriented behavior is discussed in Allee et al., 
1949 and Warden et al., 1940) . Possibly this behavior was a re- 
sponse to excessive ground moisture, Mapes and Krull (1951) 
noting that C. lubrica "is quickly activated and stimulated to 
climb up when submerged." About 100 of the Oak Park snails 
were kept under my observation for several days in a small petri 
dish, the bottom of which was covered by moistened newspaper. 
There was a noticeable tendency for the snails to concentrate in 
the center of the dish, in fact to form a pile of snails, when the 
paper was quite moist. As the paper dried out the snails tended 
to move out and distribute themselves more uniformly over the 
bottom. No mating activity was noticed during the relatively 
short observation period. Oriented behavior is discussed in Allee 
et al. ( 1949) and in Warden et al. (1940) . 

These aggregations of C. lubrica are in contrast to the difficulty 
experienced by Mapes and Krull in collecting this snail in central 
New York for experimental studies. Some of their collecting was 



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January, 1962 nautilus 115 

done near Cazenovia, the general locality of Henderson's (1905) 
observations. Mapes and KruU found it necessary to resort to the 
use of traps consisting of wet gunny sacks spread over the ground 
in suitable areas in order to obtain sufficient specimens. These 
traps, examined two or four times weekly, resulted in only 26 
specimens per trap even in favorable sites and under favorable 
conditions. These authors are of the opinion that the requisite 
factors for large populations of C. lubrica are moist soil, almost 
constant shade, and a large quantity of suitable vegetation. 

The materials required for the culture of C. lubrica are inex- 
pensive, but such cultures do require a considerable amount of 
attention. Detailed instructions may be found in Mapes and 
Krull (1951) . There is here an excellent opportunity for some 
interested amateur with the time at his disposal to make some 
very worthwhile observations on the biology of this snail. 

Allee, W. C. 1931. Animal Aggregations. Univ. Chicago Press, 

Chicago, 111. 
Allee, W. C, et al. 1949. Principles of Animal Ecology. W. B. 
Saunders Co., New York. See especially Chapter 23, pp. 393- 
Anon., 1946. Naut. 60:72. 

Binney, Amos. 1851. The Terrestrial Air-breathing Mollusks of 
the United States and the Adjacent Territories of North Amer- 
ica, vol. 1. Charles C. Little Sc James Brown, Boston. Edited 
by A. A. Gould. 
Binney, W. C, and T. Bland. 1869. Land and Freshwater Shells 
of North America, Part 1, Pulmonata Geophila. Smith. Misc. 
Coll., 194. 
Clapp, Geo. H. 1914. Naut. 28:96. 
Fromming, Ewald, 1954. Biologic der mitteleuropaischen Land- 

gastropoden. Duncker & Humblot, Berlin. 
Gould, A. A. 1870. Report on the Invertebrata of Massachusetts. 

2nd ed. Wright 8c Potter, Boston (1st ed., 1841, not seen). 
Henderson, J. B. 1905. Naut. /c9;109-l 10. 
Mapes, C. R. 1951. Cornell Veterinarian ^/. -382-432. 
Mapes, C. R., and W. H. Krull. 1951. Ibid, II. Cornell Veter- 
inarian ■/;. -433-444. 
Pilsbry, Henry A. 1948. Land Mollusca of North America (north 
of Mexico) . Vol. 2, part 2. Phila. Acad. Nat. Sci., Monogr. 
No. 3. 
Quick, H. E. 1954. Proc. Malacol. Soc. London 50.-204-214. 
Stephens, T. C. 1918 Science, n.s. 43:21]. 

Warden, C. J., et al. 1940. Comparative Psychology, vol. 2, Plants 
and Invertebrates. Ronald Press & Co., New York. 

IHi NAUTILUS Vol. 75 (3) 


The name Holopodopes (plural ot holopod-ops) is proposed 
here lor an infraorder ot the suborder Sigmurethra, to include 
the achatinoids (Achatinidae and Spiraxidae) , Streptaxidae, 
rhytidoids (Acavidae proper -{- Caryodinae, Haplotrematidae, 
Rhytididae and Chlamydephoridae) and orthalicoids (Urocop- 
tidae and Orthalicidae or Bulimulidae) . In a systematic ar- 
rangement, they should precede the Aulacopoda (arionoids, 
limacoids and testacelloids) and the restricted Holopoda (poly- 
gyroids, oleacinoids and helicoids) . Some of the reasons for the 
separation of the holopodopes have been outlined in an earlier 
paper (1955) . 

The symbols used for Puerto Rican localities were explained 
recently (1961). 

Opeas piimilum (Pfeiffer) . Es2 (coconut plantation) . 

The older name, Helix hannensis Rang, 1831, from West 
Africa, which may have been this species, now can be dropped as 
obsolete. The other widely distributed Opeas, O. pyrgula 
Schmacker & Boettger, which has less arcuate growth striae, 
probably also occurs in Puerto Rico, although it has not been 
reported, unless the older, but obsolete name, Stenogyra alabas- 
trina Shuttleworth, from near San Juan, represented it. The more 
typical Ferussaciinae, Cecilioides (Geostilbia) aperta (Swainson) 
-|- gundlaclii (Pfeiffer) and C. (Karolus) consobrinus (Orbigny) , 
found by van der Schalie, 1948:50, 51, also have been distributed 
widely by commerce. 

Lamellaxis (Leptopeas?) micra (Orbigny) and var. rnargari- 
taceus (Shuttleworth). Es2 (coconuts), Pnl (yams), Wr2 (cof- 
fee) , Wr3. All my dry shells (more in alcohol) nearer the 
smoother form, which is not limited to Puerto Rico; the obsolete 
name, Stenogyra gompharium Shuttleworth, from near San Juan, 
may have been based on the typical one. 

Martens, 1877:345, reported the Cuban L. (Leptopeas) pahi- 
dinoides (Orbigny) from Aguadilla (Ww), but I got neither 
it nor the ubiquitous L. (Allopeas) gracilis (Hutton) , which may 
mean they are more limited to cidtivated areas. 

Lamellaxis (s. s.) rnonodon (C. B. Adams) and var. opalescens 
(Shuttleworth). Pnl, Pr3, Wr2, Wn; my only dry shell (Wr2) 
the imperforate form, which also occurs in Jamaica. 

January, 19G2 nautilus 117 

Obeliscus (Stenogyra) terebraster (Lamarck) . Deep in humus, 
Er3, 5, Prl-4, 6, Wr2, 3, 1800-4000 ft.; typical larger form. 
O. (S.) TERKBRASTER RARisiNiSTER, new subspecies. Enl, Es3, 4, Jnl 
(type locality), Pnl; lowland form smaller throughout, not sim- 
ply with less whorls. The type, Pilsbry, 1906 (I6a) :pl. 32, fig. 31, 
a sinistral shell (ANSP. 59320 from R. Swift) of this variety. It 
measures: 14.4 mm. by 28 (4.1 mm.) with lOi/^ whorls. 

O. (S.) swiftianus (Pfeiffer) . Ps2 (attains length of 10.5 mm., 
with 8.2 whorls) & Ws (smaller) ; this species has gone around 
the world. 

O. (Pseudobalea) hasta (Pfeiffer) . Under leaves on ground and 
in moss on tree trunks; Er2, Jnl, Pn, Pr3, 6, Wr, 100-3400 ft. 

Since Balea dominicensis Pfr., 1853, from "I. Haiti," is identi- 
fiable only from its inclusion by Pfeiffer himself in the synonymy 
of O. hasta, the slightly older name is clearly obsolete, even if 
dated from Pilsbry, 1906 (16a) :272. 

Subulina octona (Bruguiere) . On ground under dead leaves; 
Ee, En, Es2-4, Jn, Js, Pn, Pr2, 3, Ps, Wn, Wrl, Ws, Ww; 0-3400 ft. 

The obsolete Stenogyra (Subulina) acicularis Shuttleworth, 
based on one shell from near Fajardo, seems to be represented in 
some eastern lots (Ee, Es2) from near the seashore by occasional 
dwarfed shells (with fewer whorls, but sexually mature) which 
have more deeply etched, growth striae, that do crenulate the 

Leptinaria unilamellata ( Orbigny) . Widely reported by van 
der Schalie, 1948:56, probably from cultivated places. 

Very unfortunately, I, 1945:91, adopted Orbigny's prior name 
before the more widely used L. lamellata (Potiez & Michaud) 
would have been saved by the 50 year "rule." Recently, Aguayo, 
1961:94, added the other, equally ubiquitous, but more terrestrial 
L? (Beckianum) beckianum (Pfeiffer) , which already was known 
from both the Virgins and Haiti. 

Austroselenites (Zophos) alticola H. B. Baker. Deep in leaf 
humus, Er2-4, and probably Erl and 5 (too young for exact 
identification) ; certainly above 2500 ft, on El Yunque and proba- 
bly Luquillo Mts. above 2000. 

A. (Z.) co77color (Ferussac) . Also terrestrial. En 3, 4, Jn, Pn, 
Prl, 3, 4, Wr3, and probably Pr 2, 6, Wn, Wr2 (too young) ; cer- 
tainly on lowlands, and apparently represented by a smaller, 
darker race up to 4000 ft. in the Cordillera Central, but my 
material is too scanty for certainty. 

The widely transported streptaxid, Diaphera (Huttonella) 

118 NAUTILUS Vol. 75 (3) 

bicolor (Hutton) was found by van der Schalie, 1948:68. Dia- 
phera Albers, 1850, of which Diaphora Albers-Martens, 1860, is 
a homonymic emendation, is prior to Gulella Pfr., 1856, and 
D. bicolor is the type (Nevill, 1878) of Huttonella Pfr., also 
1856, of which Indoennea Kobelt, 1904, is a subjective synonym. 

Brachypodelln (s. s.) riisei (Pfeiffer). Lowlands, En, Jn, Pn, 
Wn, Ws. 

B. (s. s.) pallida ("Pfeiffer" Philippi) . Lowlands, En, Jn; Ps2 
(beattyiy, VVs (approaching beattyi) . 

B. (Brevipedella) portoncnnn (Pfeiffer) . En, Jn, Pn, \Vn, Wr2, 
0-2300 ft. 

These 3 species of Brachypodella may occur within a few feet 
of each other on limestone rocks, but B. portoricana, much the 
poorest climber, was the only one found (Wr2) far from the 
limestone rim, and B. pallida, the best climber, was the only 
species collected at the driest station (Ps2) , where it was buried 
quite deeply under rocks. Besides their difference in habits, these 
shells from Ps2 do average smaller and commonly their last 
whorls are less widely solute (some almost adnate) but the "dif- 
fers" in riblets seem individual variation. Some are about the 
size of typical beattyi Clench, 1951:251, fig. 3, from Mona Island; 
one with all 17 whorls measures 10.3 mm. (over all) by 2.1 (not 
including aperture) . Many more were found intact but even a 
little shaking rendered them decollate. 

Pseudopineria viequensis (Pfeiffer) . In holes and under over- 
hang of cliffs, on or beneath the limestone outcrops of the north- 
ern rim, commonly in shaded places where Brachypodella was 
absent; Jnl, 2. Foot whitish; ommatophores translucent, cylin- 
dric, with black eyes; inferior tentacles small but certainly pres- 
ent; sole unizonal, with 1 or 2 waves that involve its full width. 

Macroceramiis rnicrodon (Pfeiffer). Pilsbry, 1903 (25a) : 1 15- 
1 16, pi. 24, figs. 71-72 (ANSP. 251 17 & 25115, from R. Swift) ; van 
der Schalie, 1948:map 62, pi. 7, figs. 2a, 2b. En, Jn; growth 
threads narrower than their interspaces (much more so on sub- 
apical whorls) and irregularly spaced on later whorls. 

The obsolete M. johannis Pfeiffer appears indistinguishable, 
and Aguadilla falls within the range of the typical subspecies. 

M. rnicrodon, var. sliultleiuorthi (ALirtens) , 1877:352 (?) , 
without exact locality. Pilsbry, fig. 74 (ANSP. 2572, from Bland) ; 
van der Schalie: fig. 2c. Ws: largest 18.7 mm. long with 12 whorls. 
Ps2: 1 adult 18.8 mm. long (decollate) and 2 immature, one of 
which approaches loeryi in growth threads. 

M. rnicrodon loeryi Jacobson, 1955. Ps2 (near type locality) ; 

January, 1962 nautilus 119 

one adult and 1 immature, empty shells; look as i£ started life like 
var. a, but assumed later whorls ot var. b, which see. Adult 14.1 
mm. by 55 (7.7 mm.) with 7 whorls remaining. 

M. rnicrodun loeryi, var. a. Ps2, 3. Shells about size and form of 
typical microdon; one (Ps2) measures: 14.1 mm. by 36 (5.0 mm.) 
with 11.2 whorls; ranging to 17.2 mm. long with 12.6 whorls; 
without bluish tinge when alive; 1 shell unicolor, without white 

AI. microdoji loeryi, var. b. (?) M. shuttleworthi Martens, 1891: 
132, Penuelas. Under bunch grass, but climbing 4 to 6 ft. during 
rain; near Tallaboa (Psl). Shell with growth threads, especially 
on later whorls, broader than their interspaces and quite evenly 
and closely spaced; and those on subapical whorls less widely 
spaced than in typical microdon. Predominant color of shell light 
sky-blue on living examples but now (1960) almost completely 
faded to whitish (opaque) with narrow, light brownish (trans- 
lucent) growth (axial) bands, as were empty shells found nearby 
(one of last unicolor, without whitish patches) ; with almost no 
trace of basal angulation in many; largest 21.1 by 37 (7.8 mm.), 
minor diam. 33 (6.9 mm.) with 13.2 whorls; smallest 19.8 mm. 
long wuth 1 1 whorls remaining. Living animal light slate color, 
darker near sole and on ommatophores; mantle collar dark 
brownish gray, Avith minute light spots; sole unizonal, with 1 or 2 
locomotor waves across its full width. 

In 1939, var. b. was considered a subspecies of M. miaodon 
(Cf. van der Schalie:96) but I hesitated to name it because of the 
doubt about the true M. shuttleworthi, which Martens later 
located at the nearby Penuelas. The above presents the old notes, 
with a few changes; typical loeryi and var. a were included, as 

Microceramus (s. s.) guanicus H. B. Baker. On rocks near 
ground; only known from type locality (Ps2) and Mona Island 
(Clench, 1951). 

