THE ^oy/

NAUTILUS

THE PILSBRY QUARTE]

OF CONCHOLOGISTS

VOL. 83

JULY, 1969 to APRIL, 1970

EDITORS AND PUBLISHERS

R. TUCKER ABBOTT

Delaware Museum of Natural History, Greenville, Delaware

HORACE BURRINGTON BAKER

Professor Emeritus of Zoology, University of Pennsylvania

CHARLES B. WURTZ

La Salle College, Philadelphia, Pa. 19141

MRS. HORACE B. BAKER

11 Chelten Road

Havertown, Pennsylvania 19083

PONY PRINTING, UPPER DARBY, PA.

THE NAUTILUS

Vol. 83 July, 1969 No. 1

DONAX FOSSOR: A SUMMER RANGE EXTENSION

OF DONAX VARIABILIS1

By PAUL CHANLEY2

Virginia Institute of Marine Science, Gloucester Point, Virginia 23062

In 1822 Say described two species of Donax living in die surf

zone of ocean beaches on the east coast of the United States. Donax

variabilis, the southern form, occurring in Georgia and east Flori-

da, while the more northern Donax fossor was reported from Mary-

land and New Jersey. In the original description there is very little

to differentiate the two "species." The shell of D. variabilis was

reported as "triangular; anterior margin obliquely truncated, cor-

date, suture a little convex; posterior hinge margin nearly recti-

linear." The shell of D. fossor was described as "subtriangular;

anterior margin short and rounded; posterior hinge slope recti-

linear." Other features of shape were described identically. The

size of D. fossor was given as, "Breadth from half an inch to three-

fifths." D. variabilis was described as "Length half an inch. Width

nine-tenths of an inch. Thickness seven-twentienths of an inch."

D. fossor reportedly had "yellowish" and "whitish" varieties while

the "usual varieties" of D. variabilis were "red, white, yellow or

elegantly radiated with dilated reddish brown lines upon a white

or yellow ground."

Since this original description the range of both species has been

extended considerably. D. variabilis is found from Virginia Beach

to Florida and Texas, while D. fossor is found from Long Island

to Cape Hatteras, North Carolina (J. P. E. Morrison, personal

communication) .

The two "species" are currently distinguished by most authori-

ties on the basis of size, shell color, sculpturing and thickness of

the valves at the anterior end. D. variabilis reaches a maximum

length of three-fourths of an inch, occurs in an almost infinite

variety of colors, has a marked gradient of radial shell sculpture

1 Contribution No. 312 from Virginia Institute of Marine Science.

- Present Address: Marinus Laboratories, P. O. Box 1330, Melbourne, Florida

32901.

1

2 nautilus Vol. 83 (1)

that is more pronounced posteriorly, and a thickening of the valves

at the anterior end to produce "lips." D. fossor reaches a maximum

size of one-half an inch, reportedly occurs in only two color phases,

has smooth radial sculpturing over the entire shell and does not

have the thickened valves at the anterior end.

Some authors have accepted the validity of these two species or

have at least treated them as separate species (Johnson, 1934;

Morris, 1947; Miner, 1950) . A few have suggested that the two are

the same species. Abbott (1954) , in describing D. fossor says,

"Subsequent biological studies may show that this common beach

species is a subspecies or cold-water form of variabilis. Young speci-

mens of variabilis from several southern states look suspiciously

like this so-called species." Jacobson and Emerson (1961) in dis-

cussing the periodic appearance and disappearance of D. fossor on

Long Island stated "Because of this strange periodicity, some mala-

cologists speculate that what we call D. fossor is merely D. varia-

bilis (a large Florida Donax) , the larvae of which have somehow

been swept far to the north and developed poorly in our unfavor-

able waters. It is fairly certain that D. fossor does not survive the

winter in our area." These remarks plus personal observation of

the periodic occurrence of Donax on the Eastern Shore of Virginia

prompted this investigation of D. fossor and D. variabilis to deter-

mine if they are two valid species.

This study would not have been possible without the cooperation

and help of Dr. J. P. E. Morrison of the U. S. National Museum,

Dr. Arthur H. Clark of the National Museum of Canada (who

provided the information on the Canadian collections) , Dr. R.

Tucker Abbott of the Delaware Museum of Natural History, as

well as Dr. Ruth Turner and Dr. Kenneth Boss of the Museum

of Comparative Zoology of Harvard University. Mr. M. Karl

Jacobson's data, suggestions, cooperation and encouragement were

also of utmost importance. I am especially grateful to Mr. Wade

B. Smith of the University of North Carolina for allowing me to

use the excellent data from his thesis. Finally, and perhaps most

important, I am indebted to the many members of the American

Malacological Union who so thoughtfully provided the data es-

sential to this study from their own collections.

July. 1969 nautilus 3

Materials and Methods

As a first step in this investigation a questionnaire was mailed to

several members of the American Malacological Union requesting

as much information as possible about their private collections of

Donax from the East Coast of the United States, specifically the

date and location of collections as well as the size of the specimens.

Comparable data were obtained from the collections of the U. S.

National Museum, the Academy of Natural Sciences of Philadel-

phia and the Museum of Comparative Zoology of Harvard

University.

In order to estimate the time and duration of the spawning

season, the gonad condition of Donax was determined whenever

possible by dissecting fresh material and histological sectioning of

preserved specimens. The presence of small (2-3 mm) clams in

collections was also considered evidence of recent recruitment. To

determine the length of the pelagic larval period, Donax were

spawned in the laboratory and their larvae reared to meta-

morphosis.

The direction and velocity of coastal currents that might in-

fluence the distribution of clams were determined by reviewing

pertinent literature.

Results

Donax have been collected along the Atlantic Coast of the

United States from Florida to Long Island and Connecticut. From

Virginia northward, Donax have apparently never been collected

alive from January to May 30 (Table I) . On Long Island they

cannot be found every year and sometimes as many as twelve years

elapse between appearances (New York Shell Club Notes, 1967) .

They have been collected year round in North Carolina and

southward.

North of North Carolina there is a general trend for the average

size of Donax to increase as the season progresses, as determined

by periodic observations throughout a season at one location in

New York, New Jersey and Virginia (Table II) . No such trend

is evident in collections of Donax from North Carolina.

Gonad studies of clams were limited to those taken on April 4

and August 31 in North Carolina; from July to October on the

Eastern Shore of Virginia; and on August 8 at Ocean City, New

4

NAUTILUS

Vol. 83 (1)

Jersey. Usually only heavily parasitized clams lacked mature eggs

or sperm in gonads. The occurrence of 3 mm and smaller clams

from March to December in Smith's (thesis) North Carolina

collections is further evidence of almost continual recruitment.

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Under laboratory conditions larval Donax variabilis increased

in length from 70 n to a maximum of 350 /i during their pelagic

period (Chanley, 1969). This is a greater increase in size during

the pelagic stage than is found in larvae of most bivalves. The first

larvae to metamorphose in the laboratory began "setting" at an

age of about 3 weeks. Another brood of larvae, reared under less

desirable conditions, remained pelagic for 56 days without meta-

morphosing. Mr. William Tiffany, III, of Florida State Univer-

sity, has also reared Donax larvae and has found that under

laboratory conditions some larvae remain pelagic for up to 50

days. Rate of larval development is highly dependent on water

temperature. Since laboratory temperatures were higher (20-25°

C) than those in nature when D. fossor first appear as 5 mm clams

(20° C) , it seems probable that larvae could be pelagic for even

longer periods. A reasonable estimate would be that some larvae

remain pelagic at least 6 to 8 weeks in spring and fall with a

minimum two-week pelagic period under optimum summer con-

ditions.

Dispersal of the veliger larvae of marine bivalves is accom-

plished primarily by water currents. The pertinent water currents

in the geographic range of Donax are the Gulf Stream and several

less stable inshore current and eddy systems. The Gulf Stream

moves north at about 50 miles per day (Svedrup et ah, 1942) .

Harrison et al. (1967) found the inshore current systems unstable

but nonetheless capable of carrying drift bottles southward and

shoreward from release point at average velocities of up to 24 miles

per day or north at average velocities up to 1 1 miles per day

(Table III).

Discussion

Many factors must be considered in analyzing the distribution

data. Several collections have been made of dead, even fossil, shell.

They are of no value in establishing seasonal distribution because

it is possible that these shells were transported to the point of

collection after death of the clams. The inclusion of these "irrele-

vant" collections in the study is thought to be less detrimental than

exclusion of any possibly pertinent data. Consequently, all dated

collections have been used in this analysis unless the collector has

specifically stated that clams were dead at time of collections.

NAUTILUS

Vol. 83 (1)

The average lengths of clams must also be viewed with caution.

Most collections do not represent random samples of a population

and several collections contain too few specimens to get a valid

estimate of average size. Finally, rates of growth may vary con-

siderably at different levels of the intertidal zone or in different

local colonies.

Considering these possible sources of error it is suprising how

definitive the distribution and size data are. The only two northern

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July. 1969 nautilus 7

(Long Island to Virginia) colled ions from January to June are

the McCallum collection from Assateague Island in April and the

Weingartner collection on 30 May from Beach Haven Inlet, New

Jersey. The McCallum collection was of dead hinged valves on

the bayside of the island. Since experimentally planted hinged

valves of Donax can persist on the beach for several months

(Loesch, 1957) , it seems likely that these valves were of clams that

had died the previous winter. On the other hand the clams in the

Weingartner collection were living at the time of collection and

were so large, averaging 13.6 mm in length, it seems likely they

had survived the previous winter and were not an example of 1956

recruitment. This collection is the only record of living Donax

north of Virginia before July. The Van Pelt collection in June

was of beach-worn single valves.

Perhaps the most reasonable explanation for this peculiar dis-

tribution of Donax would be that D. fossor are really D. variabilis

carried north as larvae in spring, summer and fall. These larvae

survive in seasonably warm temperatures and develop into small

clams on northern beaches. Growth and even reproduction con-

tinues until temperatures drop in the fall. Clams then become

sluggish, unable to maintain themselves in the surf zone and are

eventually killed. Living clams are not found again until subse-

quent recolonization occurs. Only rarely do conditions permit

isolated populations, such as that sampled by Weingartner, to sur-

vive the winter.

This would not be a unique phenomenon. Coe (1953), on the

west coast, observed, "By means of these currents the geographical

range of a species with pelagic larvae may be extended in favor-

able seasons far beyond the usual limits but only to have the range

restricted again when more vigorous conditions return. The popu-

lation will thereby fluctuate irregularly in distribution ."

Frequently the autumnal disappearance of Donax occurs while

water temperatures are as high as the spring temperature when the

clams first appeared. Certainly temperatures are usually higher at

the time of disappearance than winter water temperatures in

North Carolina where Donax can be found throughout the year.

Consequently factors other than temperature may also be involved.

For example, in late summer and early fall Spisula solidissima

larvae frequently settle along northern ocean beaches and tern-

8 nautilus Vol. 83 (1)

porarily establish dense populations of juveniles in Donax habi-

tats. Direct competition with juvenile S. solidissima plus increased

predation by birds, crabs, racoons and other predators contribute

to the demise of Donax populations. Such pressures would be ab-

sent in spring and early summer when smaller, less conspicuous

Donax colonize northern beaches and S. solidissima are absent.

5. soldissima are less numerous on the beaches of North Carolina,

near the southern limit of the range of this species.

Loesch (1957) observed that, in Texas, Donax moved into knee-

deep water in December. I have found no evidence to suggest that

offshore migration accounts for the fall disappearance of Donax

north of North Carolina. In fact, reports of the simultaneous

appearance of empty valves and disappearance of living clams

strongly suggests death of the clams.

The hypothesis that D. fossor is a summer range extension of

D. variabilis would not only account for the seasonal distribution

of Donax in the mid-Atlantic region but would also account for the

sporadic appearance of Donax on Long Island. Colonization of

beaches at this northern limit of the range would depend on ir-

regular fortuitous combinations of warm water currents that do

not occur annually.

Table III. Inshore Non-tidal Current Direction and Velocity of Northern Chesa-

peake Bight (From Harrison et al., 1967).

July, 1969 nautilus 9

It would also account for the seasonal increase in average size

of northern Donax that does not occur in North Carolina. Since

all clams are of the one-year class their initial average size would

increase during the year. In the south this effect would be masked

by the presence of the previous year class and almost continual

recruitment. Large North Carolina Donax frequently have a pro-

nounced "growth" ring indicating that cessation of growth oc-

curred seasonally. "Growth" rings are rare on more northern

clams and when they do occur are probably indicative of indi-

vidual cessation of growth due to injury, disease or some other

non-seasonal causes. Bi-modal length-frequency distributions of

northern populations would indicate discontinuous recruitment

during the same season rather than two year classes.

The minor conchological differences between D. fossor and

D. variabilis could be those of a population that does not over-

winter compared to a population that does. They could also be

induced by temperature or other environmental conditions. Wade

(personal communication) has found that "different factors

throughout the life cycle have effects which may give rise to much

variability in shell morphology" of Donax denticulatus. Nicol

(1967) noted that cold-water species of pelecypods had a general

lack of color and subdued ornamentation. A comparable intra-

specific phenomenon of less magnitude could account for the con-

chological differences between northern and southern Donax. The

occurrence of "both species" on the same beach would indicate

that the two morphological types had developed at different sea-

sons or under different environmental conditions rather than as

different species.

The length of the larval period of Donax plus the velocity of

ocean currents show that it theoretically is possible for larvae to

be transported from 700 to 2500 miles by the Gulf Stream during

their pelagic period. The lengthy spawning period provides ample

opportunity for larvae to be in the water for transport to the north

during months they can survive. Thus annual repopulation of

northern beaches by current transport of larvae from the entire

Atlantic range of D. variabilis appears feasible.

After the relatively lengthy larval period clams become sexually

mature rapidly and may reproduce in a few weeks. Juveniles have

doubled in size in the laboratory one week after setting. Loesch

10 NAUTILUS Vol. 83 (1)

(1957) found sexually mature males and females only 4 mm long.

Consequently, clams that have colonized a beach in late spring or

early summer are capable of producing larvae that can repopulate

still another beach the same summer. Therefore, recolonization of

northern beaches is also possible by serial advance up coast.

It is more difficult to defend the hypothesis that D. fossor and

D. variabilis are distinct species. The absence of D. fossor from

December through May might be accounted for by lack of col-

lecting effort during these months. However collecting effort should

be greater in April and May when clams are not found, than in

die fall when they have been found. It also seems unlikely that

these clams assume habits that make them immune to collection

from January to June but susceptible to capture the rest of the year.

If the two forms are distinct species the increase in average size

during the season would indicate that clams "winter-over" as

larvae or minute juveniles and resume growth when temperatures

rise. They would then die as temperatures drop the following

winter. This hypothesis does not seem likely since it requires the

assumption that larvae and minute juveniles can survive the

winter while half-grown and "adult" D. fossor cannot.

Finally Donax reproduce by releasing gametes freely into the

water. The spawning periods of the two forms overlap and both

have been collected from the same beach at the same time. It is

difficult to understand how speciation could have occurred when

there is no geographic or physical barrier and no selectivity in

mating. Some mechanism for prevention of interbreeding would

have to be demonstrated before the hypothesis that speciation has

occurred could be accepted.

Conclusions

D. variabilis larvae are pelagic for several weeks and are ex-

posed to water currents capable of transporting them great dis-

tances. In the summertime they would be expected to colonize

northern beaches that are too cold for the species in wintertime.

The distribution and growth of D. fossor are difficult to explain if

D. fossor is a distinct species, yet they are precisely what one would

expect of a summer range extension of D. variabilis. The conclu-

sion is that D. fossor is D. variabilis and that the minor conchologi-

cal differences, now used to separate them by many malacologists,

are environmentally induced.

July, 1969 nautilus II

Locality Records

The dated collection records that have been obtained are listed

as follows: Source of information (NMC, USNM, ANSP, MCZ,

PC represent, respectively, National Museum of Canada, United

States National Museum, Academy of Natural Sciences of Phila-

delphia, Museum of Comparative Zoology of Harvard University,

and Private Collection) , collector, location, year, days and months

of collection and remarks. Remarks are enclosed in parentheses

and apply only to the collection immediately preceding them.

New York

USNM: Merriam, Jones Beach, 1883 — 24 Sept. NMC, MCZ

and PC: M. K. Jacobson, Rockaway Beach, 1949 — July; 1954 —

25 July; 8, 13, 26, 29 Aug.; 6, 14, 27, 28 (collected dead) Sept.;

3 (only 3 specimens) ; 10-12 Oct. (May have been collected dead) ;

28 Nov. (Beachworn valves) : 1955 — 12, 19 July; 1, 3 Aug. PC:

E. Hartstein, Fire Island, 1967 — Mid-Aug. (present only one

day) .PC: V. Johns, Atlantic Beach, 1967 — 4 Sept. (indirect re-

port) . PC: J. Foehrenbach, Fire Island, 1968 — July (No Donax

found) .

New Jersey

ANSP: L. Woolman, Holly Beach, 1894 — 7 Oct. ANSP: P. H.

Brcnun, Wildwood, 1896 — July (single valves, probably fossil or

subfossil). ANSP: H. A. Pilsbry, Sea Isle City, 1901 — July.

ANSP: 5. S. Van Pelt, Cape May County, 1907 — 9 June (Beach-

worn single valves) . ANSP: B. Long, Ocean County, 1912 — 15-

18 Sept. (only 2 specimens). ANSP: D. McCadden, Ocean City,

1916 — 2 Aug. ANSP: H. A. Pilsbry, Cape May, 1917 — Aug.

ANSP: H. R. Roberts, Stone Harbor, 1939 — Aug. ANSP: M.

Wagner, Beach Haven Inlet, 1940 — 5 July (3 Beach-worn single

valves) . PC: R. E. Weidman, Barnegat Bay, 1947 — Aug. ANSP:

R. A. Heilman, Cape May County, 1953 — 12 July: 1954—12

Aug. PC: M. Weingartner, Beach Haven Inlet, 1956 — 30 May.

PC: G. and M. Kline, Wildwood Crest, 1966 — 8 Oct. PC: R.

Lowden, Ocean City, 1967 — 31 July; 1, 9 Aug.; 3 Sept.; 15 Oct.

(two distinct size groups) . PC: M. Hulswit, S. Hereford Inlet,

1967 — 11 Aug. PC: C. Fisher, Ocean City, 1968 — 8 Aug.

R. Fisher, Ocean City, 1968 — 2 Sept. (Donax present, not col-

lected) ; 1 Oct. (No Donax present) .

Delaware

USNM: Richards, Delaware Bay near Lewes, 1931 — 4 Aug.

(one specimen) .

Maryland

USNM: H. A. Rehder, Ocean City, 1945 — 3 Aug. USNM:

/. P. E. Morrison, Ocean City, 1963 — 7 July. PC: M. Hulwit,

Assateague Island, 1967 — 15, 16 Aug. (two distinct size groups.

Larger clams rare.)

12 NAUTILUS Vol. 83 (1)

Virginia

USNM: Fish Hawk, Lynnhaven Rd., 1916 — 15 July (probably

not live collected) ; Magothy Bay. 1916 — 22 July (3 shells) ; Vir-

ginia Beach, 1916 — 25 July (1 valve). ANSP: H. G. Richards,

Cape Charles, 1929 — 16 June (3 single valves) ; Cape Henry,

1931 — 29 Aug. (3 single valves) . MCZ: Clench, McCoy and

Russell, 10 miles S. Virginia Beach, 1934 — Aug.-Sept. USNM

/. P. E. Morrison, Northampton County, 1935 — 6 July; Vir

ginia Beach, 1946 — 22 Sept.; 1950 — 23 July. USNM

W. H. Ball, Virginia Beach, 1943 — 5 Oct. ANSP: R. 6- H. R

Robertson, Cape Henry, 1962 — 22 July. PC: P. Chanley, Cedar

Island, 1963 — 16 July to 8 Oct. (clams collected twice weekly)

1964 — 26 June; 9, 14, 21, 22, 27 July; Aug.; 17 Sept. PC: G. & M

Kline, Assateague Island, 1966 — 25 Nov. PC: G. McCallum, As-

sateague Island, 1968 — 18 April (dead shells collected on bay

side of island) . PC: M. Castagna, Near Wachapreague Inlet, 1968

— 10 June (none found) ; 17, 24 June; 16, 22 July; 1, 6, 14, 28

Aug.; 3, 16, 23, 30 Sept.; 7, 14 (none found), 21 (none found),

28 (none found) Oct.

North Carolina

USNM: Shackleford Beach, 1899 — Aug. USNM: C. W. Stiles,

Southport, 1922 — Sept. (single valve) . MCZ: /. 5. Colman, Beau-

fort; USNM: C. W. Cooke, Cape Hatteras, 1936 — May. MCZ:

H. A. Vander Schallie, Morehead City, 1937 — Sept. ANSP: R. W.

Kindle, Atlantic Beach, 1938 — 1 Nov. (6 valves) . USNM: A. S.

Pearse, 1939—17 June. ANSP: R. A. McLean, Bogue Bank, 1942

— 17 July. USNM: W. H. Ball, Nag's Head, 1943 — 29 Sept.;

Dare County, 1943 — 20 Oct. USNM: /. P. E. Morrison, Cape

Hatteras, 1943 — 20 Oct.; Nag's Head, 1957 — 26 Aug.; Cape Hat-

teras, 1957 — 29 Aug.; Emerald Isle, 1960 — 9 June. PC: W. B.

Smith, Bogue Bank, 1967 — 23, 25, 27, 28, 31 July; 18 Aug.; 11,

12 Sept.; 15 Oct.; 18, 19 Nov.; 16, 17 Dec; 1968 — 19, 20, 27, 28

Feb.; 11, 14, 15, 18, 19 March; 12, 16, 17, 18 April; 13, 14, 16,

17, 22, 23, 24 May; 18, 19 June; 1, 3 July; 6, 7, 8, 19, 20 Aug.

PC: P. Chanley, Bogue Bank, 1968 — 4 April. PC: P. Jennewein,

Wrightsville Beach, 1968 — 19 May. PC: D. Armstrong, Nag's

Head, 1968 — 31 Aug.

South Carolina

MCZ: Charleston, 1885 — May. MCZ: E. B. Chamberlin, Wav-

erly Mills, 1929 — Sept. (Beach worn single valves) . MCZ: C. W.

Johnson, Folly Beach, 1931 — 27 March (single valves) ; ANSP;

F. Harper and A. N. Leads, Myrtle Beach, 1933 — 5, 6 April (9

single valves, probably not live collected) . ANSP: E. M. Kindle,

Myrtle Beach, 1938 — Nov. (Single valves) . ANSP: H. G. Rich-

ards, Ocean Drive Beach, 1941 — 10 May. ANSP: L. Patterson,

Myrtle Beach, 1941 — June (3 single valves) . ANSP: Coventry

and McLean, Ashe Island, 1941 — 25 June (1 complete specimen

July, 1969 nautilus 13

and one single valve) . USNM: Myrtle Beach, 1946 — Sept.

USNM: H. A. Rehder, Myrtle Beach, 1946 — Oct. ANSP: H. G.

Richards, Myrtle Beach, 1947 — 4 March (single valves) . USNM:

/. P. E. Morrison, Myrtle Beach, 1955 — 3 Dec; 1959 —

22 Nov.; 1960 — 7 June; Windy Hill Beach, 1960 — 7 June; Paw-

ley's Island, 1960 — 15 Sept.; Near Little River Inlet, 1960 — 25

Sept. MCZ: A. S. Merrill, Racoon Key, 1963 — 1 July (1 drilled

valve). MCZ: G. McCallum, Pawley's Island, 1966 — 14 Aug.

MCZ: B. & D. McCallum, Litchfield Beach, 1966 — Aug. (Single

valves) .

Georgia

MCZ: Clench, Archer and Rehder, St. Simons Island, 1931 —

July, Aug. USNM: H. A. Rehder, St. Simons Island, 1938 — Oct.

ANSP: McLean and Coventry, Sea Isle, 1941 — June. NMC:

R. H. Croker, Sapelo Island, 1964 — 27 Nov.

Florida

MCZ: Clench and Okkelberg, St. Augustine, 1929 — 19 July.

USNM: B. C. Marshall, Near Mclntyre, 1931 — July (3 speci-

mens) . MCZ: R. W. Foster, Ormond Beach, 1934 — March; Key

Biscayne, 1936 — Feb. (2 specimens) Cape Canaveral, 1937 —

April (3 specimens) . USNM: /. P. E. Morrison, Duval County,

1936 — 27 April. MCZ: E. M. Davis, Fernandina, 1936 — Oct.

(single worn valves) . USNM: M. MacNeil, Volusia County, 1959

— 19 March (4 specimens) . USNM: R. O. Christenson, Daytona

Beach, 1939—19 April. ANSP: F. Harper, Ponte Vedra, 1941 —

20 May. MCZ: R. T. Abbott, Duval County, 1942 — Oct. ANSP:

E. M. Kindle, St. Johns County, 1947 — 10 March (single speci-

men) . MCZ: H. D. Athearn, Flagler Beach, 1949 — May (6 speci-

mens). NMC: P. Hooper, Daytona Beach, 1955 — 14 July.