Bulimulus (s. s.) guadalupensis (Bruguiere) . On ground and 
tree trunks up to 5 ft., Ee, Es2 (coconut plantation), Es4, Jnl, 
Ps4 (garden) , Wwl (garden) . 

For this widely disseminated pest, van der Schalie, 1948:87, re- 
turned to B. exilis (Gmelin). 

B. (s. s.) diaphanus (PfeiflPer) . More terrestrial, Pnl, Pr2, 4, 
Ps2, 3, Wn, Ws; 0-3100 ft. Shell epidermis rufous (typical) to 

This may be a native species, although described originally from 
St. Thomas. Pilsbry's, 1897 (9b) : 46-47, transference to this species 
of the Puerto Rican records of the now obsolete B. fraterculus 

120 NAUTILUS Vol. 75 (3) 

("Fer." Potiez & Michaud, 1838) is accepted; certainly nothing 
like the original (P. & M.) figures has been found since. The 
last name was nude in Ferussac, 1821, livr. 11:54, and in Beck, 
1837:67, although Beck questioningly (?) referred it to Helix 
tenuissirna Ferussac, 1832 (-|- Orbigny, 1835?). 

Drymaeus (Mesembrinus) virgulatus (Ferussac, 1821) and color 
form (?) liliaceus (Fer., 1832). Arboreal, edges of Psl (30% 
Uliaceus) and Ps2 (45% liliaceus) ; empty shells seen elsewhere 
but only near cultivated places. 

The separation of these two lots into liliaceus (complete ab- 
sence of brownish color) and virgulatus (with even a few streaks, 
up to axial and/or spiral bands of variable continuity) seems 
highly arbitrary. Confessedly, most of the "lilies" have a less 
elongate (Roding) form and a slightly swollen last whorl, but 
they include the most elongate^ shell in the series, and typical 
virgulatus covers almost the entire range of shell form. These 
may be hybrids (of an introduced with a native form?) but they 
cast doubt on any specific separation, even if it has continued 
for 130 years. 

D. (M.) multilineatus (Say) ; Cf. form osmenti Clench and/or 
eboreus Grimshawe. Along road to Wn. One shell from living 
adult resembling in form, texture and coloration those of Pilsbry, 
1946(11) :27, figs. 15b, with similar columellar blotch (absent in 
thicker and chalkier D. virgulatus) and slightly wider, sutural 
stripe, but lacking all other dark bands except 4 (plus traces) 
axial streaks near aperture, and showing no signs of characteristic 
bluish color near apex. Another shell too broken and bleached 
to be sure of its form, with similar columellar patch and axial 
streaks, but without sutural stripe. 

Neither fits the obsolete Drymaeus hjalmarsoni (Pfr.) , from 
"near Manati" (between Jn and Pn) , which, as Pilsbry (1899) 
suggested, seems to have been close to his (1946) Leptodrymaeits, 
and even might be the subsequently named D. dormani (W. G. 
Binney) . 

Simpulopsis (Eudioptusf) psidii (Martens, 1877). Not ob- 

From field notes, the young shells of Platysuccinea from the 
Cordillera Central, which approach it in height of spire, were 
confused with this arboreal species, and thus no special search 

1 More so than beattyi Clench, 1951: fig. 6, but less so than his type, figs. 
4 & 5. 

Januai7, 1962 nautilus 121 

was made tor it west of Utuado (between Wr6 and Pnl) . How- 
ever, a shell ot the sagdid at the size of S. psidii (5 by 5 mm.) 
has one less whorl (2 vs. 3) ; being small, mature 5. psidii might 
be found only near the end of a long rainy period. Pilsbry, 
1902 (31) :lxvii, included both Eudioptus Albers-Martens (not 
a homonym) and Platysiiccinea in the bulimulid Simpulopsis, 
but no geophile systematist in this century has guessed either to 
be a "Bulimulus." The type species of Eudioptus, in which 
Martens placed his "Bulimus" psidii, is Brazilian, but a dweller 
on guava leaves might be carried by commerce. 

Gaeotis nigrolineata Shuttleworth. Mainly on leaves of palms, 
but also on those of Cecropia and other trees, Er3, 4, 5; in axils 
of palm pinnae, Wr3; from Luquillo Mts. to western end of the 
Cordillera Central; 2000-3000 ft. Living animal (Er) 3 by li/g 
inches (76 by 38 mm.) ; yellowish gieen, but variable; dorsum 
of foot slaty with yellow middorsal stripe (flavolineata) which 
often becomes double anteriad; and often with frosty, whitish 
patches (like albopunctulata.}) ; sides and inferior tentacles 
lighter, but sole often with orange margin on anterior I/3 
(flavolineata); ommatophores blue green, but eyes lighter; mantle 

(over shell) pea green; often with (internal) black streaks 
visible (nigrolineata). 

Very flimsy shell "sigaretiform" but thin lower surface of 
apical whorls present and containing those of animal, so that 
separation in preserved examples means damage to one or the 
other (shells still on most of mine) . Lower surface of apical 
whorls broken away from all 6 shells studied by Pilsbry, 1899 

(35):227-231, as follows: ANSP. 4613, one labeled "albopunctu- 
lata" and one (4614) "nigrolineata" from Humacao (Bland) . 
ANSP. 26052, type lot of malleata; 2 shells from "near San Juan," 
from Swift, collected in 1855. ANSP. 4959, one juvenile labeled 
"flavolineata" by Bland (not by Pilsbry) from "Luquillo." 
ANSP. 26051, labeled "nigrolineata" from "Pto. Rico" (Swift). 

G. nigrolineata is the type species of Gaeotis, by subsequent 
designation of Kobelt, 1880, 111. Conch.:264. Although only pre- 
liminary dissections have been made as yet, all my material looks 
like one variable species, but none of it comes from the lowlands 

(secretive during dry weather?) . On El Yunque, this bulimulid 
slug was fairly common, but its coloration blends with the leaves. 

Additional references (See 1961) 
1961 (I), Naut. 74:142. 1961 (5), Naut. 75:64. 
Aguayo, Carlos G. 1961, Caribb. J. Sci. /.•89-106. 
Jacobson, Morris. 1955, Archiv f. Molluskenk. 84:91-99. 

122 NAUTILUS Vol. 75 (3) 


Julia Anna Gardner, known around the world for her work in 
stratigraphy and molluscan paleontology, died after a long ill- 
ness on November 15, 1960. She was a geologist with the United 
States Geological Survey for 32 years. 

Julia Gardner was born in Chamberlain, South Dakota, on 
January 26, 1882. Her undergraduate studies were done at Bryn 
Mawr College where she received the Bachelor of Arts degree in 
1905 and the Masters degree in 1907. Miss Gardner entered The 
Johns Hopkins University in the fall of 1907, the first woman 
regularly admitted to the Department of Geology. After receiving 
her doctoral degree in 1911, she continued there as Assistant in 
Paleontology until 1915. Her studies of the Late Cretaceous 
Mollusca of Maryland and other smaller groups were published 
by the Maryland Geological Survey in 1916. In 1915 she began 
work on the Miocene Alum Bluff Group of Florida under con- 
tract with the United States Geological Survey. 

During World War I, the broad and specific humanitarianism 
which characterized her life impelled her to go to France with 
the Red Cross in 1917. She saw strenuous service as an Auxiliary 
Nurse and after the war with the American Friends in the 
devastated areas of France. 

Upon her return to the United States in 1920, Miss Gardner 
joined the Geological Survey and took up again her professional 
career, which was to become long and distinguished. 

She was the author of over 40 reports. Pre-eminent among 
these are: 
The molluscan fauna of the Alum Bluff group of Florida: LJ. S. 

Geol. Survey ProL Paper 142, 1926-1947. 
The Midway group of Texas: Texas LTniv. Bull. 3301, 1933 

The Mollusca of the Tertiary form;ttions of northeast Mexico: 

Geol. Soc. America Mem. 11, 1947. 

Their scientific value was equaled by their usefulness in the 
field of petroleum geology, and they are standards of reference 
in the economic investigations of Tertiary strata of both North 
and South America. Her published works are foundation stones 
and bench marks in Coastal Plain stratigraphy and paleontology 

January, 1962 nautilus 123 

that insure Julia Gardner a high place with the pioneers in the 
geology ot the region. 

Among the societies of which Miss Gardner was a member are 
Phi Beta Kappa; Sigma Xi; the Geological Society of America, 
of which she was a fellow and vice president in 1953; the Ameri- 
can Association of Petroleum Geologists; and the Paleontological 
Society, which she served as president in 1952. She was a charter 
member of the American Malacological Union and was widely 
known and loved by collectors everywhere. 

She was an official delegate of the United States to the Inter- 
national Geological Congress in Madrid in 1926, and to the In- 
ternational Congress in Moscow in 1937. 

Julia Gardner made a great contribution in her chosen field of 
geology; an equally great contribution in the field of human rela- 
tionship may be exemplified by her encouragement of the Japan- 
ese scientists while on a tour of duty in Japan after World War II. 
They confirm that her kindness and generosity heartened them 
to carry on their researches. In her wide circle of friends, few do 
not have cause to remember some kindness, some encouragement, 
some tangible assistance, timely, but given unexpectedly. Julia 
Anna Gardner will live long in the hearts of her friends. — Druid 


PoMACEA PALUDOSA IN ALABAMA — In 1943, John Richardson 
II collected Pomacea paludosa (Say) at Gainesville, Florida, and 
introduced it into ponds on his farm 7 miles north of Jackson- 
ville, Florida. From here, in 1953, Frank Lyman introduced it 
into a small pond on the shore of Gantt Lake, 5 miles northeast 
of Gantt, Covington Co., Alabama. It soon escaped into Gantt 
Lake, and is now to be found along the shore for a mile in either 
direction from Lyman's pond. — Leslie Hubricht. 

S\ic<:iriea Indiana Pilsbry. 

Succinea Indiana Pilsbry, 1905, Nautilus, 19: 28. 

Siiccinea vaginocontorta Lee, 1951, Occas. Papers Mus. Zool. 
Univ. Mich. no. 533, pp. 1-7, PI. 1-2. 

A careful examination of the anatomy and shell of topotypes 

124 NAUTILUS Vol. 75 (3) 

Succinea indiana showed no character which could be used to 
distinguish it from S. vaginacontorta. The same twisted vagina 
is found in S. indiana. S. vaginacontorta Lee must be placed in 
the synonomy of S. indiana. 

Additional records for Succinea indiana are as follows: Xorth 
Carolina: Craven Co.: sandy roadside, 4.4 miles west of New 
Bern. South Carolina: McCormick Co.: below Clark Hill Dam. 
Georgia: Richmond Co.: roadside, 1600 Gordon Highway, Au- 
gusta. Alabama: Montgomery Co.: waste ground, Atlanta High- 
way and Forest Hills Ave., Montgomery. Dallas Co.: near Ala- 
bama River, opposite Selma. — Leslie Hubricht.' 

ViviPARUs suBPURPUREUs IN OKLAHOMA. — A single Specimen, 
33.0 mm. in height and 32.0 mm. in greater diameter, of this 
species was discovered in a small, mud-bottomed, heavily-vege- 
tated stream at the south boundary of Tom, McCurtain County, 
Oklahoma on 16 July, 1961. The spire was greatly eroded. Al- 
though this specimen represents a new record for Oklahoma, it 
is not a particularly surprising one. The Austroriparian Biotic 
Province extends into Oklahoma at this point but it is separated 
from the main body of that ecological division by the Red River. 
Viviparus subpurpureus (Say) is a fairly common species in ad- 
jacent Texas and Louisiana but apparently has met some diffi- 
culty in traversing the river barrier into Oklahoma, as I have 
collected rather extensively in this part of the state and have not 
found the species before. — Branley A. Branson, Dept. of Biol- 
ogy, Kansas State College, Pittsburg. 

Helisoma anceps transported by a giant water bug. — On 
5 May, 1961, I collected a giant water bug, Lethocerus americanus 
(Leidy), which had been attracted (at night) to a lOO-watt 
mercury lamp on a lawn at the University of Michigan's Erwin S. 
George Reserve, Livingston County, Michigan. The bug was 
carrying on its scutellum a specimen of Helisoma anceps 
(Menke) . The snail was alive and appeared to be immature, 
measuring 5 mm. high and 7 mm. long. Giant water bugs of this 
species and a similar species, Benacus griseus (Say) , frequently 
come to lights in early spring and later in the summer; evidently 

1 Accidentally omitted from Naut. ?'':()(). — H. B. B. 


January, 1962 



there is much movement between ponds, presumably associated 
with mating behavior. Any animal that happened to become 
attached to a bug would be similarly transported. During 1959- 
61, I examined about 500 of these bugs, but this is the only 
occasion I found an attached snail. But among several hundred 
hydrophilid beetles similarly attracted to light, I found 3 with 
immature leeches attached to their elytra. The leeches at first ap- 
peared dead, but quickly became active when placed in water. In 
view of frequent speculation as to how totally aquatic animals 
colonize isolated ponds, these records may be of interest. — D. F. 
Owen, Museum of Zoology, University of Michigan, Ann Arbor. 

The anatomy of glyphyalinia junaluskana. — Glyphyalinia 
junaluskana (Clench & Banks) generally has been treated as 
a subspecies of G. sculptilis (Bland) . However, a study of the 
genitalia shows that it is quite distinct. The following descrip- 
tion of the lower genitalia is based on a specimen collected at 

Glyphyalinia junaluskana (Clench & Banks). Three views of terminal 
genitalia, drawn with aid of camera lucida. Scale line = 1 mm. 

126 NAUTILUS Vol. 75 (3) 

4700 ft., 0.6 mile southeast of Beech Gap, GTaham Co., North 

Penis (fig.) rather short and stout, club-shaped, basal portion 
nearly white; upper portion pale brownish, globose. Penial 
retractor short and stout, inserted at apex of penis. Epiphallus 
parallel to penis, long and stout, as large as the penis; upper 
end deeply cleft into two equal lobes; lower portion pale brown, 
becoming darker at upper end. Spermatheca narrowly obovate, 
not clearly differentiated from the thick duct. Vagina and free 
oviduct short. 