USNM: /. P. E. Morrison, Daytona Beach, 1959 — 20 Nov.

Literature cited

Abbott, R. T. 1954. American Seashells. Van Nostrand, Prince-

ton, N. J. 541 p.

Chanley, P. 1969. Larval development of the coquina clam, Donax

variabilis with a discussion of the structure of the larval hinge

in the Tellinacea. Bull. Mar. Science 19(1) : 214-224.

Coe, W. R. 1953. Resurgent populations of littoral marine inverte-

brates, and their dependence on ocean currents and tidal cur-

rents. Ecology 34(1): 225-229.

Harrison, W., J. J. Norcross, N. A. Pore and E. M. Stanley. 1967.

Circulation of shelf waters of the Chesapeake Bight. Professional

Paper 3: 1-82. Environmental Sciences Services Administration.

U. S. Dept. Commerce, Washington.

Jacobson, M. K., and W. K. Emerson. 1961. Shells of the New York

City Area. Argonaut Books, Larchmont, N. Y. 142 p.

Johnson, C. W. 1934. List of marine mollusca of the Atlantic coast

14 nautilus Vol. 83 (1)

from Labrador to Texas, Proc. Boston Soc. Natural History

40(1): 1-204.

Loesch, H. C. 1957. Studies of the ecology of two species of Donax

on Mustang Island, Texas. Publ. Inst. Mar. Sci., Univ. Texas,

4(2): 201-227.

Miner, R. W. 1950. Field book of seashore life. G. P. Putnam's

Sons, New York. 888 p.

Morris, P. A. 1947. A field guide to the shells. Riverside Press.

Cambridge, Mass. 236 p.

New York Shell Club Notes. September 1967. Donax on Long

Island. No. 134: 2.

Nicol, D. 1967. Some characteristics of cold-water marine pelecy-

pods. Paleontol. 41(6): 1330-1340.

Say, Thomas. 1822. An account of some of the marine shells of the

United States. Acad. Nat. Sci. Phila. J. 2: 302-325.

Smith, W. B. Thesis (In preparation) . University of North Caro-

lina.

Sverdrup, H. V., M. W. Johnson and R. H. Fleming. 1942. The

oceans, their physics, chemistry and general biology. Prentice

Hall, Englewood Cliffs, N. J. 1087 p.

TWO INFRASUBSPECIFIC FORMS IN OLIVA

By ROWLAND F. ZEIGLER

305 West Palmetto St., Florence, South Carolina

There is perhaps more disarray in the genus Oliva than in any

other group of shells. The writer, for the past several years, has

been reviewing literature and studying collections both private

and in museums, hoping that these efforts will culminate in the

publication of a book with natural color photographs and proper

synonymies. A contemporary review should be a welcome reference

for the understanding and clarification of this misunderstood and

neglected genus. Two shells have been encountered in these studies

that appear deserving of new forma names. One of these is well-

known but without an available name, while the other is quite

uncommon and has been overlooked in the past. These new names

are being proposed at the infrasubspecific level, and do not enter

into the literature as pre-occupying subspecific names. Nor are they

available as valid specific names.

Olive (Oliva) incrassata Lightfoot, 1786 (= O. angulata Lamarck,

1811)

The well-known Angled Olive, the heaviest member of the

genus, varying in size from 50 - 88 mm. in length and up to 40 mm.

July, 1969 nautilus 15

in width, is quite common and ranges from Lower California,

throughout the Gulf of California, and down the Pacific coast to

Peru. Many of these shells in the Gulf of California contain vary-

ing amounts of a brilliant golden-yellow coloration intergrading

to a pure golden form. The only named color form is:

Oliva incrassata forma nivea Pilsbry, 1910

The rare pure-white albino found only in the northern end of

the Gulf of California. (A rare unnamed glossy black form is also

occasionally found.)

Oliva incrassata forma burchorum new form. Fig. 1.

This well-known, but heretofore unnamed form, is completely

golden. It is felt that this form is worthy of a form name, just as

the golden form of O. sayana Ravenel, 1834, on the Florida coasts

merits the name O. sayana forma citrina Johnson, 1911. In its

juvenile stage, the color is a deep-orange, tending to become more

yellowish as the shell matures and develops its angular shoulder.

Animals of both the golden and white forms are the usual buff

color of typical O. incrassata. It is completely devoid of markings,

rare, much-prized, and is known only from the northern end of the

Gulf of California. Range — longitudes 113° west — 115° west,

and latitudes 30° north — 32° north.

The holotype is in the American Museum of Natural History,

New York City, No. 147749. The type locality is San Felipe, Baja

California, Mexico. One paratype is in the collection of Mr. H. C.

Porreca, West Henrietta, N. Y., and three are in the writer's col-

lection, including the hgured specimens.

The new name burchorum is proposed to honor John Q. and

Rose L. Burch of California. No one in recent times has studied,

written about, or contributed more to our knowledge of the genus

Olwa than the Burch's. Their studies and voluminous notes on

Oliva in the Minutes of the Conchological Club of Southern Cali-

fornia (1958-60) have been of inestimable value. The Burch's re-

viewed and assembled the literature, and catalogued the Recent

and Fossil Olives in 1960. They concluded that "there are a num-

ber of species to be described as new, and others that may prove to

be variants of one species. These should be studied with care and

published with proper illustrations." Their work, as well as their

encouragement and assistance through personal communications,

has been an inspiration.

16 nautilus Vol. 83 (1)

Oliva (Oliva) tremulina Lamarck, 1811

This Indo-Pacific species has been controversial, as it is almost

identical in appearance with O. miniacea Roding, 1798, long

known as O. erythrostoma Lamarck, 1811 (non O. erythrostoma

Meuschen, 1787, non binomial) . The primary differentiating fea-

ture is that O. tremulina has a fleshy white aperture, while O. mini-

acea always has a deep reddish orange aperture. There are some

parallel color variations in both species. O. tremulina like O. mini-

acea is elongated cylindrical, large, rather thick, longitudinally

wave streaked and banded. Johnson and some other authors con-

sidered O. tremulina to be a form of O. miniacea. Burch felt that

tremulina, in addition to the differentiating fleshy white aperture,

was less swollen at the posterior or shoulder. He felt that they were

otherwise close, but separable. Dautzenberg considered O. tremu-

lina a valid species and named a color form. Oliva nobilis Reeve,

1850, is a synonym.

Distinct and recognized color forms of O. tremulina are:

Oliva tremulina forma chrysoides Dautzenberg, 1927.

A solid golden or orange form, which may have a very faint pat-

tern. It is like O. miniacea forma sylvia Duclos, 1844, except for the

white aperture.

Oliva tremulina forma concinna Marrat, 1871.

Uniform blackish brown in color, or sprinkled with a few tri-

angular markings. Its counterpart is O. miniacea forma marrati

Johnson, 1914.

Oliva tremulina forma fumosa Marrat, 1870.

Dautzenberg described this shell as "intermediate between typi-

cal O. tremulina and the variety concinna. The ground color is

tawny-gray and the dark and cloudy pattern gives to the shell a

smoky aspect. In certain examples, broad brown patches spread

over much of the surface. The aperture is white and the lip is

bordered with brown on the inside."

Oliva tremulina forma olympiadina Duclos, 1835.

A handsome Indian Ocean form which is always white, some-

times obscurely reticulated with purplish brown, but always hav-

ing a more calloused white columella and white aperture. John-

son called this shell an albinistic form of O. tremulina.

Oliva tremulina forma oldi, new form. Fig. 2.

]ul>

969

NAUTILUS

17

Fig. 1. Two para types of Oliva incrassata Lightfoot forma burchorum Zeig-

ler. Sam Felipe, Baja California, Mexico. 35 and 41 mm. in length.

This is a distinctive and constantly marked shell, characteris-

tically grayish in overall appearance, though background color is

creamy yellow. Longitudinal charcoal zigzag lines usually crossed

by two solid or broken dark transverse bands. Somewhat similar

in pattern and markings to 0. textilina Lamarck, 1810, but longi-

tudinal lines are not so closely spaced as in that species. No match-

ing color form in the species O. miniacea have been observed. Base

of columella adorned with orange, above which is a heavy dark

brown fasciole. Length 50-56 mm. Not common. Type locality:

Bougainville Island, Solomon Islands. Additional specimens from

Luzon, Philippine Islands, and Mantis Island, Admiralty Group.

Range — longitudes 120° east — 158° east and latitudes 8° south

— 15° north.

The holotype is now in the American Museum of Natural His-

tory in New York City, No. 147750. There are four paratypes, two

of which are in the author's collection (the figured specimens) and

one each in the collections of John O. Burch, Seal Beach, Cali-

fornia and Walter G. Krause, Avoca Beach, N.S.W., Australia.

Many years ago, Dr. Howard Hill, former Curator of Marine

Zoology at the Los Angeles County Museum, sent several speci-

18

NAUTILUS

Vol. 83 (1)

Fig. 2. Two paratypes of Oliva tremulina Lamarck forma oldi Zeigler. Bou-

gainville Island, Solomon Islands. Both 54 mm. in length.

mens of this shell to the writer labelled "unidentified from Bou-

gainville." More of the same shells have since been obtained from

Bougainville, as well as from Manus Island, and one from the

Philippines.

Other specimens were found in the American Museum of Natu-

ral History collection, and William E. Old, Jr., supplied several

specimens, all of which came from Bougainville. The name oldi is

here proposed for this color form in honor of William E. Old, Jr.,

who has done much work and research on the genus Oliva, and

who has been of invaluable assistance.

Literature references

Burch, John Q. and Burch, Rose L. 1960. Catalogue of Recent

and Fossil Olives. Issue 196 of Minutes of the Conchological

Club of Southern California.

. 1960. Minutes of the Conchological Club of Southern

California. Issues 181-195.

1967. The Family Olividae. Pacific Science, Vol. 21, No. 4,

503-522.

Dautzenberg, Ph. 1927. Olivides de la Nouvelle Caledonie et de ses

dependences. J. Conchy. 71: 1-72, 103-147.

Duclos, P. L. 1835, 1840. Histoire naturelle . . . de tous les genres

de coquilles univalves marins a l'eta vivant et lossile, publice

par monographies. Genre Olive. Paris.

. 1844-48. Oliva, In: J. C. Chenu, Illustrations conchylio-

logiques. Paris.

July. 1969 nautilus 19

Hill. Howard R. 1954. Variation in the Olive Shells of Tropical

West America. Nautilus. Vol. 68(2): 66-69.

Johnson, Charles W. 1910-1911. Some notes on the Olividae.

Nautilus 24: 49-51, 64-68, 121-124.

. 1915. Further notes on the Olividae. Nautilus 28: 97-103,

114-116.

-. 192S. A review of certain species of the Olividae. Nautilus

42: 6-13.

Lamarck, J.B.P.A. de M. de. 1811. (Suite de la) Determination

des especes de mollusques testaces; continuation du genre Ovule,

Tarriere Ancillaire et Olive. Mus. Hist. Nat. (Paris) 16 (for

1810): pp. 89-114, 300-328 (Jan.-Mar., 1811).

Marrat, F. P. 1870-71. Oliva Bruguiere. In: G. B. Sowerby,

Thesaurus Conchyliorum. Vol. 4, pis. 328, 329-41, 342-51 (Oliva,

pis. 1-25).

Reeve, Lovell. 1850. Conchologia Iconica: or illustrations of the

shells of molluscous animals. Monograph of the genus Oliva.

pis. 1-30. London.

Tryon, G. W., Jr. 1883. Manual of Conchology. Ser. 1, Vol. 5.

Philadelphia.

GALAPAGAN RECORDS FOR MORUM VELEROAE

(GASTROPODA: TONNACEA)

By WILLIAM K. EMERSON

American Museum of Natural History

Morion (Cancellomorum) veleroae Emerson (1968b, p. 53, pi. 1,

figs. 1-7) was recently described by the writer on the basis of five

sub-adult specimens obtained by dredging, in 1938, off Cocos Is-

land, some 325 miles southwest of Costa Rica. Shortly after my

manuscript, describing this interesting species was accepted for

publication, Mesdames Carmen Angermeyer and Jacqueline DeRoy

each reported (in litteris) taking living specimens of this species

by dredging in the Galapagos Islands. Through the kindness of

these industrious collectors, I can now provide additional data re-

sulting from the Galapagan specimens.

Mrs. DeRoy dredged two living specimens in 100 meters near

Tagus Cove, Isabella Island, on January 28, 1968. One is a large

specimen measuring 52.2 mm. in length and 27.8 mm. in greatest

diameter. This specimen, here illustrated as figs. 1, 2, possesses a

very thin, immature, outer lip and it has not developed the parie-

tal callus. The small tonnacid operculum has a marginal nucleus

and concentric growth lines. A radula is wanting. The second

20

NAUTILUS

Vol. 83 (1)

Morum (Cajicellonwrmn) veleroae Emerson, 1968

Figs. I, 2: off Isabella Island [ — Albemarle Island], Galapagos Islands, in 100

meters, DeRoy coll., X 1. Note the lack of development of a parietal callus

and tbe immature outer lip; operculum is attached to a cotton plug in aper-

ture of specimen.

Figs. 3, 4: off James Island [ = San Salvador Island], Galapagos Islands, in 69

meters; ex-Angermeyer coll., A.M.N.H. No. 152606; X 1.3. Note the well-

developed parietal callus and mature outer lip.

Jul\, 1969 NAUTILUS 21

specimen, which measures approximately 42 mm. in length, also

possesses an immature outer lip.

Mrs. Angermeyer dredged a specimen in about 69 meters on

coral and sand bottom off the southern coast of James Island, on

June 14, 1968. She noted that the animal was similar to that of

Morum (Moram) tuberculosum Reeve, 1842, with the body a light

gray with black mottling. Her specimen here illustrated as figs. 3,

4, is approximately 38 mm. in length (nuclear whorls missing)

and is about 24.3 mm. in greatest diameter. The shell, which has

a well-developed parietal callus and an apparently mature outer

lip, is more vividly colored than the type specimens, which are

somewhat faded, as they were collected some thirty years ago. The

parietal callus of this smaller, but more mature specimen, was a

glossy raspberry-red before the shell was preserved in alcohol,

whereas that of the type material is a pinkish lavender. The

dorsum of the shell is overgrown with calcareous algae.

Mrs. Angermeyer (in litteris) also reports obtaining a well-pre-

served, dead specimen off northern James Island, in 69 to 80

meters, on coral and sand bottom, June 16, 1968. This specimen,

which is approximately 42 mm. in length and 25 mm. in width,

has longer and more prominent spines on the shoulder than those

possessed by the specimen she collected alive.

With the discovery of the four Galapagan specimens of Morum

(Cancellomorum) veleroae, a total of nine examples are now

known, including the five specimens of the type lot from Cocos

Island. The largest individual, measuring nearly 53 mm. in length,

is the DeRoy specimen from the Galapagos Islands (figs. 1, 2).

The smallest specimen is a juvenile, measuring approximately

22 mm. in length, of the type lot from Cocos Island (Emerson,

1968b, pi. 1, figs. 6, 7) . The specimens were dredged from depths

ranging from 57 to 91 meters off Cocos Island, and 69 to 100

meters in the Galapagos Islands, on sand, coral, and coralline sub-

strates. Living examples of its western Atlantic analogue, Morum

(Cancellomorum) dcnnisoni (Reeve, 1842) have been collected in

62 to 139 meters, with a medium depth of 110 meters, on coral,

rocky, and sponge substrates (Dance and Emerson, 1967, p. 94) .

Future dredging operations along the continental shelf of the west

American mainland, within the Panamic faunal province, should

22 nautilus Vol. 83 (1)

reveal the presence of Morum (Cancellomorum) vcleroae living at

moderate depths on appropriate substrates.

Six living species of Morum (sensu lato) are know to occur in

warm to tropical waters of the New World, four in the Western

Atlantic and two in the eastern Pacific (Emerson, 1968a) . The

only other West American Recent representative of this genus is

Morum (Morum) tuberculosum (Reeve, 1842), a shallow-water

species of the nominate subgenus. Reeve's taxon also is repre-

sented in the western Atlantic by an analogue, Morum (Morum)

oniscus (Linne, 1767) , the type species of Morum Roding, 1798.

Literature cited

Dance, S. P., and W. K. Emerson. 1967. Notes on Morum denni-

soni (Reeve) and related species. Veliger, 10(2) : 91-98, pi. 12.

Emerson, W. K.

1968a. The American species of the genus Morum. Ann. Repts.

Amer. Malacol. Union for 1967, pp. 74-75.

1968b. A new species of the gastropod genus Morum from the

eastern Pacific Ocean. Journ. Conchyl., Paris, 107 (1) : 53-57,

Pi- 1-

NORTHERN RANGE EXTENSION AND WINTER

MORTALITY OF RANGIA CUNEATA

By JOHN L. GALLAGHER and HARRY W. WELLS

Department of Biological Sciences

University of Delaware, Newark, Delaware

The brackish water mactrid clam, Rangia cuneata (Gray) , until

recently was believed to occur no farther north than Virginia (Ab-

bott, 1968) . This species occurs at Roanoke Island, North Caro-

lina, and Wass (1963) found it was abundant in the James River,

Virginia. Pfitzenmeyer and Drobeck (1964) have more recently

reported it from the Potomac River. Because this species is har-

vested for food from various estuaries along the Atlantic and Gulf

coasts, the extent of its northward migration is of special interest.

In October, 1967, four living specimens of Rangia cuneata were

dredged from the tidal waters of North East River, Maryland, at

the head of Chesapeake Bay (39° 32' N, 75° 59' W) . This collec-

tion was made in approximately six feet of water. A year later

(December 30, 1968) , thousands of shells of this species were found

washed onto the shore of the Elk River near Thackery Point (39°

28.5' N 7,5° 58.3' W) , about five miles south of the first collection

July, 1969

NAUTILUS

23

site. The latter location is along the waterway leading to the

Chesapeake and Delaware Canal which connects Chesapeake Bay

with Delaware Bay.

The probable cause of the mass mortality observed in Elk River

was low temperature. A similar mass mortality had been observed

here in the winter of 1966-67. Heavy ice conditions occur most

winters in the Elk River, occasionally requiring ice breaker as-

sistance for ships en route to and from the Chesapeake and Dela-

ware Canal. At the time the dead Rangia were observed in 1968,

ice lined the shore although the deep water channel was free of ice.

The victims of this mortality probably came from the shallows

where the stress of low temperature would be greatest. Under these

physical conditions, clams in deep water could survive and main-

tain the population in this area.

Salinity conditions within the Elk River (0.3% in January,

1969) are an unlikely alternative cause of death, because Rangia

cuneata has been reported living in the Potomac River in a simi-

lar salinity regime. However, low salinity may have increased

stresses upon clams at low temperatures. Pollution is considered

unlikely as a cause of the mortality because no other species was

10

20 30 40

SHELL LENGTH (MM)

Fig. 1. Length distribution of Rangia cuneata from Elk River, Maryland,

December 30, 1968.

24 NAUTILUS Vol. 83 (1)

found to have succumbed. As a crude assay, water samples taken

at the time of collection were found to support developing frog

embryos and tadpoles.

From a representative sample of 244 specimens collected from

a 200-yard section of the beach, the length-frequency distribution

shown in Fig. 1 was constructed. Shell lengths ranged from 1 1 to

46 mm., with a mean of 26 mm. and a standard deviation of 5.2

mm. In comparison with data presented by Pfitzenmeyer and

Drobeck (1964) for clams from the Potomac River, the shells from

Elk River have a greater average length. At present, however, it is

not known if this mortality were selective, or if the dead clams

were indeed representative of the total living population in that

river. Lengths reported by Wolf and Petteway (1968) for Rangia

cuneata at different ages, and observations of growth rings on these

shells indicate that the Elk River mortality involved individuals

ranging from one to four years old. In most cases, the flesh of the

clams was badly decomposed at the time of collection.

Pfitzenmeyer and Drobeck (1964) suggested that Rangia cune-

ata may have been introduced to the Potomac River with seed

oysters. Because the oyster bars closest to the Elk River are approxi-

mately 30 miles away in Chesapeake Bay, it is conceivable that this

clam has spread northward from the oyster growing area. Pfitzen-

meyer and Drobeck considered the possibility that, after many

years' presence elsewhere in the Potomac River drainage system,

this species had expanded its distribution to areas where it at-

tracted biologists' attention. However, they discounted this possi-

bility since no sudden environmental changes had been noted

which could be correlated with the clams' discovery. In the absence

of marked recent changes in the local environment, their conclu-

sion seems applicable to the Elk River situation. If future surveys

show this population to be relatively isolated, it would increase

the probability of man's introduction of Rangia cuneata into the

immediate area, possibly in waste from dredge gear or spoil barges.

On the other hand, winter kills in the Elk River suggest that

this species is approaching its natural northern geographic limit

under present climatic conditions. Richards (1938) has reported

fossil Rangia cuneata in New Jersey, indicating former Pleistocene

extension into that state, but he could find no living specimens.

The sea-level canal connecting Chesapeake Bay to Delaware Bay

July, 1969 nautilus 25

presents an opportunity for the transport of larvae and reproduc-

tive stages of aquatic organisms tolerant to brackish waters. The

larvae of Rangia cuneata, described by Chanley (1965) , can read-

ily be transported in this fashion. The proximity of the canal to

the Elk River collection site makes it probable that Rangia cune-

ata will be found living in brackish waters of the Delaware Bay

system, paralleling the record of its Pleistocene distribution in

that area.

Specimens have been deposited in the United States National

Museum, Division of Mollusks.

Literature cited

Abbott, R. T. 1958. Seashells of North America. Golden Press,

New York. 280 pp.

Chanley, P. E. 1965. Larval development of the brackish water

mactrid clam, Rangia cuneata. Chesapeake Sci. 6 (4) : 209-213.

Pfitzenmeyer, H. T., and K. G. Drobeck. 1964. The occurrence of

the brackish water clam, Rangia cuneata, in the Potomac River,

Maryland. Chesapeake Sci. 5(4): 209-212.

Richards, H. G. 1938. Animals of the seashore. Bruce Humphries,

Inc., Boston. 273 pp.

Wass, M. L. 1963. Check list of the marine invertebrates of Vir-

ginia. Va. Inst, of Marine Science, Spec. Sci. Rept. 24. 56 pp.

Wolfe, D. A., and E. N. Petteway 1968. Growth of Rangia cuneata

Gray. Chesapeake Sci. 9(2): 99-102.

26 NAUTILUS Vol. 83 (1)

A NEW CAECUM FROM THE TROPICAL

WESTERN ATLANTIC1

By DONALD R. MOORE

Institute of Marine Sciences, University of Miami

In the summer of 1968 I was asked by George Rachvin, then a

predoctoral fellow at the U. S. National Museum, to check his

identification of some marine micromollusks. The material came

from Payardi Island on the Atlantic side of Panama. The island

is adjacent to the city of Colon, in the Canal Zone.

An unusal specimen of the family Caecidae was included in

the material. As it was a single specimen, it was put aside. A few

weeks later, I was invited by Helmer Odd and Harold Geis to look

over the Caecidae collected in their Texas off-shore survey. There

was a wealth of material and again a single specimen of the un-

usual Caecum. Since the two specimens were clearly conspecific,

it was decided to describe the species.

Caecum condylam, new species

Description: The teleoconch of C. condylum is cylindrical and

only slightly curved for most of its length. The anterior part near

the aperture is more strongly curved. Near the aperture there is a

broad rounded swelling or varix. Shortly beyond this the shell

terminates at the aperture.

The septum is slightly depressed and the low, somewhat pointed

mucro is angled to the right. The aperture is circular and is about

the same diameter as the shell on the proximal side of the varix.

The ground color is white with several diffuse bands of light

brown.

The sculpture consists of approximately 100 low close set annu-

lar ridges. They continue over the varix on to the aperture.

The ridges do not increase in size on the varix. The protoconch

and second stage are unknown. The operculum was not observed.

Material. Holotype. Payardi Island, near Colon, on the Atlantic

side of Panama. Length, 2.40 mm. Division of Mollusks, U. S. Na-

tional Museum, no. 679348. Paratype. A single specimen from

27° 54' 30" N., 93° 35' 45" W., East Flower Garden Bank, Gulf of

Mexico, coral debris, 18.3-23.8 m., length, 3.36 mm. Division of

Mollusks, U. S. National Museum, no. 679349.

l Contribution No. 1033. from The Institute of Marine Sciences, University

of Miami, Miami, Florida.

July, 1969 NAUTILUS 27

Fig. I. Caecum condylum new species. The holotype as seen from the right

side. There is a slight break in the aperture.

Name. From condylus, the enlarged end of a bone.

Remarks. There is a considerable difference in size in the only

two specimens available. They are contrasted below.