It differs from G. sculptilis in being without the penial appen- 
dix of that species and in having the epiphallus divided into two 
distinct lobes. It apparently belongs in section Glyphyalinia. s. s 
rather than in section Glyphognomon H. B. Baker. — Leslie 


CoRBicuLA FLUMiNEA (Miiller) from the Ohio River — On a 
field trip last August 30, 1960, to the Ohio River just east of 
Fort Massac and Metropolis in Massac County, Illinois, I col- 
lected 1 1 recently dead specimens of Corbicula (Corbicula) flu- 
minea. They were 15-20 feet from the north bank in 6'' of water. 
The bottom was more gravel than sand with very little mud 
present. The areas on either side were muddy bottoms and 
about one foot deep, yet no specimens were found. 

Corbicula fiinninea, a native of eastern Asia, has not been 
reported from the northeast or central states as of this writing; 
therefore, I can mention those collected at Metropolis, Illinois, 
as new in this part of the country. Sinclair and Ingram (1961, 
Naut. 7-/:l 14-1 18) reported the species' presence in the Tennessee 
River in Hardin County, Tennessee. Previously, the species was 
known only from the south to north western states, i.e., Arizona, 
California, Idaho, Nevada, Oregon and Washington (op. cit.) . 
Corbicula flumiyiea seems to be widely distributed over the 
United States. Records of its further spread are to be expected 
with additional collecting. 

Since I am not an expert on foreign sliells, the specimens were 
submitted to Dr. Fritz Haas of Chicago Natural History Museum 
for identification. They are now Chicago Natural History Mu- 
seum No. 103678. Sincere appreciation is extended to Dr. Haas 
for his assistance. — Frederick R. Fechtner. 



Vol. 75 APRIL, 1962 No. 4 


By a. a. OLSSON and D. R. MOORE 

In 1949, Mr. Leo A. Burry described and figured (Shell Notes, 
December, 1949) , a most unusual Strombus under the name of 
S. canaliculntus, reported as having been taken in a fish-trap set 
at a depth of 175 fathoms off Hopetown, Elbow Key in the 
Bahamas. The whereabouts of the type is unknown. The figure 
shows an immature specimen in which the lip is not yet fully 
formed. Subsequent to the description of the species, the junior 
author saw 3 additional specimens which Burry had obtained 
from the same source. Two specimens were half grown and 
smaller than the holotype, the third was fully matured with a 
flared and thickened outer lip. This last specimen, a topotype, 
is in the reference collection of the Marine Laboratory of the 
University of Miami (Accession no. 30:1839). It measures 242 
mm. in length. (Plate 11.) 

The full-grown shell has a high, broad, conic spire of 8 or 
more whorls, each edged with a sharp, peripheral keel or carina, 
the main surface of each spire-whorl smooth, except for the fine 
lines of growth. There are no axial riblets at any stage. The 
peripheral keel becomes more strongly emphasized on the shoul- 
der of the body-whorl and at its termination on the edge of the 
outer lip it forms into a deep notch. The wall of the shell is 
quite heavy, interior of aperture with a pink coloration and the 
surface is covered with the usual, light-brown, peeling perio- 
stracum. The report that this shell was taken from a fish-trap 
seems very questionable. There seems to be notliing closely re- 
lated to S. canaliculatus in the living fauna; S. leidyi Heilprin of 
the Florida Pliocene is perhaps the nearest, but even in this case, 
the resemblance is not close. 

The collection of at least 4 typical specimens of S. canaliculatus 
in a relatively short time and from the same general area seems 
to preclude a chance anomaly or a deformed or diseased shell. 
The absence of axial ribbing on any part of the whorls of the 


128 NAUTILUS Vol. 75 (4) 

spire is again emphasized. More material with accurate data as 
to station and environment is needed and we are calling re- 
newed attention to this interesting species so that collectors 
visiting the Bahamas can be on the watch for it. 



Assistant Curator of Mollusks, 

Academy of Natural Sciences of Philadelphia 

Olsson & Moore (1962) have claimed that Strombus canalicu- 
latus Burry (1949) is a distinct, "neglected" species, not closely 
related to any other living species. They have discounted the idea 
that the 4 known specimens are abnormal. 

5. canaliculatus was originally described on the basis of a sin- 
gle, sub-adult shell, purportedly from a fish trap in 175 fathoms 
off Hope Town, Elbow Cay, Great Abaco, Bahama Islands. The 
trustworthiness of this type locality has bearing on the validity 
of this species. If, as Olsson has suggested (m litt.), it is a deep 
water, "outer shelf" species, it could have been overlooked. 

Unfortunately, Burry gave no information as to how he ob- 
tained the holotype. According to D. R. Moore {in litt.) , Burry 
obtained it from a man named Ford [not Rev. Paul D.], who had 
a shop in Riviera Beach, Florida, and who imported shells from 
the Bahamas. Possibly, Ford bought the shell from a fisherman. 
There are four reasons to doubt that this shell (collected alive) 
came from 175 fathoms off Hope Town: (1) . Although detritus 
feeders can live at great depths, I doubt that such a large, 
herbivorous gastropod (Robertson, 1961) could live at 175 
fathoms, well below the photosynthetic zone.^ (2) . The escarp- 
ment bounding the Little Bahama Bank off Hope Town is 
precipitous. The margin of the Bank is at a depth of 55-70 
fathoms (personal observations) ; soundings of about 1000 
fathoms are recorded as near as 5 miles to the Bank (see H.O. 
chart 0026e) . (3). The currents off Abaco are so strong (2-3 
knots) that retrieval of a fish trap from 175 fathoms would be 

' Various Strombus species, all smaller than 5. canaliculatus, are known 
definitely to live in depths of at least 30-43 fathoms, but not deeper. A tiny 
species has been dredged alive in the Hawaiian Islands between 43 and 66 
fathoms (see Abbott, 1960, pp. 83-84) . 



ligs. 1:1 Siioinhu.s canaliculalus Bmry. Length: 242 



^ i 

^' L.^k 

Spires of iiialloniu'd, tarinalc Mxnnbus (Ti i<(>i iii.s) i^/iic/.s Linn. MalKima 
Islands (?) . Fij^. I. Holotype of S. caintlii iihil us lluiiy. Drawing l)v (lilbcrt L. 
Voss in Burrv (1949). Fig. 2. Pui ported "topoiype" of ,S. raiialirulatus. Same 
slicll as tliat figured l)\ Olsson & Moore (19(32). Both figs, approx. natural size. 

April, 1962 nautilus 129 

impossible without equipment not available to local fishermen, 
Avho do not place their fish traps ("pots") outside the reef. 
(4) . During 3 summers (1953-1955) while collecting shells at 
Hope Town, 1 was unable to get any information from the local 
fishermen about the finding of 5. canaliculatus. These fishermen 
remembered details about shells of commercial value collected 
years before. 

In view of these 4 reasons to doubt that the holotype came 
from 175 fathoms off Hope Town, it is improbable that the 
other 3 specimens of S. canaliculatus (see Olsson & Moore) came 
precisely "from the same source [locality]". Olsson & Moore 
concede that they doubt that the "topotype" figured by them 
was taken from a fish trap. Lyman (1951) reported that the 
adult specimen (presumably the "topotype") "was brought in 
from the Bahamas and delivered to Mr. Burry." Thus, there is 
no compelling evidence that any of these 4 shells came from deep 
water, or even from the vicinity of Hope Town. 

The shell figured by Olsson & Moore, identified by them as 
5. canaliculatus, differs significantly from the holotype. Oriented 
conventionally, the holotype (Plate 12, fig. 1) has a high spiral 
ridge projecting vertically from each whorl of the spire, forming 
a deep channel near the suture. A similar ridge projects laterally 
on the shell figured by Olsson & Moore (Plate 12, fig. 2) ; there 
is no deep channel. In addition, the spires of the 2 shells are of 
different heights. 

In my opinion, the name 5. canaliculatus has been applied to 
malformed shells of S. (Tricornis) gigas Linn. The holotype of 
S. canaliculatus (255 mm. long) and the shell figured by Olsson 
& Moore (242 mm. long) are both the normal size of adult and 
sub-adult S. gigas. A congenital defect of a small part of the 
mantle laying down the shell material of the posterior part of 
the outer lip, or an injury at an early age, would account for all 
the morphological differences (including the suppression of 
spines and axial sculpture) , and for the differences in the form 
of the spiral ridge if the position of the defective part of the 
mantle varies. There are other striking malformations of S. gigas 
(see M. Smith, 1940, and M.C.Z. no. 168409; Salisbury, 1953), 
and similar carinate malformations due to mantle defects occur 
in many other gastropods (Pelseneer, 1920; for more examples. 

130 NAUTILUS Vol. 75(4) 

see Crosse, 1881; Sykes, 1903; Dautzenberg, 1911). 

A malformed shell of 5. (Strombus) pugilis alatus Gmelin 
(Plate 13) corresponds in part to S. canaliculatus. On the last II/2 
whorls a vertical spiral ridge forms a deep channel near the 
suture. Growth was normal until the outer lip was twice broken. 

La Fontaine (1875) and Abbott (1960, p. 112) discuss 2 more 
malformed, carinate Strombus, one (S. vittatus Linn.) canali- 

A simple experiment might prove beyond question that S. 
canaliculatus is a malformed shell. The appropriate portion of 
the mantle of juvenile S. gigas could be artificially damaged in 
order to observe the effects on the subsequently formed outer 
lip of the shell. 

In 1951, 2 years after describing 5. canaliculatus and about 
11/2 years before his death (in Dec, 1952), Leo A. Burry ad- 
mitted to William J. Clench and Ruth D. Turner that he then 
believed he had named a malformed shell. The additional speci- 
mens were in Burry's possession at that time. 

Acknoioledgment: Axel A. Olsson graciously pro\'ided the ex- 
cellent photograph of the spire of S. canaliculatus, and courte- 
ously allowed me to read the manuscript of his paper co-authored 
with Donald R. Moore. 


Abbott, R. T. 1960. Indo-Pacific Mollusca, 1 (2) . 

Burry, L. A. 1949. Shell Notes [publ. Frank Lyman, Lantana, 
Florida], 2:106-109. 

Crosse, H. 1881. J. Conchyl., 2P;341-342, pi. 11, fig. 3. 

Dautzenberg, Ph. 1911. J. Conchyl., 55:209, pi. 10, figs. 12-13. 

La Fontaine, J. de. 1875. Ann. (Mem.) Soc. Malac. Belgique, 
;0:21, pi. 2. figs. 1-2. 

Lyman, F. 1951. Shell Notes, 2:195-196. 

Olsson, A. A. & D. R. Moore, 1962. Naut., 75:127-128. 

Pelseneer, P. 1920. Acad. roy. Belgique; classe sciences; Mem. 
(8vo), (2)5:22, 45. 

Robertson, R. 196L Notulae Naturae (Acad. Nat. Sci. Philadel- 
phia) , no. 343. 

Salisbury, A. E. 1953. Proc. Malac. Soc. London, 30A7AS, pi. 8 

Smith, M, 1940. World-wide sea shells, p. 131. 

Sykes, E. R. 1903. Proc. Malac. Soc. London, 5:260. 

April, 1962 nautilus 131 



Surface sculpturing on the shell of Anomia acule.ata Gmelin 
varies considerably (Verrill and Smith, 187S; Jackson, 1890; 
Whiteaves, 1901; and Foster, 1937) ranging from smooth to 
highly spinose. In fact, lacking intermediates, the variants at the 
extremes could be easily mistaken for separate species. One, ac- 
cordingly, might expect to find grave nomenclatorial difficulties 
in the literature but this is not the case. The smoother of the in- 
dividuals are still occasionally listed with "form" or "varietal" 
names but it is generally well understood that they are part of a 
single species complex. 

With ample material available, this study was undertaken sim- 
ply to describe the degree and complexity of sculptural variation 
normally occurring in this species. As the study progressed I 
foimd that spinosity, if it is expressed, develops at widely differ- 
ing shell sizes. In the usual lamellibranch, a character which 
develops in the postlarval form becomes apparent at about the 
same age in all specimens. But in A. aculeata, this laboratory has 
individuals whose variability for the characteristic spinosity may 
be expressed any time after metamorphosis. 

This paper, then, describes variability in A. aculeata and pre- 
sents data to show the change in ratio of smooth to spiny indi- 
viduals with reference to shell size. 

Methods of Collecting and Measuring. Most of the material 
used in this study came from navigation buoys stationed in off- 
shore waters south of Nantucket, Massachusetts. Since buoys are 
brought in periodically for cleaning and servicing, the many 
forms of life found attached can be removed for study. Anomia 
often homesteads the convenient buoy surfaces, after the initial 
pelagic larval stage, instead of settling in its usual habitat on the 
bottom. Because smaller Anomia were predominant on the buoys 
sampled, the bottom was dredged to obtain representatives of 
the larger adult sizes. 

A. aculeata usually maintains a suborbicular shape during 

^ U. S. Department of the Interior, Fish and Wildlife Service, Bureau of 
Commercial Fisheries Biological Laboratory, Woods Hole, Massachusetts. 

132 NAUTILUS Vol. 75(4) 

growth. Those which, by some accident of position, meet an ob- 
stacle in the immediate vicinity assume various distorted shapes 
in an effort to circumvent the impediments. This apparently does 
not affect the morphology gieatly other than to shift the axis of 
growth. The shape may become transversely elongated, or, more 
commonly, irregularly rounded. Thus, to better indicate true 
shell size, measurements of both the height and length were 
taken, added together, and the result divided by 2. Over 4,000 
specimens, from newly settled larvae to mature adults, were 

Variation in surface structure. A. aculeata derives its specific 
name from the spinous and/or foliaceous character of its surface. 
While most individuals exhibit some degree of surface ornamen- 
tation, many individuals are entirely smooth. Foster (1937), 
reporting on specimens brought up attached to shells from fairly 
deep water on Georges Bank, described considerable variation in 
the spines, but, lacking smooth individuals, he did not describe 
this condition. Verrill (Verrill and Smith, 1873) mentioned vari- 
ations in A. aculeata from the Bay of Fundy and the Casco Bay 
areas and described a variety with the scales "more or less 
absorptive or even entirely absent." Jackson (1890) and Whit- 
eaves (1901) briefly discussed the smooth form. 