The chief difference in the two specimens is size. In all other re-

spects they are closely similar. Both appear to be fully mature.

I have examined thousands of specimens of Caecidae from the

western Atlantic. In all that material, it seems strange that there

would be only two specimens of the new species from localities

some 1500 miles apart. A possible explanation might be that it

occupies a restricted habitat, and that this habitat has been little

sampled for micromollusks.

C. condylum does not appear to be very closely allied to any

western Atlantic species. C. subvolutum Folin and C. veracruzanum

Folin are similar in shape, but they do not have the axial ridges.

Those species with axial ridges, such as C. pulchellum Stimpson,

have them much larger and fewer in number. C. clava Folin has

an even larger terminal varix and longitudinal sculpture. Even

immature specimens of C. condylum should be fairly easy to

identify.

At first glance it would seem that this species should go into the

genus Micranellum Bartsch 1920. The type of the genus is Caecum

crebricinctum Carpenter 1864. In 1866, Carpenter noted that C.

crebricinctum had the aspect, but not the sculpture of an Elephan-

tulum. In this latter genus, the sculpture supposedly consists of

longitudinal ridges only. Unfortunately, there are the two ex-

28 NAUTILUS Vol. 83 (1)

tremes of sculpture development, and, as more species are exa-

mined, it is seen that there is a gradual transition from one type

to another.

A species being described from the Virgin Islands as Caecum

insularum (Moore, in press) is very similar in appearance to

C. crebricinctum. Chief differences are fewer axial ridges and the

presence of very weak longitudinal ridges in C. insularum. This

species is in turn very close to C. imbricatum Carpenter, a species

in which axial and longitudinal sculpture is nearly equal.

C. crebricinctum is also quite similar to C. imperforatum Kan-

macher, the type species of Caecum. Even the coloration is much

the same and the principle differences are size and number of

axial ridges. The entire family seems quite homogenous, and all

subdivisions should probably be at the subgeneric level. It may

not be possible to separate Caecum s.s. from Micranellum. Bartsch

(1920) mentions that five of the species he placed in Micranellum

have bulbous swellings at the end of the teleoconch. This would

seem to justify the establishment of a new subgenus based on

C. condylum and the possibly related Pacific coast species, but

Bartsch did not really describe any of his species nor did he figure

any. Until the Pacific coast material can be examined carefully,

it is best to refer the new species herein described to the genus

Caecum without increasing the list of supraspecific names.

Acknowledgements

I am grateful to the Division of Mollusks, U. S. National Mu-

seum, for the opportunity to describe the species. I also wish to

thank Harold Geis and Helmer Ode for contributing the paratype

from Texas. This work was supported by National Science Foun-

dation Grant GB-5055 (UM N8753) . The excellent drawing of

the holotype was made by Mrs. Barbara Lidz.

Literature cited

Bartsch, P. 1920. The Caecidae and other marine mollusks from

the northwest coast of America. J. Wash. Sci., 10 (20) : 565-572.

Carpenter, P. P. 1864. Supplementary report on the present state

of our knowledge with regard to the Mollusca of the west coast

of North America. Rept. British Assoc. Adv. Sci. 1863: 517-686.

. 1866. Descriptions of new marine shells from the coast of

California. Pt. III. Proc. Calif. Acad. Sci., 3: 207-224.

Moore, D. R. (in press) A new Caecum from Puerto Rico and the

Virgin Islands. Bull. Mar. Sci.

July, 1969 nautilus 29

NOTES ON THE BIOLOGY OF THE SNAIL

SONORELLA ODORATA IN ARIZONA1

By LANCE H. GILBER TSON

Orange Coast College, Costa Mesa, California

Most species of Sonorella inhabit rock slides of the arid and

semi-arid slopes of the Lower and Upper Sonoran Life Zones in

Arizona, New Mexico and Sonora, Mexico. Sonorella odorata

Pilsbry & Ferriss is exceptional ecologically, being a forest snail of

the mesic Canadian Life Zone (usually between 7,500 and 9,000

feet) . It is definitely known only from the Santa Catalina Moun-

tains near Tucson, Arizona. Its supposed occurrence in the nearby

Rincon Mountains has not been confirmed in recent years (Miller,

1967).

Pilsbry 's name "Sonorella odorata" was based on populations

which were later segregated as the nominate subspecies, viz., Son-

orella odorata odorata. Another subspecies 5. o. marmoris inhabits

the upper talus slopes of Marble Peak in the same mountain range.

"S. odorata" will refer here to the nominate subspecies only.

The present study was conducted at Bear Wallow, located near

the north-facing slope of Mt. Lemmon at approximately 8,000 feet.

This is near the type locality of 5. odorata, "at the head of Alder

Canyon." The forest floor here is generally shaded, damp, and

deep with decaying plant litter.

When disturbed, 5. odorata emits an unpleasant, mephitic odor

which accounts for its specific trivial name. However, it was never

observed ejecting drops of odoriferous fluid as Ferriss reported.

Feeding habits — S. odorata is nocturnal, and only under arti-

ficial conditions has it been observed feeding during daylight

hours. Therefore, to investigate its normal feeding habits under

natural conditions, several trips were made at night to Bear

Wallow. The results of these field observations and subsequent

laboratory analyses of digestive tract contents indicate that the

principal food items are: (1) decaying wood, bark, needles and

forest litter in general, and (2) lichens, especially the plentiful

foliose Usnea sp. as well as the more crustose forms. In addition,

the snail has been observed feeding on a large red-capped mush-

l This paper is an abridgment of a portion of the biological data in a thesis

submitted for the M.S. degree in Zoology at the University of Arizona,

Tucson, in January 1965.

30 nautilus Vol. 83 (1)

room, a rotting mushroom, a fallen green leal (apparently of a

columbine) and debris of a Douglas Fir cone. A specimen in the

laboratory was observed consuming its epiphragm upon emerging

from estivation.

Mating — Mating occurs only during summer months in the

field, but throughout the year in the laboratory. In both places it

was observed primarily at night, but on occasion also during day-

light hours. Copulation is reciprocal and simultaneous, and has

been timed to last from 2 min. 45 sec. to 9 min. No special pre-

ovulatory behavior was observed. Mating individuals were ob-

served in Bear Wallow on the night of September 21, 1964, very

near the end of their active sesaon.

Oviposition and Development — Egg laying has been observed

only after drenching summer rains. It was observed by the author

on August 4, 6, and 12, 1964. The snail burrows with the anterior

portion of its body about 1.5 cm. below the surface of the soil.

The eggs are then laid in spiral fashion with each layer consisting

of six to seven eggs surrounding a central one. The amount of time

needed for oviposition was not determined; however, one individ-

ual was observed in the typical egg-laying position for almost 5

hours. The length of time between copulation and oviposition

remains uncertain, due in part to the reluctance of S. odorata to

lay eggs in the laboratory. In one instance, however, it apparently

was about 45 days.

The incubation period in the laboratory at 75° F ranged from

15 to 20 days. Under natural conditions, however, incubation pos-

sibly lasts longer; in one observed cluster the eggs showed no signs

of advanced development after 17 days. When the embryonic snail

is fairly well developed, it rotates incessantly within the egg mem-

brane. Many structures are easily visible at this time, including

the shell, body proper, tentacles with eye-spots and even the beat-

ing heart. After several hours of rasping at the egg membrane, the

young snail pulls free from the egg and accompanying mucus. The

new-born snail weighs just slightly under 0.01 gm. Its shell has

already acquired the characteristic brown color and is covered

with fine, hair-like projections.

Hibernation — S. odorata spends most of its life in hiberna-

tion. Only during the summer rainy season does it emerge to feed

and reproduce. It is found hibernating under large branches, logs,

July, 1969

NAUTILUS

31

amongst debris of wood and bark, under rocks, in rock crevices,

in the soil, and perhaps most often under large flattened pieces of

bark. Typically, the snail lies with the aperture facing upwards.

The aperture is closed with a brilliant orange muco-calcereous epi-

phragm; during the course of hibernation, additional, much lighter

and thinner epiphragms are formed internally. One snail, found

on November 4, 1964, had already formed two such additional

epiphragms.

5. odorata often hibernates gregariously. For example, in March

1963, Dr. Walter B. Miller uncovered a "pocket" of some 20 in-

dividuals hibernating in close proximity in the soil. On another

occasion, the author discovered nine hibernating S. odorata under

a large, flat piece of bark within 10 cm. of each other. However,

many have been found hibernating alone.

Near the end of hibernation, individuals weigh less than active

ones, but more than those which had estivated in the laboratory

(at 75° F) for an equivalent length of time (Fig. 1) .

2.00

oo

o

o

UJ

<

1.60-

1.20-

0.80-

0.40

0

18

21

SHELL DIAMETER IN MM.

Fig. 1. Comparative weights of S. odorata in (1) the active state, (2) after

ca. 8 months of hibernation, and (3) after ca. 8 months of estivation.

32 nautilus Vol. 83 (1)

Estivation — Estivation occurs rather commonly in natural as

well as laboratory based populations. In both situations, a few

individuals usually could be found estivating even when environ-

mental conditions seemed ideal for active life. This phenomenon,

therefore, may be cyclic or periodic in nature as suggested in the

research of Howes and Wells (1934) .

Estivating S. odorata, unlike those in hibernation, are usually

sealed to some object by their calcareous mucus. This mucus is

never the brilliant orange color characteristic of the primary epi-

phragm secreted by hibernating individuals; rather, it is much

lighter and resembles the more internal secondary epiphragms.

The snail is able to withstand prolonged periods of estivation; but

this capability is not essential for survival under prevailing en-

vironmental conditions. Of 13 individuals, 7 survived 8 months

of estivation in the laboratory (at 75° F and low humidity) .

Mature individuals lost from 39 to 59 percent of their body weight

(less shell) . However, one small, immature snail lost only 10 per-

cent of its body weight.

Association with fungi — The role of land pulmonates in the

germination and dissemination of fungal spores has been com-

mented on by several investigators. The present study represents

an effort to define those types of fungi, found in the digestive tracts

of 5. odorata, which were able to grow on artificial nutrient media.

Five laboratory based specimens and six specimens collected at

Bear Wallow on August 4, 12, and 18, and September 16, 18 and

21, 1964 were used. The entire digestive tracts were removed,

rinsed in sterile water and triturated with a sterile mortar and

pestle. Petri dishes containing Malt-Yeast Extract Agar with a pH

of 4.0 were then inoculated and incubated at room temperature.

As colonies developed, they were transferred to test tubes with

slants of Malt- Yeast Extract Agar for subsequent identification.

Several species of fungi were isolated from both groups. Those

isolated from the digested tracts of laboratory based 5. odorata

were: three Mucor Mich, ex Fr. spp. including Mucor raman-

nianus Moeller; three Mortierella Coemans spp.; two Penicillium

Link ex Fr. spp.; Syncephalastrum Schroet. sp.; and Trichoderma

lignorum Harz. Fungi isolated from snails collected in their natural

habitat were: three Trichoderma Pers. ex Fr. spp., including

T. album Preuss and T. koningi Oudemans; four Mucor Mich.

July, 1969 nautilus 33

ex Fr. spp., including M. ramannianus; six Penicillium Link ex

Fr. spp. including the strongly fasciculate P. clavigerum Demelius;

Pulhdaria Berkh. sp.; Diplodia Fries sp., and Cephalosporium

Corda sp.; Mucor ramannianus and one other Mucor sp. were

the only two species common to both groups. M. ramannianus

showed up in every laboratory based snail and in three of the six

snails collected in their natural habitat. This was particularly

significant, since it had not been previously isolated from this

area despite extensive soil sampling by Dr. Adelaide Evenson

Riker. A third Mucor sp., characterized by a yellowing of the

mycelium with age, was isolated from the digestive tracts of all

six specimens taken from the natural habitat, but never showed

up in the laboratory based snails. This suggests either a definite

relationship between S. odorata and this fungus, or an ubiquitous-

ness of its spores. The regular isolation of at least three Mortierella

spp. from the laboratory based group also seems significant, since

species of this genus are not commonly obtained from soil samples.

Unfortunately, however, they were not isolated from any of the

six specimens comprising the natural habitat group. The genera

Penicillium and Trichoderma were well represented in both

groups, but by differing species. T. album and T. koningi were

both isolated twice from different snails in the natural habitat

group. No myxomycete growth was observed.

I wish to thank Drs. Albert R. Mead and Joseph C. Bequaert

for their guidance and encouragement throughout the course of

this investigation and the preparation of this paper. I also wish to

express my indebtedness to Dr. Adelaide Evenson Riker of the

Department of Microbiology, University of Arizona, who directed

and assisted me in all phases of the mycological studies, and made

all of the identifications.