Plate 14 illustrates the variation found in the surface sculpture 
of this species. On individuals with well-developed spines, the 
spines may radiate to the ventral margin at regular, well-spaced 
intervals. On others, they may be irregularly or more closely 
spaced. The spines may be seated on raised ridges or may simply 
arise from the surface. 

Concentric lines form at intervals as the shell grows. The 
spines terminate at the line formed at the end of each of these 
growth intervals, and other spines commence to form with 
renewed shell growth. The degree of fluted, frilled, or other 
transverse sculpture depends on the degree of development of 
concentric processes. For example, on those with closely spaced 
and conspicuous growth lines, the spines are blunted. Often those 
with weaker growth lines tend to have spines which project from 
radiating ridges. The spines may also lack radial symmetry and 
simply form along concentric lines apparently at random over the 
shell surface. 

April, 1962 nautilus 133 

On those individuals with smooth surfaces, the concentric 
growth lines are slightly undidate causing the surface to appear 
craggy or scaly. Very few have the smooth and glittering surface 
of the allied species, A. simplex Orbigny; usually the surface 
appears dull and chalky. 

Practically every conceivable combination occurs between 
plain smooth and fully scaly individuals. Those not quite smooth 
may have faint ridges, perhaps just a ray or two in some; others 
may have a full fan of strong ridges with no sign of spines. On 
some with ridges, there may be a raised spine here and there, or 
possibly one of the ridges will be fidly spined with none of the 
other ridges showing the slightest tendency to spininess. Some 
will be smooth to a certain stage in growth, then suddenly 
begin producing nicely developed spines. Others may do this and 
then just as suddenly become smooth again. There is seemingly 
no end to the possible combinations. For the purpose of this 
paper, the individuals have been separated into two recognizable 
categories: those lacking surface ornamentation, and those show- 
ing surface sculpture to various degiees. The latter include 
individuals whose sculpture ranges from faintly rayed to heavily 

Size-frequency data by station: As mentioned earlier, popula- 
tion samples of A. aculeata were taken from 3 offshore buoys as 
well as from the bottom by dredging. Table 1 lists the frequency 
distribution from all stations as well as the percentage of smooth 
Anomia in each size category. The station locations and other 
data relating to them are listed below. 

Nantucket Shoals Lightship Buoy (NSLS). The NSLS buoy, 
located 1 mile, 55° true, north of the permanent Nantucket 
Shoals Lightship at 40° 33' N. latitude and 69° 28' W. longitude, 
was placed on station October 8, 1957. When removed May 10, 
1958, it had been in the water for 7 months. An estimated 
25,000-30,000 specimens of A. aculeata were attached to the buoy. 
The sample used for this study consisted of 2,423 specimens. 

Davis Shoals Buoy (4DS#1). The station occupied by the 
}DS#I buoy is about 25 miles west-northwest of the NSLS buoy 
at 40° 57' N. latitude and 69° 55' W. longitude. This buoy was 
on station 1 year from May 7, 1957, to May 15, 1958. The sample 
taken from this buoy included 537 specimens of A. aculeata. 



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April, 1962 nautilus 135 

Davis Shoals Buoy {4DS#2). The 4DvS#2 buoy succeeded the 
4DS#1 buoy. It remained on station for 6 months from May 15, 
1958, until November 10, 1958. The sample included 983 speci- 
mens of A. aculeata. 

Channel Station. At a station about 44 miles northeast of 
Nantucket Shoals Lightship (41^03' N. latitude; 68°45' W. 
longitude) several bottom tows were made during the course of 
sea scallop investigations on September 16, 1959. From the 
dredged material 246 A. aculeata were picked from shells, stones, 
and debris. 

Discussion: The larvae of A. aculeata pass through a pelagic 
stage in the upper waters during which time they are subject to 
variable currents and winds. Hence, they are apt to settle hither 
or yon in due time on the ocean floor. It is not surprising, then, 
to find samples made up of individuals that vary from one 
another widely. Verrill (Verrill and Smith, 1873) observing 
variation in this species thought it might possibly be a variety 
of the true A. ephippium which has a smooth surface. He was 
sure it was not A. glabra [= A. simplex] of our coast. Whiteaves 
(1901) remarked of the smooth form of eastern Canada as fol- 
lows: "The little smooth form of this region that has been 
hitherto referred to A. ephippium, is regarded by Verrill as most 
probably a form of A. aculeata." Jackson (1890) was able to 
separate the two species of Anomia in our area by means of prc^- 
dissoconchal characteristics. He found that larval A. aculeata 
had a byssal notch both in the left and right valve, while larval 
A. glabra had a notch only in the right valve. By this means, we 
all are sure of which species he was referring when he stated, 
". . . associated specimens are frequently wanting in the spinous 
character, although unquestionably belonging to the same 
species." Bousfield (1958, 1960) followed W^hiteaves' lead (per- 
sonal communication) in listing the smooth form as A. ephip- 

Two authors (Jackson, 1890; Foster, 1937) have remarked on 
the fact that the surface of A. aculeata may be smooth for a 
period of time before spines begin to form. Jackson was referring 
to the young growing for a brief period of time before any spines 
were produced; Foster was commenting on occasional large speci- 
mens on which the spines were represented only on the outer 



Vol. 75(4) 



- ^ 




I i r 







■: \ 
\ \ 
\ \ 

\ \ 


I \ L_J I \ I I I 

6 8 10 12 



Fig. 1. Data from Table 1 plotted to show decrease in percentage of smooth- 
surfaced Anomia aculeata with increase in shell size. 

edge oi the shell. These miscellaneous facts can now be con- 
nected and a clear picture of variation presented by means of 
the data. 

A. aculeata, in truth, is spineless for a brief period. The first 
signs of spines appear on occasional specimens at about 0.7 mm. 
By this time the shell has possibly tripled in height since setting. 
As can be seen on the graph in Figure 1, the percentage fre- 
quency of smooth individuals decreases rapidly at first and levels 
off at about 25 percent above 8 mm. In large spiny individuals 



.\[all()niic'(l. caiinatc Sliuuihus < Sty(niil>u.\j piiolli.s aliiliis (.nielin. jiucnile. 
Shell ct)ated with MgO to show sculptuiL- bin not color pattern. Saiiibel 
Island (?), Florida, .\merican Miiseiiin ol Xaiinal History (no. 46068). Fig. 1. 
\]>ical \iew. x'-^"- I'ig- -■ Vix'rtural \iew. Fig. .S. ()l)li(iiic \iew. Figs. 2-3 
iiainral size. 

NAUTILUS 75 (4) 


A)iomui nnilctilti C.mcliii. Rcpicsciilativc individuals arranged in show 
variation in shell stidpturc. 

April, 1962 nautilus 137 

one can easily trace back along the shell and see when spines 
were first laid down. Although this usually is found to be within 
the first few millimeters of giowth, as reflected by the graph, it 
may be found on particular specimens at any size. Even a speci- 
men of almost maximum size may finally express potential to 
produce spines. 

It should be pointed out, in regard to using material mostly 
from buoys for this study, that the environment on the buoy is 
probably more uniform than on the bottom. However, the small 
amount of data from the bottom collections fits in well with the 
data from buoy samples. This tends to confirm the possibility 
that variation in surface sculpture in A. aculeata might be a 
genetic rather than an environmentally induced phenomenon. 
Conclusion and Summary 

1. Surface sculpture in the shell of Ajiomia aculeata varies 
considerably depending, in part, on size (or age) . The sinface 
may be smooth or, to a highly variable degree, spinose. A long 
series of similarly-sized specimens from single buoy samples can 
be formed in which graded variability in surface structure is 
shown easily. Therefore, in keeping with the usual procedures in 
taxonomy, no distinction shoidd be shown the extreme variants. 

2. All early post-larval A. aculeata have smooth siufaces and, 
although a percentage may remain smooth throughout life, many 
start early to put on spines, the percentage of spiny individuals 
increasing with shell size. 

3. The fact that smoothness is always expressed in the early 
dissoconch shell implies that spinosity is a natural later develop- 
ment. The early smooth shell undoubtedly represents the gen- 
eralized or ancestral form. The generalized form found along the 
western Atlantic coast is A. simplex, or along the European 
coast, A. ephippium (type species of Anomia) . Spinosity, a 
divergence from the generalized, may be considered a derivative. 

Literature cited 
Bousfield, E. L. 1958. Proc. Nova Scotian Inst. Sci. 24 (3) : 303-325. 
1960. Canadian Atlantic sea shells. National Museum of 

Canada, Ottawa, 72 pp. 
Foster, Richard W. 1937. Naut. 50(3) : 102-103. 
Jackson, Robert Tracy. 1890. Phylogeny of the pelecypoda, the 

Aviculidae and their allies. Alem. of the Boston Soc. of Nat. 

Hist. 4 (8) :277-400. 

138 NAUTILUS Vol. 75(4) 

Whiteaves, J. F. 1901. Catalogue of the marine invertebrates of 
Eastern Canada. Geological Survey of Canada, Ottawa, 
271 pp. 

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, 448 pp. 


c/o Transspace Laboratory, Box 111, Princeton Junction, New Jersey 

At the close of a recent period of collecting and experimenting, 
an interesting observation was made pertaining to the fluores- 
cence of shells (1) which is deemed worthy of preliminary 
publication at this time. 

All specimens were obtained at and below (using scuba) the 
"low tide line", Northwest Harbor, Deer Isle, Maine, August 
28-29, 1961. The ultraviolet light used was one originally de- 
veloped for underwater work and littoral exploration (2) emis- 
sion being from a BLB lamp (approximately .8500 to 4000 
angstroms) . 

While attempting to ascertain the suitability of glycerine as a 
preservative for the color and normal fluorescence of "coralline 
algae" (3) , a small "limpet" adhering to one specimen was inad- 
vertently coated with the glycerine, whereupon (in the dark) 
under the ultraviolet light the limpet shell burst into a strong, 
glowing, deep red fluorescence. 

Fifteen living specimens of Acmaen testudinalis testudinalis 
(Miiller) (limpets) were subsequently either coated with or 
immersed in glycerine. In every case a deep red fluorescence 
resulted over the entire shell while under the glycerine. Prior to 
treatment these specimens had no apparent visible red fluores- 

Unfortunately time did not permit an examination of the 
effect of glycerine on the fluorescent areas (feeding?, homing?) 
surrounding such limpets when they are found living on "bare" 
rocks — an observation made by the writer some years ago (un- 
published) . 

Twenty-seven living specimens of Littorina littorea (Linn^) 

April, 1962 nautilus 139 

(the common periwinkle) were treated with glycerine. In all 
cases, a similar deep red shell fluorescence of considerable inten- 
sity resulted, particularly strong in small specimens. Location of 
the area of maximum shell fluorescence was somewhat variable, 
appearing to have a tendency to localize in areas in the proximity 
of the apex. Fifteen specimens kept immersed in glycerine ap- 
pear to have retained their red fluorescence unchanged for two 

In the case of the periwinkles, the writer has felt for some 
years that occasionally some sort of dark red shell fluorescence 
could be observed but it was so illusive and ephemeral that such 
observations could be considered at best as highly questionable; 
however, it has led the writer to refer to the phenomena as 
"enhancement" of fluorescence, rather than, say, "inducement." 

Vigorous scraping with a knife blade of both limpet and peri- 
winkle shells subjected to the above glycerine treatment re- 
sulted in no apparent diminution of shell fluorescence until the 
shells were deeply abraded — suggesting that the phenomena is 
not due to a superficial shell coating. 

Three small, living specimens of Bucciniim undatum Linn^ 
(the common welk) coated or immersed in glycerine showed no 
such shell fluorescence. 

A small specimen of Clanculus puniceus from the writer's col- 
lection of fluorescent shells showed no particular enhancement 
of its normal brilliant rose fluorescence. 

Woodbridge, R. G. Fluorescent Shells, a Monograph, etc. Pub- 
lished by Transspace Laboratory, 1961. 
Ibid. Underwater Ultraviolet Lights, etc. Published by Trans- 
space Laboratory, 1960. 
Woodbridge, R. G. and Woodbridge, R. C., The Application of 
Ultra-Violet Lights to Underwater Research, Nature 184:2b^ 
(July 25, 1959) . 



Scripps Institution of Oceanography 

La Jolla, California 

Olivella mutica (Say) occurs commonly in warm, shallow 

^Contribution no. 171 from the Oceanographic Institute, Florida State 
University, Tallahassee, Florida. 

140 NAUTILUS Vol. 75(4) 

waters from North Carolina to Texas, and in the West Indies 
(Abbott, 1954) . However, little appears to be known about its 
general biology. The following observations are primarily con- 
cerned with the external form of the egg capsvde, length of the 
developmental period, and duration of the spawning season. 
Some data are also presented on the nature of the occupied 
habitat and population density. This study was made possible 
through use of the facilities of the Alligator Harbor Marine 
Laboratory of Florida State University, in Franklin County, 
Florida. Menzel (1956) has cited O. mutica as a common snail 
in the immediate region. Olivellas were collected from the south 
end of a sandbar lying across the mouth of Alligator Harbor. 
Extensive portions of this bar become exposed at most low tides. 

Thirty- two snail egg capsules of similar appearance but of 
unknown parentage were collected, primarily from the valves of 
a brachiopod which occurs commonly on the bar. All but 7 
of these embryos were reared successfully in filtered sea water, 
and many of them were then maintained until they were suf- 
ficiently large to be identified. Most were identified from Abbott 
(1954) and Olsson's monograph (1956) as O. mutica. However, 
one O. pusilla (Marrat) which is also present at Alligator Har- 
bor, was subsequently discovered included with the specimens of 
O. mutica. Because, unfortunately, laboratory-raised and field- 
collected specimens were mixed, I do not know whether all the 
eggs were O. mutica, or whether a few might have been O. pu- 
silla. This confusion of two very similar species will not affect 
those conclusions based on egg capsules in which O. mutica is 
known to have predominated. However, no distinction was 
drawn between these two olivellas in the field and thus the 
density and habitat information includes both species. 