Literature cited

Howes, N. H. and G. P. Wells. 1934. The water relations of snails

and slugs. I. Weight rhythms in Helix pomatia L. II. Weight

rhythms in Arion ater L. and Limax flavus L. Jour. Exp. Biol.,

11: 327-351.

Miller, Walter B. 1967. Anatomical revision of the genus Sonorella

(Pulmonata: Helminthoglyptidae) , 293 p. (Ph.D. dissertation,

Univ. of Arizona, Tucson) .

Miller, M. C. 1961. Distribution and food of the nudibranchiate

34 nautilus Vol. 83 (1)

mollusca of the South of the Isle of Man. J. Anim. Ecol.

30: 95-116.

Moore, George M. 1968. Personal Communication, University of

New Hampshire, Durham, N. H.

Odhner, Nils Hj. 1929. Aeolidiiden aus dem nordlichen Nor-

wegen. Tromso Museums Arsh. 1: 1-22.

Odhner, Nils Hj. 1939. Opisthobranchiate mollusca from the

western and northern Coasts of Norway. Norske Vid. Selsk. Skr.

1: 1-93.

Sars, G. O. 1878. Bidrag til kundskaben om Norges Arktiske Fauna.

I. Mollusca Regionis Arcticae Norvegiae. Christiania.

Swennen, C. 1961. Data on distribution, reproduction and ecology

of the nudibranchiate molluscs occurring in the Netherlands.

Neth. J. Sea Res. 1: 191-240.

Thompson, T. E. 1964. Grazing and the life cycles of British

nudibranchs. Brit. Ecol. Symp. 4: 275-297.

FURTHER ADDITIONS TO THE UNIONID FAUNA

OF THE GULF DRAINAGE OF ALABAMA,

GEORGIA AND FLORIDA

By RICHARD I. JOHNSON

Museum of Comparative Zoology

In 1967 and 1968, I reported on the additions to the unionid

fauna of the Apalachicolan region which have been made since

the study of Clench and Turner (1956) . Recently, through the

kindness of Dr. William H. Heard of Florida State University,

Tallahassee, who sent some specimens to the Museum of Compara-

tive Zoology, I am able to report on two more species previously

unknown from the Apalachicolan region, and on the extension of

the ranges of two others.

Anodonta suborbiculata Say is widely distributed throughout

the Interior Basin, extending south to Louisiana. It occurs in the

Escambia River system at Gantt Lake, an empoundment on the

Conecuh River, Clearview, Covington Co., Alabama. Although I

am not aware of any records of this species in the neighboring

Alabama-Coosa River system, it would be surprising if it does not

occur there.

Anodonta couperiana Lea was previously thought to be limited

to the Southern Atlantic Slope and Peninsular Florida regions

(Johnson, 1965, pi. 1) . It occurs in the neighboring Apalachicola

and Ocklockonee river systems. In the former, at Apalachicola

July, 1909 nautilus 35

River, Ocheesee Landing, 6 miles North of Blountstown, Calhoun

Co., Florida and in the latter at Ocklockonee River, 1 1 miles

North of Tallahassee, Leon Co., Florida. The discontinuous dis-

tribution of this species like a number of others, may be explained

by a former confluence of the headwaters of the Apalachicola and

Savannah river systems.

Fusconaia escambia Clench and Turner was known only from

the Escambia River system when it was described. It also occurs in

the Yellow River system, at Yellow River, 4 miles West of Crest-

view, Okaloosa Co., Florida.

Obovaria rotulata (B. H. Wright) was formerly known only

from the type specimen from the Escambia River system. It has

since been found there, in the Escambia River, Molino, Escambia

Co., Florida and in the Conecuh River, 1 mile South of East Brew-

ton, Escambia Co., Alabama.

Literature cited

Clench, William J. and Ruth D. Turner. 1956. Freshwater mol-

lusks of Alabama, Georgia, and Florida, from the Escambia to

the Suwannee River. Bull. Florida State Mus., 1: 97-239, pis. 1-9.

Johnson, Richard I. 1965. A hitherto overlooked Anodonta (Mol-

lusca: Unionidae) from the Gulf drainage of Florida. Breviora,

Mus. Comp. Zool., No. 213: 1-7, pis. 1-2.

. 1967. Additions to the unionid fauna of the Gulf drainage

of Alabama, Georgia and Florida. Breviora, Mus. Comp. Zool.,

No. 270: 1-21, pis. 1-3.

1968. Elliptio nigella, overlooked unionid from Apalachicola

river system. Nautilus, 82 (2) : 22-24, plate.

NOTES AND NEWS

Dates of the Nautilus. — Vol. 82, no. 1, pp. 1-36, was mailed

July 24, 1968. No. 2, pp. 37-76 on Oct. 28, 1968. No. 3, pp. 77-114

on Jan. 31, 1969. No. 4, pp. 115-148 on April 28, 1969. — R. T.

Abbott.

Ecological Distribution of the Non-marine Gastropods of the

Olympic Peninsula, Washington, is the subject of a Sigma Xi 1969

grant recently made to Dr. Branley A. Branson of Eastern Ken-

tucky University.

36 NAUTILUS Vol. 83 (1)

Concerning rawson's physa. — Bland and Binney, in 1873

(Ann. Lye. Nat. Hist. N. Y., 10: 255-257) , described and figured

the lingual dentition of a species of Physa from Guadeloupe. They

obtained the specimen from Governor Rawson of Guadeloupe

who had earlier received it from the naturalist H. Schramm. The

snail was thought to have been collected in the vicinity of Point-a-

Pitre and was unusual in that the jaw and radula more nearly

resembled those of the Lymnaediae and Planorbidae than of the

Physidae. The shell, however, was typically physoid.

Bequaert and Clench (J. Conch., 21: 175-178) compared Bland

and Binney 's figures with material from Plesiophysa ornata (Haas),

and concluded that Rawson's Physa was a member of the genus

Plesiophysa; they placed the species in synonymy with Plesiophysa

guadeloupensis (Maze) .

In examining a series of old slides deposited in the Museum of

Comparative Zoology at Harvard University, I found one labeled

"Rawson's Physa" and signed W. G. Binney. The date and journal

citation of Bland and Binney's article also appeared on the label.

Although the slide was badly broken and the mounting medium

dried and cracked, details of the radula and jaw were observable.

The rachidian tooth had 6 cusps, typical of the genus Plesiophysa .

The marginals were not as figured by the original authors, but

were morphologically similar to those of P. ornata as figured by

Bequaert and Clench (loc. cit.) and by Hubendick (Arkiv. f.

Zool., 42: 1-10) . There were marked vertical striations on the

dorsal portion of the jaw, and chitinous material suggestive of

lateral processes were noted. These two characters are absent in

P. ornata, but have been observed in Puerto Rican P. hubendicki

by Richards and Ferguson (Trans. Am. Microscop. Soc, 81: 251-

256) . There appears to be little doubt that Rawson's Physa was

a species of Plesiophysa, possibly P. hubendicki. Until the mor-

phology of P. guadeloupensis is known, however, it is uncertain

whether P. guadeloupensis and P. hubendicki are the same or not.

Thus, the specific identity of Rawson's Physa remains question-

able. — Edward H. Michelson, Dept. Trop. Public Health, Har-

vard School of Public Health, Boston. (Under Training Grant

Al-00046 from the U. S. Public Health Service.)

July, 1969 nautilus iii

R. Tucker Abbott, who has been acting editor of the Nautilus

during 1968, has resigned from the Academy of Natural Sciences

of Philadelphia to join the newly founded Delaware Museum of

Natural History, Greenville, Delaware 19807. He will fill the

du Pont Chair of Malacology and also serve as an Assistant Direc-

tor of the museum. Indo-Pacific Mollusca is now being published

by the Delaware Museum of Natural History, and Dr. Abbott

will continue to serve as editor of that journal and the Nautilus.

Dr. H. B. Baker will continue to review the non-marine papers

submitted to the Nautilus. From July 1 to December 31, 1969,

Dr. Abbott will spend part of his time revising American Seashells

at the Academy of Natural Sciences, 19th and Parkway, Phila-

delphia, Pa. 19103. Suggestions and corrections to American Sea-

shells are welcomed, and Nautilus manuscripts may be sent either

to Dr. Abbott in Philadelphia or to Dr. Baker in Havertown.

George Mitchell Moore. 1906-1968. — Professor George

Mitchell Moore died at his home in Durham, New Hampshire, on

May 19, 1968. Born in Shelby County, Ohio, August 5, 1906, he

received the B.Sc. (1928) from Otterbein College, M.S. (1932)

and Ph.D. (1938; Zoology, Limnology) from the University of

Michigan. He taught in Ohio and Minnesota and was assistant

Professor at Bowling Green State from 1938-42. He conducted re-

search on the biology of Volsella demissus and Crassostrea virginica

as an associate biologist for the Virginia Fisheries Laboratory, Col-

lege of William and Mary in 1942-44. In 1944 he became associate

professor at the University of New Hampshire, was made full pro-

fessor in 1949, and served as Chairman of the Department of

Zoology from 1947-63. While at UNH, Dr. Moore developed grad-

uate and undergraduate courses in littoral marine invertebrate

zoology, revitalized interest in estuarine research and was instru-

mental in establishing a Ph.D. curriculum in zoology. During

1958-59 he was Fullbright Lecturer at Chulalongkorn, Bangkok,

Thailand, where he studied marine mollusks. The object of his

research in recent years was the nudibranchs of New England and

his comprehensive collection of these shell-less opisthobranchs has

been placed in the Division of Mollusks, U. S. National Museum,

Smithsonian Institution, Washington D. C. — Joseph Rosewater.

iv nautilus Vol. 83 (1)

PUBLICATIONS RECEIVED

McMillan, Nora F. 1968. British Shells, xii + 196 pp., 80 pis.

(24 in color) . 50 shillings ($6.00) . Frederick Warne and Co.,

Ltd., London. An excellent handbook covering the marine, land

and fresh-water mollusks of the British Isles.

Grasse, Pierre-P. (editor) . 1968. Mollusques Gasteropodes Scapho-

podes, vol. 5, fascicule 3 of the Traits de Zoologie — Anatomie

Systematique, Biologic 1083 pp., 517 text figs., 1 color plate.

Masson et Cie., 120 Boulevard Saint-Germain, Paris 6, France.

About $58.00. An extensive coverage of the prosobranchs and

opisthobranchs (by Andre Franc) ; evolution and paleontology

of the gastropods (by G. Termier and H. Termier) ; neurose-

cretion and endocrines in mollusks (by M. Martoja) ; and the

scaphopods (by E. Fischer-Piette and A. Franc) . The nomen-

clature of the prosobranch taxa is, in some parts, out of date.

Thompson ,Fred G. 1969. The Aquatic Snails of the Family Hy-

drobiidae of Peninsular Florida. Univ. of Florida Press, Gaines-

ville, $8.00, 268 pp., 69 pages of excellent figures of shells, oper-

cula, radulae, verges, soft parts and distributional maps of 36

Florida hydrobiids, including 23 new species and the new genera

Heleobops, Onobops, Hyalopyrgus, Aphaostracon and Spilo-

chlamys.

Purchon, R. D. 1968. The Biology of the Mollusca. Pergamon

Press, London and N. Y., $27.00, 560 pp., 185 figs. Extensive

and well-illustrated chapters on the Mantle Cavity, Feeding

Methods, Adaptive Radiation, Digestion, Reproduction, Dis-

tribution, and reviews of the seven classes (includes Aplaco-

phora) of Mollusca.

Boss, Kenneth J. 1969. The Genus Strigilla; The Subfamily Tellin-

inae in the Western Atlantic. Johnsonia, vol. 4, no. 47, pp. 345-

366, pis. 164-171. Excellent coverage of the six known species,

including pscudocarnaria Boss, n. sp.

Johns, Veronica P. She Sells Seashells. 198 pp. Plates on front

and back covers. Funk & Wagnalls division of Reader's Digest

Books. $5.95. A charming and witty account of Mrs. Johns'

experiences in her New York shell shop. Delightful reading —

B.B.B.

WILLIAM H. WEEKS SHELL COLLECTION: New price lists

of this famous collection, with full scientific data, are in prepa-

ration. Many new additions of fine and rare species are also

included. To obtain free copies write:

George E. Jacobs, 853 Riverside Drive, New York 32, N. Y.

Vol. 83 OCTOBER, 1969 No. 2

THE

NAUTILUS

THE PILSBRY QUARTERLY

DEVOTED TO THE INTERESTS OF CONCHOLOGISTS

EDITORS AND PUBLISHERS

Horace Burrincton Baker, 11 Chelten Road, Havertown, Pa.