Adult snails were noted from September, 1959, to August, 
I960, indicating year-round occupancy of the intertidal portions 
of the sandbar's southern end. The bottom sediment here is 
predominately a mixture of fine-grained quartz sand M'ith an 
average diameter of 0.125 to 0.250 mm., and a few shells or shell 
fragments. The silt and clay fractions of the sediment are mini- 
mal, and little decaying organic matter has accumulated. The 
marine grass Diplanthera wrightii (Ascherson) , occurring in 
patches, has consolidated portions of the bottom sediment but 
has not changed the physical characteristics of the substratum 

April, 1962 nautilus 141 

appreciably. Olivellas were collected from both consolidated and 
unconsolidated sandy sediment, with the maximum density of 
178 snails per square meter in the latter. Snails of this genus 
were not found in samples from soft bottom areas overlain by 
quantities of organic material. 




Fig. 1. Usual egg capsule ot Olivella mutica. 

The encapsulate eggs of O. mutica (Figure 1) are laid separ- 
ately, one ovum per capsule, with no trace of nurse eggs, and are 
attached to any solid object that protrudes above the surface of 
the sand. The geneial cliaracteristics of the egg and the site of 
oviposition are thus similar to those described by Marcus and 
Marcus (1959) for O. verrcauxii (Ducros) . The base of the egg 
capsule of O. mutica is roughly circular and 1.0 to 1.1 mm. in 
diameter. From this base rises a smooth, spherical, transparent 
dome about 670 microns in diameter. Near the top a distinct 
ridge indicates the line along which the capsule lid separates 
when the young snail escapes. In general form the capsules ap- 
pear indistinguishable fiom those figured by Perry and Schwen- 
gel (1955) for O. pusilla. In contrast, the egg capsule of O. ver- 
reauxii (Marcus and Marcus, 1959), though entirely similar in 
gross shape, can readily be distinguished by the surface sculptur- 
ings — fine, raised, parallel lines. 

The majority of the eggs of O. mutica were found attached to 
the anterior end of the valves of the inarticulate brachiopod, 
Glottidia pyramidata (Stimpson) , with from 1 to 5 capsules per 
brachiopod. Freshly-deposited eggs or developing embryos were 
obtained on 25 March and 8 April, 1960, when the water temper- 
ature ranged from 16° to 20° C. The maximum developmental 

142 NAUTILUS Vol. 75(4) 

period, based on laboratory-reared eggs, was 19 or 20 days for 
both collections. The cleavage and veliger stages are spent in the 
capsule, and upon hatching the capsule lid is broken off and a 
young snail, 750 microns in length crawls away. On the other 
hand, Marcus and Marcus (1959) have shown that O. verreauxii 
hatches as a veliger after 8-9 days. 

Large samples of the brachiopod were also closely scrutinized 
on 26 February, 10 and 12 March, 22 and 23 April, and 3, 8, 25, 
26, and 28 May; they were devoid of Olivella egg cases. The first 
egg to hatch in the laboratory did so on 6 April. If one extrapo- 
lates back 20 days, the suggested period of embryonic develop- 
ment, the initial spawning took place about 17 March. Egg- 
laying appears to have ceased sometime between 8 and 22 April. 
These data suggest that O. miitica spawns for approximately one 
month during the spring, and that egg-laying is closely limited 
to this period. No information is available on spawning in the 
fall, or latitudinal variations in the duration of egg-laying. 

Literature cited 
Abbott, R. T. 1954. American Seashells. Princeton: D. vanNos- 

trand Co. 541 pp. 
Marcus, E. and E. Marcus, 1959. On the reproduction of Olivella. 

Univ, Sao Paulo, Fac. Filos. Cienc. e Letras, Bol. Zool. 22: 

Menzel, R. W. 1956. Annotated check-list of the marine fauna 

and flora of the St. George's Sound - Apalachee Bay Region, 

Florida Gulf Coast. Fla. State Univ. Ocean. Inst., Contrib. 

Olsson, A. A. 1956. Studies on the genus Olivella. Proc. Acad. 

Nat. Sci., Philad. 108:155-225. 
Perry, L. M. and J. S. Schwengel, 1955. Marine shells of the 

western coast of Florida. Ithaca: Paleon. Res. Inst. 318 pp. 



The symbols used for Puerto Rican localities were explained 
recently, 1961 (1) . 

Yunquea denselirata H. B. Baker. Only known from type 
locality (Er3) ; on last whorl of type (only dry) shell, "very fine" 
growtli threads average 1 1 microns from crest to crest, which 
means 91 per mm., and that the "very slightly stronger" ones run 
13 to 15 per mm. Y. monteplatonis (Pilsbry) has similar threads. 

Odontosagda sp. "Yunquea denselirata" van der Schalie, 1948: 

April, 1962 nautilus 143 

71, pi. 6, fig. 5. Probably an unnamed species, from about 3 km. 
south of Aguas Biienas. 

The anatomy of Odontosagda is unknown. Although its shell 
has internal lamellae (like some Sagdinae) , in general form and 
size, it does resemble more closely that in Yunquea (Yunqueinae). 

Hyalosagda (Microsagda) subaquila (Shuttleworth) . Among 
dead leaves in disturbed places. En, Jn, Pn, Prl (2500 ft.) , Wn, 
Wr2, Ws, Ww2; mainly lowlands. Foot light near sole but almost 
black above, with lighter middorsal stripe and triangle between 
ommatophores, which are very dark; eyes light with dark centers. 

H. (M.) subaquila, var. a. Psl-3; shell more depressed, with 
larger umbilicus (about I/5 major diameter) . 

H. (Lacteoluna) selenina (Gould) . Probably distributed by 
commerce; Es2, Jn, Pn, Pr2-6, Wn, Wr, Ws, 0-3400 ft., but not 
found in Luquillo forests. 

H. (Aerotroc/ius?) krugiana (Martens) . Size (1877:346) : 3.5 by 
171 (6 mm.) with 5 whorls, subangulate, narrowly umbilicate. 
Caguana, west of Utuado. 

Martens compared the last species to the Cuban Helix turbini- 
formis Pfeifier, which at the time was confused with type species 
of Aerotrochiis, the Jamaican Hyalosagda subpyramidalis (C. B. 
Adams) ; the last is a higher, but angulate shell, which lives on 
trunks of trees. These 3 Puerto Rican species belong in the sub- 
genus (or genus?) Lacteoluna; Hyalosagda (s. s.) and Stauro- 
glypta appear to be mainly Jamaican. 

Aquebana (s. s.) velutina (Lamarck) . Living animals very rare, 
under rocks, 5-6 inches below surface (Jnl) and under dead 
leaves between limestone ledges (Enl) during and at end of 
drouth, but dead shells numerous (Jn2) which may indicate 
that it has periods of abundance. Fresh shells with greenish tinge. 
To 1940c: 59, is added: foot pale, with chrome spots, obscured 
by chalky ones. 

Platysuccinea portoricensis (Shuttleworth) . Terrestrial, under 
upper layer of dead leaves, especially in ruderal places; Er, Pr, 
Wr, 1500-4000 ft. Fresh shells, with adherent dirt except in 
columellar region, greenish corneous on El Yunque and often 
with very low spires, but often more hyaline with higher spires 
in Cordillera Central. Animal (added to 1940c:61) rather slug- 
gish; side of foot very warty, ochraccous to brown, darker in 
wrinkles, dorsad and on ommatophores; mantle collar not over- 
lapping peristome but with umbilical lobe over columellar area; 
sole dark slate, only trizonal, with numerous rapid waves in 
middle zone. 

P. portoricensis, var. a. Under rocks, Enl; also empty shell 
from Pnl ; less than 200 ft. Shells much smaller, more hyaline and 

144 NAUTILUS Vol. 75 (4) 

with higher spires, but not distinguishable from young ones from 
Cordillera Central; also quite similar in form to "Simpulopsis" 
aenea Pfeiffer, 1861, as figured by Fischer & Crosse, 1877, Miss.. 
Mex. Moll. 7;pl. 24, figs. 12; "S." cumingi Pfr., P. Z. S., 1861:pl. 3, 
fig. 2, from Mexico, looks like typical P. portoricensis (some El 
Yunque shells even larger) ; and ruderal snails do get around. 

Varicella (Vagavarix) portoricensis (Pfeiffer) . Terrestrial; En, 
Es, Jn, Pn, Pr3,6, Wn, Wr2,3, Ws, 0-3000 ft., but not found in 
Luquillo Mts. 

V. (Vagavarix) calderoni H. B. Baker. On ground; Pr6, Wr2, 
2000-3000 ft., Cordillera Central. 

V. (Vagavarix) sulculosa (Shuttleworth) . Terrestrial; En, Es, 
Jn, Pn, Pr 1,5,6, Ps2,3, Wn, Wr, Ww, 0-4000 ft., but not found in 
Luquillo Mts. Larger lots intergrade completely with smoother 
form terebraeformis (Sh.) , which has more widely and irregu- 
larly spaced growth striae; mostly size of sulculosa but one (Ps2) 
12.1 mm. long with 8.2 whorls. 

V. (Vagavarix) sporadica H. B. Baker. On ground; Es3,4, Er2, 
200-2500 ft., only found in eastern Puerto Rico. 

Oleacina (Laevaricella) interrupta (Shuttleworth) . Usually 
subarboreal, but also seen alive under leaves at base of trees; 
Er2, Pr2,3, Wr2,3, 2000-3400 ft., ranging from Luquillo Mts. to 
western Cordillera Central. Animal (1941b:29) seen attacking 
Nenia; lateral zones of foot narrow. 

Oleacina (Boriquena) glabra (Pfeiffer, 1846) . Terrestrial, Er2- 
5, above 2000 ft. in Luquillo Mts. 

O. (B.) playa (H. B. Baker) . On ground; En, Es3, Jn, Pn, Wn, 
Ws, lowlands, 0-800 ft. This "large, elongate, smooth species com- 
mon to Puerto Rico" first noticed and called "gracilior" by Shut- 
tleworth, 1854:51; all I did (1940a) was to prove it a distinct 
species and name his "var. beta" in Varicella; if his descriptive 
term be considered a trinomial, it would be preoccupied in 
neither Gla7idina nor Oleacina. 

Although the animal of Oleacina s. s. is still unknown, Bori- 
quena (possibly only a section) and Laevaricella (a very distinct 
subgenus) now are made congeneric with it, because of the 
close resemblance in the embryonic shells of Oleacina, Boriquena 
and Varicella s.s., which was pointed out by Pilsbry, 1907 (12a): 
127, and because the differences in shell form between O. voluta 
(Gmelin) and O. (voluta var.?) flexuosa (Pfr.) from Haiti so 
closely parallel those between the Puerto Rican O. glabra and 
O. playa. In 1956:133, 135, after vacillating in I941d and 1943c, 
the almost strictly Antillean Oleacinidae (-f Varicellarum) , with 
1 Cuban species in Florida, were separated from the Spiraxidae 
(including Streptosylinae and Euglandininae) , which are most 

April, 1962 nautilus 145 

differentiated on the mainland, but have invaded Jamaica (Spir- 
axinae) and other Antilles [Streptostylinae: Salasiella (Laevolen- 
cina & Flavoleacina), Streptostyla (Rectoleacina), etc.]. 

These two families have almost nothing in common except 
their carnivorous habits and the correlated convergences in the 
radulae of their most specialized groups. Even in their radulae, 
the Spiraxidae almost intergrade with the Achatinidae (sens. 
lat.) , which also have little epiphallic differentiation and some- 
times (Ferussaciinae) have similar "heterurethrous' " pallial com- 
plexes. The changes in the radulae within Varicella (especially 
Melaniella) show that the convergences (uniformly aculeate 
teeth) have come from different sources, and the Oleacinidae 
have sigmurethrous pallial complexes and truly penial epiphalli, 
often with well developed verges, much more like those in the 
Sagdidae. The unroofing of the adrectal limb of the ureter, which 
is paralleled in both the Oleacinidae and the bigger Spiraxidae 
(and in many other groups of geophiles) is found also in 
Yunquea, which has the most generalized radula in the oleaci- 
noids. Incidentally as far as known, outside the small species of 
Hojeda, and the riideral Hyalosagda selenina and (possibly) 
Platysuccinea, the Sagdidae also seem to be restricted to the 

Additional references (see 1962) 
1961 (1), Naut. /•/ .-142-149. 1962, Naut. 75:116-122. 


Department of Biology, Kansas State Teachers College, Emporia 

The subfamily Neoplanorbinae was established by Hannibal 
(1912) to accommodate 5 species in two unusual genera of fresh- 
water mollusks. These species demonstrate an extreme in endem- 
ism, all known specimens having been collected from the lower 
Coosa River bordering Coosa, Chilton, and Elmore counties in 
central Alabama. As far as I am aware, only t^vo men, H. H. 
Smith and A. A. Hinkley, have ever seen these animals alive, and 
it is unlikely that living specimens will be seen by anyone else. 

The first mention of these snails was by Hinkley in 1904, who 
listed both "Planorbis tnntillus Pilsbry" and an undescribed 

' Supported by grant G-14125 from the National Science Foundation. 

146 NAUTILUS Vol. 75 (4) 

form, "N. g., n. sp." among a group of Alabama shells collected 
the previous year. Pilsbry (1906) recognized these shells as 
". . . quite unlike any fresh-water snails hitherto known in this 
country, having affinities with the Ancylidae." He established 
two new genera for them, describing Amphigyra alabamensis and 
Neoplanorbis tantillus as new species. Two years later. Walker 
(1908) named 3 additional species of Neoplanorbis: N. cari- 
natus, N. umbilicatus, and N. smithii, and these have remained, 
without change or addition, to the present. 

In 1914 Lay Dam, the first of a series of major hydroelectric 
power installations, was completed across the Coosa River be- 
tween Coosa and Chilton counties, 13 miles east of Clanton, Ala- 
bama. This dam imponds an area of 6,000 acres, with a back- 
water extending to a point 13 miles below Childersburg (Pierce, 
1955) . Sixteen miles downstream Mitchell Dam was completed 
in 1923, with 5,800 acres of backwater extending to Lay Dam. 
In 1929, Jordan Dam was built 8 miles north of Wetumpka, 
Elmore County. This dam supports a lake of 4,900 acres, extend- 
ing about 18 miles upstream to Mitchell Dam. The large lakes 
formed by these dams, which provided power and recreational 
facilities for the people of central Alabama, also destroyed the 
habitats (and presumably caused the extinction) of all known 
species of Neoplanorbinae. These tiny mollusks lived on stones in 
the swift current of the Coosa (See Goodrich, 1944) , in localities 
now covered by many feet of impounded lake water. An attempt 
in June, 1959, by the author and Dr. John B. Burch to find living 
specimens in the Coosa was not successful (Basch, 1959) . 

Through the kindness of Dr. Henry van der Schalie of the Uni- 
versity of Michigan Museum of Zoology I have been given access 
to the specimens of Neoplanorbinae in his care, with permission 
to extract and prepare radulae from a sample of the specimens. 
The radulae were extracted and mounted using techniques out- 
lined in a previous paper (Basch, 1961) . A study of these radulae 
was undertaken in an attempt to learn more about the systematic 
position of the Neoplanorbinae by comparison of their radulae 
with other groups of Bosommatophora. 

In Neoplanorbis, the shell is planorboid and extremely sim- 
ilar in conformation and dimensions with shells of Micro- 
menetus of the family Planorbidae. Since Menetus (Micro- 

April, 1962 nautilus 147 

menetus) alabamensis is known from the state, 1 considered that 
possibly Neoplanorbis represented a special group of Menetus- 
like planorbid snails. Ecologically, Menetus is an inhabitant of 
quiet, plant-filled pools in a situation vastly different from the 
rocky rapids of the old Coosa River. Shells and radulae of 
Micromenetus are illustrated by F. C. Baker (1945). 

The following radulae have been prepared (UMMZ numbers): 

#102672 — A^ tantillus, 2 specimens. 
#102664 — N. carinatus, 2 specimens. 
#102671 — ^A^. smithii, 2 specimens. 

The species studied were all similar in radular characters. Neo- 
planorbis tantillus, carinatus, and umbilicatus are obviously 
closely related on the basis of shell similarities and may in fact 
represent a single variable species. The radulae of A^. tantillus 
and N. carinatus are indistinguishable (it was not possible to 
obtain a suitable specimen of A'^. umbilicatus). Neoplanorbis 
smithii, whose shell is easily differentiated from the other three 
by its lack of spiral sculpture, has a radula differing in minor 
characters, particularly the angle formed by the medial surface 
of the lateral teeth with the longitudinal axis of the ribbon. In 
all three species of Neoplanorbis the radular formula is 17-1-17, 
with the inner cusp of the laterals long and dagger-like, bearing 
a small accessory cusp on its medial surface. The marginal teeth 
are characterized by an increasingly rectilinear upper edge and a 
prominent needle-like cusp located just medial to the center of 
the tooth (Page 148, Figs. 1 and 2) . The extreme smallness of 
these teeth must be pointed out, for some of the finer features lie 
beyond the resolving power of a light microscope. The optical 
equipment used in this study consisted of a Leitz triocular 
Labolux microscope with a lOOX N. A. 1.30 oil immersion ob- 
jective and paired 20X oculars. An internal prism factor of 1.25X 
provided a final magnification of 2500X. Even at this extreme 
magnification, some details were not readily distinguishable in 
the bacteria-sized teeth. 

The radula of Amphigyra is strongly bicuspid (cf. Pilsbry, 
1906) and similar to that of Neoplanorbis. The formula is 
18-1-18, and lateral teeth 7 to 11 resemble planorbid and some 
ancylid laterals rather closely (Page 148, Fig. 3), perhaps estab- 
lishing a point of homology. Two radulae were prepared from 



Radulae of Neoplanorbinae. Fig. 1, N. carinatus Walker; Fig. 2, A^. smithii 
Walker; Fig. 3, Amphigyra alabamensis Pilsbry. 

specimens in UMMZ lot #\Q2Q11. 

Judging by the radula, apparently Amphigyra and Neoplan- 
orbis are closely related to each other, but they have diverged 
considerably from other groups of Basommatophora. With 

April, 1962 nautilus 149 

a dearth oi anatomical data concerning the Neoplanorbinae, its 

position within the Ancylidac is not firmly supported by radular 

structure alone, but no other evident relationships are revealed. 
Literature cited 

Baker, Frank C. 1945. The Molluscan Family Planorbidae. 
Urbana. Univ. of Illinois Press. 530 pp. 

Basch, Paul F. 1959. The Coosa Revisited. Amer. Malac. Union 
Annual Rept. for 1959: 17. 

1961. Radulae of North American Ancylid Snails. I Sub- 
family Rhodacmeinae. Naut. 75:97-101. 

Goodrich, Calvin. 1944. Naut. 5.9 (1) :11-15. 

Hanibal, Harold. 1912. Proc. Maloc. Soc. London. 70:1 12-211. 

Hinkley, A. A. 1904. Naut 18(5) :54-57. 

Pierce, Laurence B. 1955. Hydrology and surface-water resources 
of east-central Alabama. Geol. Surv. of Alabama, Special 
Rept. 22:1-318. 

Pilsbry, Henry A. 1906. Naut. 20 {5):49-5\. 

Walker, Bryant. 1908. Naut. 2/ (1 1): 126-129. 


National Science Foundation, Washington, D. C. 

The hooked or curved mtissel, Brachidontes reciirvus (Rafin- 
esque) of the family Mytilidae, occurs in large numbers on oyster 
bars in the tipper Chesapeake Bay and its tributaries. This species 
is an important fouling organism on oyster bars and affects the 
growth habits of the oyster. It was reported by Frey (1946) as 
forming much of the cultch on oyster bars in the Potomac River 
where most of the marketable oysters weie completely covered 
with mussels. As a result, the oysters were misshapen from grow- 
ing among the mussels. Beaven (1947) states that this species is 
most abundant on bars of the upper portion of the Bay and its 
tributaries where the salinity is low. 

Under the name Mytilus recurvus, this species has been re- 
ported (Nelson, 1928a) as occurring along the New Jersey coast. 
Chestnut (1949) noted the appearance of the curved mussel in 
North Carolina. Englc (1945) observed that the hooked mussel, 
Mytilus Jiaynatiis Say, was present on oyster bars in Alabama, 

* Work done while with the Department of Zoology, University of Mary- 
land, College Park, Maryland. 

150 NAUTILUS Vol. 75(4) 

and later (1948) the same investigator reported its presence on 
bars in Mississippi and Louisiana. According to Abbott (1954) 
the geographic range of Brachidontes recurvus is Cape Cod to the 
West Indies. 

Despite the wide distribution of this species and its impor- 
tance as a fouling organism on oyster bars, a survey of the lit- 
erature reveals that with the exception of the observations on 
distribution previously mentioned, little information is available 
concerning its biology. This situation is in sharp contrast to that 
existing in regard to other mussels, such as Mytilus edulis Linne; 
Mytilus calif ornianus Conrad; Modiolus demissus (Dillwyn) ; and 
Modiolus modiolus (Linne) , where considerable data are avail- 
able on numerous aspects of the biology of the species. 

The investigation reported here was part of a cooperative pro- 
gram between the Department of Zoology of the University of 
Maryland and the Shellfisheries Investigations of the United 
States Fish and Wildlife Service, located at Annapolis, Maryland. 
Appreciation is expressed to Dr. Robert A. Littleford for his 
suggestions and criticisms; and to Mr. James B. Engle and Staff 
of the Fish and Wildlife Service for collection of materials and 
for data on temperature and salinity. 

Procedure: Specimens of B. recurvus used in this study were 
collected from Hackett's Bar, located on the western side of upper 
Chesapeake Bay and approximately two nautical miles north of 
the Severn River. This bar is 1/2 to % of a mile offshore from 
Hackett Point, in water varying in depth from 2 to 35 feet. Sam- 
ples were taken at two week intervals from January 16, 1950' 
through April 16, 1951, except when prevented by weather con- 
ditions or mechanical difficulties with the boat. All the mussels 
were collected with a standard oyster dredge from the same gen- 
eral location on the bar and at a depth of 16 feet. 

With the shell closed and the specimen held in a lateral posi- 
tion (Newcombe and Kessler 1936), shell dimensions were 
measured to the nearest tenth, using a sliding vernier caliper 
calibrated in millimeters. The dimensions used are defined as 
follows: length, the greatest distance between the anterior and 
posterior ends of the shell; width, the greatest distance between 
the dorsal and ventral edges; and thickness, the greatest distance 
between the right and left valves. The term height is commonly 


April, 1962 nautilus 151 

used in place of thickness as a more meaningful designation. 

Specimens of the entire size range were selected from each 
collection for microscopic study. The individuals were fixed 
within the shell by severing the posterior muscles and then drop- 
ping the entire animal into Bouin's alcoholic fixative (Galigher, 
1934) . Other fixatives were tried but Bouin's proved most satis- 
factory. After fixation for several days, they were removed from 
the shell, washed in 70% alcohol and then stored in 80% alcohol. 
They were dehydrated in alcohol and xylol, embedded in Tissue 
Mat, sectioned at 10 micra, stained with either Harris's hema- 
toxylin and eosin or Mallory's triple stain, and mounted in 

Plankton samples were collected simultaneously with the col- 
lection of mussels by pumping 100 liters of water from just above 
the bar through a No. 20 silk plankton net. Each sample was 
concentrated to 100 ml. and preserved with formalin. Examina- 
tion of these samples to determine the presence or absence of 
mussel larvae was made with a Sedgewick-Rafter Cell (Little- 
ford, Newcombe, and Shepherd, 1940) . 

Obseiuations. The gross microscopic morphology of the curved 
mussel is identical with that of Mytihis edulis as described by 
Field (1922) . However, the alimentary tract is sunounded by 
the visceral mass which contains the other organ systems of the 
animal. Immediately surrounding the stomach and intestine is 
the so-called liver, or digestive diverticula. The latter structure 
also extends into the mesosoma, which is an elongate, ventral 
extension of the visceral mass, except during the period of gonad 
development, when the mesosoma is filled with the gonad tissue. 

The gonadal tissue of B. recurvus consists of a series of ducts 
and canals, lined with germinal epithelium, which end in pockets 
or follicles. The mature tissue occupies almost the entire mantle 
(Plate 15, fig. 4) and the mesosoma to the edge of the foot and 
penetrates into the digestive diverticula, where it is found ad- 
jacent to the so-called liver canals, the stomach, and the direct 
and recurrent intestine. Apparently the early gonad development 
begins in the mantle and then spreads into the mesosoma. The 
vesicular tissue which fills in between the follicles is replaced by 
the gonad tissue as it matures. Thus, when the sex cells are 
mature, nearly the entire structure of the organism is devoted to 

152 NAUTILUS Vol. 75(4) 

reproduction. As sexual maturity approaches, the mantle and 
mesosoma become either a bright yellow or a stippled brown 
in color. 

The craved mussel is dioecious, and of the 896 specimens exam- 
ined, there was no indication of the existence of hermaphrodit- 
ism or alternation of sex. The gonads of both male and female 
show parallel development in time and in the position of the 
gonads. From those studied, the sex ratio apparently is evenly 
distributed. The reproductive potential of these mussels is de- 
termined by their size rather than by their age. Studies were made 
of those forms which were as small as 20 mm. in length, and their 
gonad tissue had the same appearance as that of the larger 
specimens. Presiuiiably they produce as many reproducti\c cells 
as their anatomical structure will allow. This also leads to the 
conclusion that sexual maturity occurs the summer following 
setting, regardless of size of the mussel. 

Seasonal Developmerit in the Male. The male follicles vary a 
great deal in size, depending upon whether the immature or 
mature condition is observed. From January until mid-April, the 
follicles are found in tlie mantle, and to a lesser decree, in the 
mesosoma. They appear at this time to be loosely filled with cells, 
but by mid-April they become more dense in appearance and 
they take a deeper and a darker stain (Plate 15, fig. 1) . As they 
mature, the follicles become filled with spermatozoa which ra- 
diate from the periphery of the follicle toward the center (Plate 
15, fig 2). These cells appear as small oval or spherical bodies 
which are packed too closely together for their detailed structure 
to be clearly visible. They matiue by June and remain mature 
until October. Observation on the density of the sex products 
in the follicles leads one to believe that spawning may occur 
from the mantle first, then proceed from the mesosoma. However, 
the appearance indicates that the animal could very well spawn 
simiUtaneously from both areas. Spawning from each follicle ex- 
tends over a considerable period of time so that some follicles 
do not have the appearance of being spent, while others are 
mature. As the follicles mature, there are light, radiating areas, 
which under increased magnification show that the cells adjacent 
to these areas are concentrated in short rods, apparently similar 
to the condition observed in Mytilus edulis (Field, 1922). 

April, 1962 nautilus 153 

While the follicle increase in size and maturity, the supportive 
vesicular tissue surrounding them gradually disappears almost 
completely. After spawning, the supportive tissue again appears, 
filling the spaces between the spent follicles. 

Seasonal Development in the Female. The immature follicles 
of the female are separated by vesicular connective tissue and 
vary in size. By the first week in May, the follicles are lined with 
small cells containing nuclei which with hematoxylin and eosin 
stain a dark blue. By the third week in May, the cells have in- 
creased in size so that the prominent nucleoli can be seen, but the 
cells are still immature. By mid-June, the mantle and the 
mesosoma are occupied by follicles which are filled with mature 
and immature eggs (Plate 15, fig. 3) . Some of the vesicular sup- 
portive tissue is present, but has decreased considerably or dis- 
appeared as the eggs approach maturity. The eggs vary a great 
deal in size and the nucleoli are large and prominent (Plate 15, 
fig. 3) . When the eggs mature, they break from the follicle and 
are inoved along by ciliated ducts. The eggs are not considered 
mature until they break free. As is the case in the male, the 
female gonad tissue penetrates into the digestive diverticula and 
is in close proximity to the liver canals. (Plate 15, fig. 3) . The 
condition of the gonad from July to October is similar to that in 
Jime. After October, however, the ova are not as numerous and 
some retain the same appearance as those of June. 

On July 3, the small eggs appeared in the mantle and the 
larger ones in the mesosoma which contained many follicles. In 
one specimen from the September group, one lobe of the mantle 
contained only a few eggs, while the other lobe and the mesosoma 
were dense with them. Although the condition of the maturity 
of the ova is such that the entire gonad tissue is subject to spawn- 
ing at the same time, the intensity of spawning and the time of 
spawning of these different areas depends upon the individual. 