(Emeritus Professor of Zoology, University of Pennsylvania)

R. Tucker Abbott, duPont Chair of Malacology

Delaware Museum of Natural History, Greenville, Del. 19807

Charles B. Wurtz, Biology Department

La Salle College, Philadelphia, Pa. 19141

CONTENTS

On the feeding of the nudibranch, Coryphella verrucosa

ruftbranchialis, widi a discussion of its taxonomy. By M.

Patricia Morse 37

A new northern record for Bursatella leachii pleii Rang

(Opisthobranchia) , with notes on its biology. By W. L.

Kruczynski and Hugh J. Porter 40

Succinea bakeri Hubricht. By Leslie Hubricht 42

Nonmarine mollusks of the Katalla region, Alaska. By S. J.

Tuthill and Robert L. Johnson 44

Illustrations of Lamarck's types of North American Unionidae,

mostly in the Paris Museum. By Richard I. Johnson 52

Nomenclatorial note: Arcticidae versus Cyprinidae (Mollusca;

Bivalvia) . By Kenneth J. Boss 61

Marine mollusks of Port Moller Bay, Alaska Peninsula. By

James X. Corgan 65

Notes and news 66 Publications received 75

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Mrs. Horace B. Baker, Business Manager

11 Chelten Road, Havertown, Pennsylvania 19083

Second-Class Postage paid at Spring House, Pa.

NAUTILUS:

A Quarterly Journal devoted to the study of Mollusks, edited and published

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Matter for publication should be sent to the acting editor, Dr. R. Tucker

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THE NAUTILUS

Vol. 83 October, 1969 No. 2

ON THE FEEDING OF THE NUDIBRANCH, CORYPHELLA

VERRUCOSA RUFIBRANCHIALIS, WITH A DISCUSSION

OF ITS TAXONOMY

By M. PATRICIA MORSE

Northeastern University, Marine Science Institute, Nahant, Massachusetts

The nudibranch mollusk, Coryphella verrucosa rufibranchialis

(Johnston 1832) as defined by Odhner 1939, has been observed

to feed upon the ascidian, Botryllus schlosseri (Pallas 1766) . The

observations and experimentation on this prey-predator relation-

ship reported in this paper were made at the Marine Science In-

stitute, Nahant, Massachusetts, during January, February and

March, 1968.

Coryphella verrucosa rufibranchialis has been reported to feed

on hydroids by Miller (1961), Swennen (1961) and Thompson

(1964) . In the investigations of these authors, there is no record

of any member of the family Coryphellidae feeding on an ascidian.

Swennen (I.e.) suggests a definite list of criteria for the acceptance

of a food-organism as important to the nudibranch. The criteria

are: 1) association of the nudibranch with the food-organism in

the field, 2) observation of the nudibranch feeding in the labor-

atory or in the field and 3) subsistence on the food-organism alone

for some period of time.

The nudibranchs were first seen feeding on the ascidian on the

sides of the 12' by 4' laboratory floor tanks and observations over

several months revealed small and large specimens of C. v. rufi-

branchialis associated with the colonies of ascidians. It was noted

that the cerata of the feeding nudibranchs all appeared chocolate

brown in color. In order to observe actual feeding and color

change of the cerata, Coryphella verrucosa rufibranchialis were

collected from the Nahant rocky shore and placed in 10" finger

bowls with the food organism, B. schlosseri. Growths of the ascid-

ian on Ascophyllum nodosum were utilized for the experiments.

Observations of the actual feeding process were made under a

dissecting microscope.

37

38 nautilus Vol. 83 (2)

The nudibranchs approached the food-organism and used the

sensory oral tentacles for positioning the mouth opening. Utilizing

the radula, the mollusk grazed on individual zooids of the star-

shaped cluster, scraping away the tunic and ingesting the softer

portions of the ascidian. Food was seen passing through the

esophagus into the stomach. Contents of the stomach and digestive

glands (which extend into the cerata) became dark chocolate-

brown in color, approximating the color of the prey, and the fecal

pellets were dark red-brown in color. (It is interesting to note that

Dendronotus frondosus (Ascanius 1774) has also been observed

feeding on B. schlosseri and ingests the tunic, individual zooids

and common matrix of the prey with no color changes in the

nudibranch.) Nudibranchs used in the experiments showed varia-

tion in the color of their cerata from a bright red to a darker red-

brown. In the case of those with bright red cerata, the color pro-

gressively turned to dark red-brown as the nudibranchs continued

to feed on B. schlosseri. Animals maintained on the ascidian for

three weeks copulated and deposited numerous strings of eggs.

Observations on C. v. rufibranchialis in January 1969 showed

that when animals with bright red cerata were placed in a finger

bowl with Syncoryne mirabilis Agassiz 1862, the cerata turned a

salmon-red color which approximated the color of the hydroid.

Discussion

In 1909 Balch recognized two varieties of Coryphella rufi-

branchialis on the basis of distinct color variations among nudi-

branchs collected at Gloucester, Massachusetts. He assigned the

names C. rufibranchialis mananensis (Stimp.) and C. rufibranch-

ialis chocolata var. nov. The late Dr. George M. Moore (personal

communication) collected the variety (C. r. chocolata) during the

month of April in Rye, New Hampshire and Kittery, Maine. He

compared the radulae of the two varieties and did not find any

differences. An examination of his radular preparations verifies

this. Balch (I.e.) postulated that C. r. chocolata might be "a mere

physiological phase" and the evidence herein bears that out.

It must be noted that all specimens of Coryphella verrucosa

(M. Sars, 1829) recorded from New England have been the vari-

ety C. verrucosa rufibranchialis as designated by Odhner (1939) .

In Odhner's synonomy and consequent acceptance of two sub-

species to account for the extreme forms of C. verrucosa, he relied

October, 1969 nautilus 39

on the reports and investigations of Friele and Hansen (1875),

Sars (1878), and Loyning (1922) . It is evident in the records of

these latter investigators that Coryphella verrucosa verrucosa is a

variety found only in Norwegian waters and thus has a very

limited distribution while Coryphella verrucosa rufibranchialis

has a wide boreal-arctic Atlantic distribution. In discussion of

intermediate forms of C. rufibranchialis and C. verrucosa and the

radula of the two typical forms Loyning (I.e.) stated that there

were no distinguishing characteristic differences in the radulae.

The figures of the radula of Coryphella verrucosa by Friele and

Hansen (1875) , of Coryphella rufibranchialis by G. O. Sars (1878)

and of Coryphella rufibranchialis by Odhner (1929) are mutually

similar and agree with observations made by the present author

of the radulae from specimens of Coryphella verrucosa rufibranch-

ialis along the New England Coast. However it is most unfortu-

nate that Odhner (1939) figured the radula of a Coryphella ver-

rucosa rufibranchialis (Fig. 21, p. 59) which is not compatible

with his 1929 figure nor with any other morphological report on

the radula of the subspecies.

It appears that the variety names accepted by Odhner (1939)

should be maintained in the literature until additional knowledge

of the breeding cycle, veliger shells and other characteristics verify

the relationship between Coryphella verrucosa and Coryphella

rufibranchialis.

Sincere thanks are due Dr. E. Deichmann for aid in translation

of the Norwegian papers.

Literature cited

Balch, Francis N. 1909. A spring collecting trip. Notes on New

England Nudibranchs II. Nautilus 23: 33-38.

Friele, H. and G. A. Hansen. 1875. Bidrag til Kundskaben om de

norske nudibranchier. Vidensk. Selsk. Forh. Christiania.

Loyning, Paul. 1922. Nudibranchfaunaen i Drobaksundet. I. Fam.

Aeolididae. Vid. Selsk. Skr. I. Mat.-Nat. Klasse, No. 6. Kristiania.

Miller, M. C. 1961. Distribution and food of the nudibranchiate

mollusca of the South of the Isle of Man. J. Anim. Ecol.

30:95-116.

Moore, George M. 1968. Personal Communication, University of

New Hampshire, Durham, N. H.

Odhner, Nils Hj. 1929. Aeolidiiden aus dem nordlichen Nor-

wegen. Troms Museums Arsh. 1: 1-22.

Odhner, Nils Hj. 1939. Opisthobranchiate mollusca from the

40 NAUTILUS Vol. 83 (2)

western and northern Coasts of Norway. Norske Vid. Selsk. Skr.

1: 1-93.

Sars, G. O. 1878. Bidrag til kundskaben om Norges Arktiske

Fauna. I. Mollusca Regionis Arcticae Norvegiae. Christiania.

Swennen, C. 1961. Data on distribution, reproduction and ecology

of the nudibranchiate molluscs occurring in the Netherlands.

Neth. J. Sea Res. 1: 191-240.

Thompson, T. E. 1964. Grazing and the life cycles of British

nudibranchs. Brit. Ecol. Symp. 4: 275-297.

A NEW NORTHERN RECORD FOR BURSATELLA

LEACHII PLEII RANG (OPISTHOBRANCHIA),

WITH NOTES ON ITS BIOLOGY

By WILLIAM L. KRUCZYNSKI and HUGH J. PORTER

University of North Carolina Institute of Marine Sciences,

Morehead City, North Carolina

Bursatella leachii pleii Rang, the ragged sea hare, can now be

added to the list of tropical animals known from the coast of

North Carolina, U. S. A. The specimens here reported were iden-

tified from descriptions of Eales and Engel (1935) and Henry

(1952) .

Circumtropical in distribution, the species is divided into six

geographical subspecies (Eales and Engel, 1935) . B. I. pleii is

common throughout the West Indies. Henry (1952) gives its

previous northern record as Alligator Harbor, Florida. Abbott

(1954, 1968) reports it only from the west coast of Florida and

the Caribbean.

A population of 27 individuals was found in two of six newly

constructed concrete holding ponds at the Institute of Marine Sci-

ences, Morehead City, North Carolina, and was kept under casual

observation from 21 September until 26 December, 1968. Each

pond measures 29 x 19.5 x 2.7 feet and contains approximately

11,400 gallons of sea water, which is continuously pumped from

nearby Bogue Sound.

Salinity and temperature measurements in the ponds were re-

corded throughout the observation period. Salinity fluctuated

slightly about 30 ppm. Water temperature dropped steadily from

29° C. (24 September) to 3.5° C. (21 December) when it re-

mained constant for nearly a week.

October, 1969 nautilus 41

On 23 October, all specimens were removed from the ponds and

measured. Mean total length was 138 mm.; mean width across the

branchial cavity, 52 mm. Henry (1952) found the average length

and width of 100 living specimens from Alligator Harbor to be

135 and 75 mm. respectively. Although our specimens were not

measured initially, we believe they grew appreciably after first

observation.

Eight individuals were remeasured on 12 December when the

water temperature was 10° C, but the remainder could not be

found because of cloudiness of the water. All animals found were

alive, but sluggish. Average length and width had decreased to

123 and 65 mm. respectively. However, because of low temperature

the specimens may not have been fully extended. We noted later

that an indication of death was noticeable decrease in size. Wet

weights of the eight individuals were found to average 134 grams

which is above the averages (111 and 80 g.) recorded between

March and June in Alligator Harbor by Henry (1952) .

Copulatory chains were observed from 21 September through

3 December. Although egg masses were deposited throughout this

period, and at temperatures as low as 10° C, incubation in the

laboratory showed that eggs did not hatch in water colder than

15° C. We observed that animals cleared an area on the side of the

pond by rasping away algae before depositing eggs. Characteristics

of egg masses agreed with description of Henry (1952) . Cylindri-

cal egg strings contained a mean number of 53 spherical loculi

per cm., and within each loculus there was an average of 28 eggs

or embryos.

Although individuals remained alive at 8° C, they were sluggish

and were not observed to feed. The entire population died when

temperature dropped to 3.5° C, for several days. Before death

each animal extended the gill through the branchial opening.

(Preserved specimens, catalogue #2532, are in the mollusk col-

lection of the University of North Carolina, Institute of Marine

Sciences.)

It is not known how these animals entered the two ponds. Possi-

bly they were attached as eggs to the shells of Glycymeris ameri-

canus Defrance placed in the ponds on 29 June, 1968. These bi-

valves, dredged off Cape Fear, North Carolina, had remained out

of water two days before being placed in the ponds. A more proba-

42 nautilus Vol. 83 (2)

ble method of entrance was as larvae through the sea-water system.

Because the walls of the ponds are covered with barnacles, tuni-

cates and other encrusting organisms, it is known that larval forms

can pass unharmed through the pumping system. Regardless of

which mode of entrance is correct, the finding of B. I. pleii in

North Carolina is a new geographical record for the subspecies.