Mature eggs are present in December, but show evidence of 
degeneration or death, as shown by the condition of the nucleus, 
the absence of nucleoli, and the general appearance of the cyto- 
pla.-,m. WHiile the destruction of the gonadal tissue appears evi- 
dent, there is no similarity to the comparable mechanism of re- 
sorption noted in some pclccypods by Loosanoff and Davis 

154 NAUTILUS Vol. 75(4) 

The development ot tlie gonads indicates that the state of sex- 
ual maturity exists from June through October. Release of the 
sex products occurs over a relatively long period of time, from 
early June through October, and in some cases, through Novem- 
ber. Individual follicles develop independently of each other in 
both sexes. In other words, the gonads do not reach a high peak 
of maturity and immediately spawn out. 

Seasonal Distribution of the Larvae. Plankton samples col- 
lected from January 16, 1950, through April 16, 1951, were ex- 
amined to determine the presence, relative abundance, and stages 
of development of the larval forms of B. recurvus. 

Since the rate of development of the larval stages shows con- 
siderable variation, in part, at least, because of the protracted 
spawning period, I decided to divide the forms present into three 
groups: pre-hinge stage, hinge stage, and post-hinge stage. The 
hinge stage was defined as "the straight-hinge-line embryonic 
shell" (Field 1922), between the circular immature larval stage 
and the triangular ovate form of the mature larva. In the area 
of Hackett's Bar, the larval stages of this species could be con- 
fused only with the larval stages of Congeria leucophaeta, Cras- 
ostrea virginica, or possibly with Tagelus plebeius and Mya 
arenarin. However, as is pointed out by Sullivan (1948) , care- 
ful examination will show numerous diagnostic differences be- 
tween the mussel larvae and other lamellibranchs. Therefore, the 
larval stages of different groups may be easily separated from 
each other, although some confusion might possibly arise in 
regard to the larvae of Brachidontes recurvus and that of Con- 
geria leucophaeta. However, considering the rare occurrence of 
the latter form on Hackett's Bar, such confusion should not 
invalidate the data. 

The results of the observations on larval distribution are pre- 
sented in Table I. The relative abundance of the various larvae 
is expressed as present, common abundant, and very abundant, 
rather than by using some numerical statement. I felt that such 
an expression of results had more validity than an exact numer- 
ical expression, since the method of collection was not subject to 
exact quantitative analysis (Littleford, Newcombe, and Shep- 
herd, 1940). 

Larvae appeared in the plankton for the first time on June 5, 

April, 1962 










June 5 




June 19 




July 3 




July 17 




July 31 




August 14 




August 28 




Sept. 11 

Very Abundant 

Very Ab'jndant 

Very Abundant 

Sept. 25 




October 9 




October 25 




Nov. 13 




Nov. 27 




Dec. 11 




1950 when pre-hinge and hinge forms were observed in the 
samples. By the middle of June, the pre-hinge and hinge stages 
had quadrupled in number over the earlier part of the month. 
The post-hinge stage was observed for the first time on June 19, 
1950. During July and August there was, generally speaking, a 
continued rise in abundance of the larval stages. However, the 
pre-hinge stage was not found on July 17, 1950 and the post- 
hinge stage was not observed in collections of July 31 and August 
14. All three stages were present in large numbers in the collec- 
tion of August 28, and the peak of their numerical abundance in 

156 NAUTILUS Vol. 75(4) 

the plankton was ob,ser\cd on September 11, 1950. 

Some idea of the relative seasonal abundance of the larval 
stages may he obtained from the fact that the number of pre- 
hinge larvae in June was 27.50% of the total number observed on 
September 11. Similarly, the number of the hinge larvae in June 
was 15.15% and the number of post-hinge was 0.54% of the 
total observed on September 1 1 . 

(To be concluded) 


Institute Nacional de Endemias Rurais, Brazil 

This species was described by Pfeiffer (1839, p. 354), on the 
basis of the shell characters, as follows: 

"43. Planorbis albicans Pfr. — Testa orbiculari, utrinque um- 
bilicata, solidula, albicante vel pallide fulvicante, anfract. 3 
teretibus; labro subincrassato albo; apertura subovata. — Diam. 
21/2, alt. 1'". — Dem PI. albus (hispidus) am nachsten verwandt." 

Clessin's desauption (1884, p. 119-120) is more detailed and 
extends the species range to other Antillean islands: 

"87. Planorbis albicans Pfeiffer. Taf. 11. Fig. 14. T. depressa, 
utrinque centro profundissime immerso; subtiliter striatula; 
lutida, pallide-cornea; anfractus 4, rapide accrescentes, rotundati; 
superius valde convexi, sutura profundissima separati; inferius 
planulati, sutura paulo immersa disjuncti; idtimus penultimo 
paulo latior, aperturam versus descendens; apertura late-lunata, 
albo-labiata, valde obliqua; peristoma acutum, marginibus con- 
junctis. Diam. 5,5, alt. 1,8 Mm. 

Planorbus albicans Pfeiffer Wiegm. Archiv I 1839 p. 354. 

Planorbus albicans Reeve conch. Icon. XX f. 117. (?) 

Gehause gedriickt, ober- und unterseits in de Mitte tief 
eigenesenkt, sehr fein gestreift, glanzend, hellhornfarben, Um- 
gange 4, rasch zunehmend, nach oben sehr gewolbt und durch 
eine sehr tiefe Naht getrennt; nach unten flacher und durch 
eine seichtere Naht verbunden; der letzte Umgang nur i/gmal 
breiter als der vorletzte; gegen die Miindung etwas herabstei- 
gend; Miindung breit-mondformig, weissgelippt, sehr schief; 
Mundsaum scharf, mit verbundenen Randern. Vaterland: Die 

^ Work made in cooperation with Institnto Oswaldo Cruz and Service 
Especinl tie Saiulc i'uhlica. Aided by the C'onsellio Nacional de Pes(|uisas of 
Brazil, which defrayed the expenses of a trip by the senior author to the 
type locality of A. albicans and also provided additional facilities for the 
study of the material collected there. 

NAin ILIIS 75(4) 


I. Ii)(i( liidoulcs icrun'us in niicl-A|)ril. Imiiuiuirc t'ollicles of (lie male 
I^oiukI in the niaiitle. H. and E. slain. X2l)(). 2. in June. Male follicles lilled 
with spermatozoa. Note the i^adiation from the perijiheiN of the follide lo 
ihe (enter, (iross section through the niesosoma. Malloi\'s Triple slain. \2()(). 
3. in mitl-Jiuie. Matiue female follicles adjaceni lo the so-called liver canals 
in the di;^esti\c di\erli( iila. H. and I-',, stain. X2()U. 4, in Jidy. General view 
of cross section in region of the gills. Note the presence of mature and im- 
mature eggs and the prominence of the nucleoli. H. and E. stain. X50. 



Fig. 1: Shell of Auslralot his albicans horn Laguna Somorostro, Havana, 
lig. 2: Raclula teeth of Aiislralorbis albicans from Lagiina La Canoa, Pinai 
del Rio {C'. = central, I intcinicdiale, L ^ lateral, M - marginal). Fig. 3: 
(ieiiital organs of Australorbis albicans from I.agiina La C.anoa. Pinar dci 
Rio (ag= fragment of albumen gland, ta ^ carrefoin-, ng - nidamenlal 
gland, od " proximal segment of ovispermidnct, od' = distal segment of 
ovispermiduct, ot - ovotestis, o\ ^" oviduct, po = pouch of oviduct, pp = 
prepuce, pr ^ prostate gland, ps = penis sheath, rm = retractor muscles of 
penial complex, sd - spermiduct, sp - spermatheca, .sv = seminal vesicle, 
ut = uterus, va - vagina. \tl \as deferens). 

April, 1962 nautilus 157 

Antillen, liisel Cuba, bei Cardenas; Portorico, St. Thomas. 
(Coll. Dunker) . 

Die Art gehort zur Sect. Armigerus, welche durch im Innern 
der Umgange angebrachte Zahne und Lamellen ausgeszeichnet 
ist. — Diese sitzen (meist zu 2 oder 3 an der Miindungswand, und 
zu 3 an der gegeniiber liegenden Gaumenwand) und zwar in 
ziemlicher Entfernung von der Miindung, so dass sie gewohnlich 
erst beim Zerstoren des Gehauses zu entdecken sind, wenn sie 
nicht durch die Umgange durchscheinend von aussen an der 
weissen Farbe erkannt werden. Auf der Miindungswand steht 
gewohnlich in de Mitte derselben ein grosserer starkerer Zahn, 
dem nach unten gegen die Ecken ein kleinerer sich anreiht. Auf 
der Gaumenwand befindet sich in der Mitte eine lamellen — 
oder faltenartige Schmelzleiste, wahrend zu beiden Seiten dersel- 
ben in nahezu gleicher Entfernung von ihr und etwa in der 
Mitte des Raumes von der Mittelleiste bis zu den Anschlussecken 
zwel Hocker — oder zahnartige Schmelzstiickchen angebracht 
sind. — Ueber den Zweck dieser Gehauseverstarkungen fehlen bis 
jetzt noch Beobachtungen." 

Among the planorbids collected by the senior author in Cuba, 
in December, 1956, there were specimens whose shell agreed, in 
shape and dimensions, to the two above transcribed descriptions, 
inclusive as to the apertural lamellae. Some of them, dissected 
for previous observation, proved young forms of a species 
anatomically undistinguishable from Planorbis peregrinus Or- 
bigny, 1835. As then shown (Paraense and Deslandes, 1958b), 
the shape of the shell underwent a process of remodelling along 
with growth, with eventual resorption of the five lamellae. 

A sample from Puerto Rico, sent us by Drs. F. F. Ferguson and 
Charles S. Richards in 1959, and regarded by them as albicans, 
was conchologically similar to the aforesaid Cuban material, 
but the specimens stopped growing when they reached about 
7 mm. in diameter, developed permanent lamellae and showed 
remarkable anatomical differences. 

Those facts led us to examine our remaining material, among 
which we found 1 1 specimens anatomically similar to the Puerto 
Rican ones. Of such specimens, 7 were collected at Laguna La 
Canoa (between Artemisa and Mangas, Candelaria, Province 
of Pinar del Rio), and 4 at Laguna Somorostro (Havana) . The 
largest specimens were 5 mm. in shell diameter, and only two 
from La Canoa showed apertural lamellae. 

Taking into account Pfeiffer's and Clessin's descriptions, and 

158 NAUTILUS Vol. 75(4) 

the fact that we are dealing with topotypic specimens, we agree 
with Drs. Ferguson and Richards in considering this species as 
Phinorbis albicans. Since the generic problem of the group to 
which it belongs was submitted to the International Commission 
on Zoological Nomenclature, the species is here placed in the 
genus Aiistralorhis, according to the reasons presented by Para- 
ense (1961). 

The following description is based on the above-mentioned 
Cuban material, of which four shells and one dissected specimen 
were deposited in the collection of Instituto Oswaldo Cruz (No. 
7907) . 

Description: The empty shell (PI. 16, fig. 1), is yellow or 
amber, finely oblicjuely striate, has about 31/9 whorls, and usually 
shows 6 internal lamellae deeply situated in the apertural region. 
The whorls widen rapidly, but at the region of the lamellae they 
stop expanding or even slightly decrease, and then rapidly widen 
toward the opening. The outer whorl shows various degrees of 
downward deflection at the apertural region. The upper (right) 
side is flattened and deeply umbilicate. It shows round smooth- 
walled whorls, of which the inner one is obscured at the bottom 
of the umbilicus. The under (left) side is concave and has a 
vortex-shaped central depression shallower than the umbilical 
one. On this side the whorls are bluntly carinate and the inner 
whorl is plainly visible at the bottom of the central depression. 
The aperture is directed forwards in young specimens, but as the 
shell grows larger the lamellae arise and gradually develop, and 
the aperture gradually bends downwards. In older specimens a 
callous thickening of the lip may be found. A complete set of 
lamellae consists of two parietal and four palatal units. Not only 
as concerns the lamellae, but also the shell, this species is similar 
to Australorbis janeirensis (see Paraense and Deslandes, 1956), 
and to young A. glabratiis and A. peregriniis from some popula- 
tions whose specimens may develop, at early age, a transient set 
of apertural lamellae (Paraense, 1957; Paraense and Deslandes, 

The animal shows no appreciable difference from that of other 
small species of the genus. The pseudobranch is simple, thin and 
flat. The rectal ridge extends from the pseudobranch into the 

April, 1962 nautilus 159 

pulmonary cavity, disappearing at the level of the stomach. There 
is no renal ridge. A dorsolateral ridge, facing the rectal one, runs 
parallel to the left side of the renal vein. 

The genital organs are shown in fig. 3. 

The ovotestis is composed of numerous sac-like diverticula, 
which are nearly always simple and less frequently bifurcate. The 
diverticula open into the ventral collecting canal, which con- 
tinues into the thin proximal segment of the ovispermiduct. 
Then follows the seminal vesicle, whose parietal diverticula 
are poorly developed, as in A. philippianus (see Paraense and 
Deslandes, 1958a). The seminal vesicle gradually merges into the 
distal segment of the ovispermiduct, which is about 3-4 times as 
long as the proximal segment. 

The spermiduct, highly sinuous at the beginning, follows a 
flexuous course in contact with the oviduct and, then, traverses 
the furrow formed by the pouch of the oviduct. At the point 
where it emerges from that furrow, it receives a single row of 
prostrate diverticula. The number of diverticula varied from 8 
to 16 in our material. They are mainly bi- or trifurcate, less fre- 
quently unbranched or arborescent. Although some of them have 
a short stalk, they are mostly sessile, their branches arising al- 
most directly from the prostate duct. The prostate duct continues 
into the vas deferens, which has no special characteristics. 

The penis sheath varies from a little longer to about 4 times 
as long as the prepuce. It contains a penis that shows no essential 
differences from that of congeneric species. The prepuce is only 
a little wider than the penis sheath and is internally separated 
from the latter by a muscular diaphragm. 

The oviduct is similar to that of other congeneric species, also 
having a large pouch of bosselated walls. The nidamental gland 
and uterus, taken conjointly, are from one and half times to 
thrice as long as the oviduct. The vagina is tubular and smooth- 
walled. The spermatheca is pear-shaped or club-shaped and has 
a narrow duct about as long as the spermathecal body. 

The jaw is somewhat T-shaped, consisting of a wider upper 
piece, vertically striate, and two narrower lateral pieces. The fol- 
lowing characteristics were observed in 5 specimens from Pinar 
del Rio (fig. 2) : radula formula, 14-1-14 to 15-1-15; horizontal 
rows, 80 to 90; central tooth bicuspid; 4 to 6 laterals, 2 to 3 

160 NAUTILUS Vol. 75(4) 

intermediates, 6 to 8 marginals. 

Comparison loitli related species: In a conchological diagnosis 
between A. albicans and other congeneric species, there will al- 
ways be place for uncertainty. In fact, even if no doubt arises as 
to the distinction between adult A. albicans and young shells 
from large species of Australorbis, there remain to be considered 
the similarities between the former and other lamellate shells. Of 
the Neotropical species studied by us, the adult A. janeirensis is 
always provided with a permanent set of apertural lamellae 
(Paraense and Deslandes, 1956) . Under special environmental 
conditions, young individuals from other species, such as A. 
glabratus and A. peregrinus, may develop a transient set of 
lamellae which are resorbed as the shell grows larger (Paraense, 
1957; Paraense and Deslandes, 1958b) . Thus, a reliable diagnosis 
must be based on the anatomical characters associated, of coinse, 
with those of the shell. 

A. albicans may be readily distinguished from A. glabratus by 
the absence of renal ridge, or of the pigmented line that precedes 
it in young specimens, and of the vaginal pouch (Paraense and 
Deslandes, 1955a, 1959) ; from A. tenagophilus (formerly called 
A. nigricans by us), A. peregrinus, A. andecolus and A. prorius, 
by the absence of tlie vaginal pouch (Paraense and Deslandes, 
1955b, 1957, 1958b, c) ; and from A. stramineus (^ centimetralis) 
by the absence of vaginal corrugation (Paraense and Deslandes, 

Despite the great similarity of its shell, when lamellate, to that 
of A. janeirensis, there are remarkable anatomical differences 
bet^\'een the two species (for comparison, see Paraense and Des- 
landes, 1956) . The penial complex, the vagina and the sperma- 
theca of A. janeirensis are very long, the penial complex being 
about half as long as the whole female duct (oviduct, nidamental 
gland, uterus and vagina together) ; moreover, the penis sheath 
is from 4.7 to 7.6 times as long as the prepuce. In A. albicans, the 
penial complex, the vagina and the spermatheca are relatively 
very short, the penial complex being from 14 to 1/7 (about 
1/^.5) as long as the female duct and, the penis sheath, from a 
little longer to about 4 times as long as the prepuce. 

Of the Australorbis species so far studied by us, the most closely 
related to A. albicans, from the anatomical standpoint, is A. 
philippianus (see Paraense and Deslandes, 1958a) . The latter 

April, 1962 nautilus 161 

also has a small penial complex, the length ot which is about 
1/^ that of the whole female duct (range from 1/2.5 to 1/7) . The 
two species may be distinguished by the characters of the penial 
complex and the prostate. The penis sheath is from a little longer 
to about 4 times as long as the prepuce in albicans; the two or- 
gans are about the same length in philippianiis, the ratio between 
the former and the latter varying from 0.9 to 1.5. And the number 
of prostate diverticula is much smaller in philippianiis: from to 
about 6, as against 8 to 16 in the present sample of albicans. 

Acknowledgement: We are indebted to the distinguished Cu- 
ban naturalists Miguel Jaume and Manuel Barro, for their help- 
ful assistance to the senior author during his work in Cuba. 

A description of the planorbid species Australorbis albicans 
(Pfeiffer, 1839) is presented. The following of its characters are 
considered as having diagnostic significance: 

Shell up to about 7 mm. in diameter and 2.5 mm. in height, 
usually with 6 apertural lamellae in the adult; whorls about 814, 
rapidly widening, bluntly carinate underneath. Absence of renal 
ridge. Ovotestis diverticula simple or, less frequently, bifurcate. 
Seminal vesicle with poorly developed diverticula. Prostate di- 
verticula mainly bifid or trifurcate, less frequently unbranched 
or arborescent, and arranged in a single row. Penis sheath from 
a little longer to about 4 times as long as the prepuce. Well de- 
veloped pouch of oviduct. Vagina tubular and smooth-walled, 
without pouch or corrugation. 

Clessin, S., 1884. Die Familie der Limnaeiden enthaltend die 

Genera Planorbis, Limnaeus, Physa und Amphipeplea. In 

MARTINI k CHEMNITZ: Systematisches Conchylien- 

Cabinet, v. 17. Bauer & Raspe, Niirnberg. 
Paraense, W. L., 1957. Proc. Malac. Soc. London, 32 (4): 175-179. 
. 1961. Shell versus anatomy in planorbid systematics. I. 

Australorbis glabratiis. Rev. Brasil Biol., in the press. 
Paraense, W. L. and Deslandes, N., 1955a. Mem. Inst. Oswaldo 

Cruz, 53(1) :87-103. 

. 1955b. Mem. Inst. Osioaldo Cruz, 53(1) :121-134. 

. 1955c. Rev. Brasil. Biol, 15(3) :293-307. 

. 1956. Rev. Brasil Biol., 16(1) :81-102. 

. 1957. Rev. Brasil. Biol., 17 (2) : 235-243. 

. 1958a. Rev. Brasil. Biol., 18(2) : 209-2 17. 

. 1958b. Jour. Conchyliol., 98 (3) : 152-162. 

. 1958c. Rev. Brasil Biol., 18 (4):367-373. 

. 1959. Am. J. Trop. Med. & Hyg., 8 (4) : 456-472. 

Pfeiffer, L., 1839. Arch. f. Naturgesch., I. Zool., 5 (1) :346-358. 

162 NAUTILUS Vol. 75(4) 


Department of Chemistry, University of Southern CaUfornia 

Burch (1959) has discussed a problem concerned with identity 
of two forms [Cf. plate 17] of Oliva spicata (Roding, 1798) from 
La Paz, Baja California, which were brought to his attention by 
Mr. Naylor of San Diego, who stated that one form was rejected 
by Indian artisans on the basis that they shatter while being 
worked, the other form being acceptable in making various arti- 
facts. Although at first glance both lots appeared identical, addi- 
tional study revealed differences which may be summarized as 

Used shells: Discarded shells: 

Base of columella Base of columella 

white or pink, light purple, 

more obese, more slender, 

lower spire. higher spire. 

For convenience, Burch temporarily identified the acceptable 
shells as Oliva spicata var. venulata, the discarded ones as var. 
ustulata, and concluded that, since the two forms were easily 
separable, the lumping of all these variations under one name 
was odd. O. venulata (Lamarck, 1811) and O. ustulata (La- 
marck, 1811) are usually included in the synonomy of O. spicata, 
e.g., by Johnson (1915) and Keen (1958). 

In the hope of throwing additional light on this question, we 
have examined, by the method of x-ray diffraction, samples taken 
from both forms. These samples were taken from broken frag- 
ments, and the locations from which the samples were taken is 
indicated in Fig. 1, which presents an equatorial cross section of 
a characteristic shell. Shells from both lots showed the structure 
shown in Fig. 1, i.e., a translucent outer layer which encloses the 
pigment, a somewhat thicker, white, inner layer which also en- 
closes pigment that, however, is not visible because this layer is 
thicker, and finally, a white, opaque, middle layer which con- 
tains no pigment. The outer and inner layers coalesce at the 
outer lip. 

Samples taken from the outer and middle layers of both forms 
showed almost the same x-ray powder diffraction patterns. All 

April, 1962 



Translucent outer layer 
Opaque middle layer 


I cm 

Fig. 1. Equatorial cross section of a characteristic specimen, showing the 
region from which the samples were selected. The dark areas represent the 
pigmentation, which was avoided in taking the samples for diffraction 

their major and minor features exactly match the pattern re- 
ported by Swanson, Fuyat, and Ugrinic (1954) for the aragonite 
polymorph of calcium carbonate. An important difference occurs, 
however, in the case of both kinds of samples from the form 
acceptable to the Indians in that faint additional lines character- 

164 NAUTILUS Vol. 75(4) 

istic of the powder pattern of the calcite polymorph of calcium 
carbonate (Swanson and Fuyat, 1953) may also be seen. We 
estimate that about 5% of calcite occurs in the outer layer, and 
somewhat more than half that much in the middle layers, of the 
utilized form. This calcite content is significantly different from 
that found in the samples from the form discarded by the artisans 
where none at all could be detected in the middle layer and only 
a trace in the outer layer. 

It would thus appear that, in addition to the differences in 
shape and color between the two forms, there is also a difference 
in chemical constitution. There does not seem to be any a priori 
explanation as to why this difference should result in the differ- 
ence in physical character noted by the artisans. Although this 
may or may not be relevant, calcite is considerably softer than 
aragonite (hardness 3, as compared to 4, Mohs scale) , but pos- 
sibly the shattering property may be the result of some other 
effect, e.g., differences in the orientation of the aystallites in the 

The question of which are the appropriate names to assign to 
these forms is discussed by Burch and Burch (1962) . 

Experimental Details. The x-ray diffraction patterns were re- 
corded photographically by the standard techniques in a Norelco 
powder camera, with CuKa radiation, filtered through nickel 
foil. All lines out to spacings of d > 1.15 A were measured. Rel- 
ative intensities were estimated visually. The 38 lines in this 
region reported for aragonite were all observed at the expected 
spacings and with the correct relative intensities for the four 
different types of samples. The calcite lines observed in the case 
of the samples from the "accept" form, all with very faint in- 
tensity, were those at d = 3.86 A and 3.035 A. These are two 
of the strongest lines reported by Swanson and Fuyat for calcite. 
All the other strong calcite lines listed by them are coincident 
with lines due to aragonite and were not observed. The only 
calcite line observed in the case of the samples from the outer 
layer of the "reject" form was the one at spacing 3.035 A, which 
is by far the strongest line of that pattern; the next most intense 
lines were too weak to be observed. The calcite content was 
estimated by comparison of the intensities of the 3.035 A line 
of calcite with the 2.871 A line of aragonite. 



Upper and lower, Icli ligs.. Iiolotxpc ol Ulii'a rejecla Burch .^- Burch. Upper 
aiul lower, right figs., Oliva I'onilaia Lamarck. 



Maxwell Smith, 1888-1961 

April, 1962 nautilus 165 

Fig. 1 was prepared by Mrs. Maryellin Reinecke. We wish to 
thank Mr. John Q. Burch for bringing this problem to our 
notice and for furnishing the specimens of both forms. 

Burch, J. Q. 1959. Minutes Conch. Club So. Cal. 184:20. 
Burch, J. Q. and Burch, R. L. 1962. [See next paper]. 
Johnson, C. W. 1915. Nautilus 2<?.-115. 
Keen, A. M. 1958. Sea Shells of Tropical West America. Stanford 

University Press, Stanford, Calif, p. 421. 
Swanson, H. E., and Fuyat, R. K. 1953. Nat. Bureau of Standards 

Circular 539, 2;51. 
Swanson, H. E., Fuyat, R. K., and Ugrinic, G. M. 1954. Nat. 
Bureau of Standards Circular 539, 3:53. 


The literature on the history and many variations of the 
species commonly known as Oliva spicata (Roding, 1798) is 
voluminous. The disposition of most modern authors has been to 
lump all varieties as forms of one species. In our opinion, the 
last word has not been written on this assemblage. 

In our work, Minutes of the Conchological Club of Southern 
California, no. 184, p. 20, Jan. 1959, we discussed the problem of 
the two species studied by Donohue and Hardcastle. Mr. W. E. 
Naylor of San Diego, a dealer in commercial shells, referred this 
problem to us. His trade in olives is largely to the Indians who 
use the shells to make various artifacts. The Indians rejected one 
variety claiming that they shatter when worked and are unsatis- 
factory. The acceptable variety is known to the trade as Oliva 
venulata Lamarck, 1811. These shells are heavier, with lower 
spire, and more obese than the discarded lot. These specimens 
consistently have the base of the columella with a white color 
and perhaps a tinge of pink. Shells of the rejected lot are more 
slender, higher spired, and the base of the columella is consis- 
tently a light purple instead of white. The interior of both is a 
bluish white. At first glance the shells in question seem to be very 
similar, but after a little study they readily separate into two 
forms. The studies of Donohue and Hardcastle have shown that 
the two forms differ in the chemical analysis of the shell. Both 

166 NAUTILUS Vol. 75(4) 

forms are found on the same tide flats at La Paz, Baja California, 
Mexico, with no intergrades in many hundreds of specimens. 
Certainly many species are recognized upon much more meagre 

In our preliminary discussion, for convenience, we assigned 
the discarded form to Oliva ustulata Lamarck, 1811. Several very 
diff^erent forms have been recognized by various authors under 
this name. Tryon in the Manual of Conchology thought it to be 
a variety of the Atlantic Olivn reticularis Lamarck, 1811. Charles 
Johnson thought it to be a form in which the pattern of the 
shell is obscured by a dark brown layer. In our opinion, the name 
Oliva ustulata Lamarck cannot be used for this form. Seemingly 
the species of Oliva under discussion has been considered a fonii 
of the typical by all authors we have consulted. Therefore, we 
feel justified in giving it a name and the status of a new species. 

Oliva rejecta, new species. Plate 17, left figs. 

Shell cylindrically oblong; spire exserted, prominent; color 
white with a closely reticulate pattern of chocolate; base of 
columella purple, swollen, with 4 plaits; interior of aperture 
bluish white. Length: 35.5 mm. Greatest diameter: 15 mm. Type 
locality: tide flats at La Paz, Baja California. 

A comparison of the dimensions of the type of the new species 
with those of a normal specimen of Oliva vemilnta Lamarck is 
of interest. Both specimens measure exactly 35.5 mm. in length, 
but the greatest diameter of Oliva rejecta is 15 mm., while that 
of Oliva venulata is 17.5 mm. Also, the gieatest diameter of 
Oliva rejecta is much lower on the body whorl. 

The holotype is to be deposited in the California Academy of 
Sciences, San Francisco, no. 12400. Fifty paratypes will be dis- 
tributed to various institutions. 


BuLiMULUs DEALBATUS joNESi Cleuch appears to be Bulimulus 
mooreanus (Pfeiff^er) which has been introduced into Alabama, 
It is somewhat smaller than is usual for the species in Texas, but 
is otherwise typical. Unlike B. dealbatus (Say) , it estivates on 
the weeds where it can be found in large numbers even in the 
hottest part of the summer. — Leslie Hubricht. 


liJH 17XP T