Acknowledgments

We are grateful to Dr. F. Schwartz, University of North Caro-

lina Institute of Marine Sciences, who first called this population

to our attention.

Literature cited

Abbott, R. T. 1954. American Seashells. D. Van Nostrand Co., Inc.,

Princeton, New Jersey.

. 1968. Seashells of North America, A Guide to Field Identi-

fication. Golden Press, New York, New York.

Eales, N. B. and H. Engel. 1935. The genus Bursatella de Blain-

ville. Proc. Malac. Soc. London, 21: 279-303.

Henry, L. M. 1952. Observations on the sea hare Bursatella leachii

pleii Rang. Stud. Fla. Univ., 1 (7) : 8-14.

SUCCINEA BAKERI HUBRICHT

By LESLIE HUBRICHT

4026 - 35th St., Meridian, Miss. 39301

In a recent paper, Browne & Bruder (1968), reduced Succinea

bakeri Hubricht (1963) to the synonomy of Succinea grosvenori

Lea because they: "were unable to draw distinctions either in

measurements or form of shell, from some specimens of this fauna

and forms described by Lea, Pilsbry, and others, as 5. grosvenori."

As a result, I feel that I should go into more detail as to my

reasons for believing that the Pleistocene fossils of the loess which

have been called S. grosvenori are not that species.

Pilsbry (1948) figured 12 shells as being S. grosvenori (fig.

444a-f, page 820) ; of these shells a and d are S. greeri Tryon. The

shells e and / from Arizona do not look like S. grosvenori, and it is

doubtful that it occurs so far west. These records, and that from

Ogallala, Nebraska (fig. 444c) need to be anatomically verified.

Thus, only the two shells b, the presumed paratypes from Lea,

can be accepted as 5. grosvenori. Pilsbry also reproduced Lea's

figure (fig. 452i, page 833) which one could safely use. Pilsbry

October, 1969 nautilus 43

records the species as far north as Great Slave Lake, Canada. That

it occurs so far north is very doubtful. The most northern records

for anatomically identified material are from Kansas, but there

has been no recent work on the genus from farther north, so the

northern limit of the species is unknown.

There are three species of Succinea found in the southern and

central United States which, although they are quite distinct

anatomically, cannot be identified by their shells with any real

certainty. They are: S. grosvenori, S. greeri, and 5. indiana (Pils-

bry) . These three species occur in similar habitats. They are all

found on bare or sparsely weedy ground in sunny situations. I have

never found any of them in the woods. Of the three, 5. grosvenori

prefers the wetter habitat. It is usually found in the vicinity of

water, even near pools which remain for a few days after a shower.

5. greeri occupies the driest habitat. I found it in Oklahoma crawl-

ing on a sheer, south-facing cliff after a shower. It is frequently

found on loess banks with a southern exposure. Shells of Catinella

vermeta (Say) and C. texana Hubricht have also been misidenti-

fied as S. grosvenori.

There are some shell differences between S. grosvenori and

5. bakeri which, though not always reliable in the Succineidae

where shell differences between species can be very small, do in-

dicate a strong probability that they are distinct species. In S. gros-

venori the start of the nuclear whorl is raised; in S. bakeri it is

depressed. In 5. grosvenori the whorls are usually somewhat shoul-

dered; in S. bakeri they are not. Under the microscope the surface

of 5. grosvenori shows some spiral sculpture. None was seen on

any of the shells of 5. bakeri examined. The habitats of the two

species are quite different. 5. grosvenori is a sun-loving species of

open ground, while 5. bakeri, judging by its associated fauna, must

have been a woodland species.

Literature cited

Browne, Ruth G. & Pamela M. Bruder. 1968. Wisconsin mol-

luscan faunas from Henderson County, Kentucky. Bull. Amer.

Paleont. 54: 191-275.

Hubricht, Leslie. 1963. Some Succineidae, with a new species.

Nautilus 76: 135-138.

Pilsbry, Henry A. 1948. Land mollusca of North America (north

of Mexico) . Acad. Nat. Sci. Philadelphia, Mon. 3, 2: 521-1113.

44 nautilus Vol. 83 (2)

NONMARINE MOLLUSKS OF THE

KATALLA REGION, ALASKA

By SAMUEL J. TUTHILL1 and ROBERT L. JOHNSON2

In July of 1967, casual collections of terrestrial and freshwater

gastropods were made along the coast of Katalla and Controller

Bays (northeastern Gulf of Alaska, 60°00T0" N by 144°30'00" W,

see U.S. Geological Survey maps, CORDOVA A-l, A-2, B-l, and

B-2, scale 1:63360) . The molluscan fauna of the coastal region is

different from that of the adjacent Copper-Martin-Bering Rivers

region. Lymnaea palustris, Haplotrema sportella, Vespericola Col-

umbiana, Vertigo modesta, Succinea cf. S. rusticana, var. alaskana,

Oxyloma cf. O. retusa, Deroceras laeve, and Euconulus fulvus alas-

kensis were found at Katalla. Only D. laeve and E. fulvus alaskensis

are part of the widely distributed molluscan fauna typical of the

Copper-Martin-Bering Rivers region. The coastal region around

Katalla, the Cordova region, and the course of the Copper River

and Northwestern Railroad in the Copper River trench was the

site of much American (non-native Alaskans) activity between

1898 and 1938 and to a lesser degree between 1938 and the present.

The region east of the Copper River proper, the Martin River

valley and all but the coastal part of the Bering River valley (only

12 miles from the village site of Katalla) has seldom been visited

by non-Alaskan Americans. The molluscan fauna of the two

regions is known from the investigations of Bickley (in press) in

the Tasnuna Valley along the former right of way of the C.R.&NW

RR; Tuthill (1963) which reported the results of investigations

in the Martin River valley in 1962; Tuthill, Field, and Clayton

(1968) which reported the nature and structure of the molluscan

population of the Sherman and Sheridan Glacier valleys; investi-

gations by the senior writer in 1963-5, 1967-8; and the studies re-

ported here.

Differences in the molluscan fauna in the two regions show a

strong association with the activity of man. We interpret these

distributional phenomena to be the result of accidental introduc-

tions of Lymnaea palustris, Haplotrema sportella and Vespericola

columbiana by non-Alaskan American developers during the first

'Iowa Geological Survey, Iowa City, Iowa 52240

2 Muskingum College, New Concord, Ohio.

October, 1969 \ ujtilus 45

third of the twentieth century. It is likely that Vertigo modesta, V .

ovata, Succinea rusticana and Oxyloma retusa were also introduced

by newly arriving Americans, but the general distribution of these

taxa is such as to reduce our conviction from that we hold with

respect to the three species mentioned above.

Table 1 shows the species composition of the faunas in the

Katalla and Copper-Martin-Bering Rivers regions respectively.

Table 2 shows the estimated population of Lymnaea palustris

in a brackish tidal pool at Katalla.

Lymnaea palustris (Miiller, 1774) .

Taylor (1965, p. 598-599) has pointed out the fact that animals

which had shells conforming to the conchological definition of this

species were studied by Jackiewicz (1959) in Poland and found to

constitute three distinct neontologic entities. The shells we col-

lected at Katalla resemble the descriptions and figures of the fol-

lowing taxa in the indicated reports: Galba palustris, Baker (1911,

p. 298-322), Lymnaea palustris, Hubendick (1951, p. 119-122)

and Stagnicola palustris, La Rocque (1968, p. 443-446) .

L. palustris was found only in brackish pools high on the beach

east of the Katalla River. A series of interconnected pools, about 2

feet in maximum depth, parallel to the coast at the shoreward

margin of the beach, about i/2 mile from normal high sea strand-

line was found to contain an abundant, crowded population of

the taxon. During periods of heavy rain and shortly following

them, groundwater supplies a sufficient amount of water to these

pools to effect flow in the interconnecting streams and water flows

out of the system into an incipient back-beach lagoon and ulti-

mately into the Katalla River estuary. It is conceivable that dur-

ing heavy spring and fall storms and periods of very high tides

that sea water is delivered directly to this part of the beach, but

this was never observed by us. The salt content of the water is prob-

ably the result of leaching from the beach-sand shoreward of the

system of pools. During periods of low or no rainfall the water

level in these pools drops sharply. The entire population of one

pool was collected. This pool had a water level which was 10

inches below the high strandline and the snails were concentrated

in the upper 6 inches of the water. The few snails which were

observed in the deeper water (the pool was 24 inches deep at its

deepest point) were the largest individuals in the population.

46 nautilus Vol. 83 (2)

Several specimens were observed crawling about on the moist

ground and on grass stalks that grew at the pool edge. The area

of beach in which the pools occur has been captured by grass,

moss, and shrubs. Where sand bottoms occurred snails were scarce.

The junior writer collected the entire population of a pool

which had a surface area of 90 sq. ft and an estimated volume of

160 cubic ft. At its highest stand the pool had had an area of 120

sq. ft and an estimated volume of 240 cubic ft. The structure and

density of the population of snails was estimated from a 10 per-

cent sample of the 1900 individuals collected. There were 360

empty shells in the area collected and these were examined to

determine whether mortality had been more prevalent at any one

age class. A general living population density of 16 individuals/

ft2 of the plane at high strandline is not representative of the

actual concentration of animals in the habitat. As mentioned

above, only larger snails were common below 6 inches depth from

the water surface at time of collection and the number of indi-

viduals above the water surface was also few. An estimated actual

density of 23 individuals per sq. ft. of water surface and 36 indi-

viduals per cubic ft. of water is closer to the conditions which

actually existed. Table 2 shows the distribution of L. palustris

with number of whorls and length classes of 2 mm. A few shells

were seen to have been drilled, presumably by a radula. Thus,

although we did not observe it, cannibalism may possibly be a

behavior common to much crowded L. palustris.

During six field seasons of collection in the region between Cor-

dova, Chitina, and Katalla we have encountered this snail only

at Cordova, the Tasnuna Valley, and at Katalla, all localities where

white American activity has been intensive. This pattern of dis-

tribution inclines us to believe that the species has been acciden-

tally introduced into the region.

Vertigo modesta (Say, 1824).

This tiny terrestrial snail was found on logs under stands of

mature Sitka spruce growing on a raised beach ridge which is

now about 35-40 ft. above the sea and about i/2 mile inland. The

Katalla region has been tectonically active during the past 14,000

radiocarbon years and as recently as 1964 was raised from 6 to 8

ft. during the Alaskan Earthquake. Thick accumulations of de-

caying alder leaves, spruce needles, and mosses cover the forest

October, 1969 nautilus 47

floor. The snails were found only on logs which lay in the leaf

litter. Twelve living snails were collected in one locality. Despite

thorough search in similar habitats elsewhere in the region this

species was not encountered again. Also of surprise to us was the

absence of Vertigo columbiana which is an abundant and vigorous

pioneer species in the recently deglaciated regions near the Sher-

man and Sheridan Glaciers and is found in large numbers on the

stagnant, drift-insulated terminus of the Martin River Glacier.

Oxyloma cf. O. rctusa (Lea, 1834) .

The genera and species of the family Succineidae are differen-

tiated on the morphology of their soft parts. Thus, assignment to

genus and species is tentative, but the shells conform to the taxa

indicated and as defined in Pilsbry (1948, p. 785-788) . The three

specimens of this taxon which we collected at Katalla were found

on the moist margins of the pools described as the habitat of

L. palnstris. They were among specimens which we have assigned

to Succinea rusticana. On an occasion previous to the time when

we collected them, the senior writer observed much larger num-

bers of the family in these habitats. Presumable predation by

birds, tracks of which were numerous, explains the reduction in

the succinid population.

Succinea cf. 5. rusticana var. alaskana Dall, 1905.

Four specimens of this taxon were collected with the Oxyloma

around the pools mentioned above. The specimens from Katalla

are smaller than the measurements given by Pilsbry (1948, p.

824-825) . The number of whorls they contained suggests they were

not yet fully grown when we collected them.

Haplotrema sportella (Gould, 1846).

This species was collected from Whale Island at Cape Martin,

about 3 miles west of Katalla and in the village itself. At Whale

Island (a tombolo-tied island since the uplift accompanying the

Alaskan Earthquake of 1964) the snails were found, both live

animals and shells, in the grass and leaf litter at the top of 30-50

foot-high sea cliffs. They were living in a zone which receives direct

splash and much spray during the frequent violent storms of this

part of the Gulf of Alaska. At Katalla they were found under the

rotting logs and lumber of the delapidated buildings. Specimens

of this species were collected from logs and lumber trash in the

harbor of Cordova in 1963. Pilsbry (1946, p. 226) cites Vancouver

48

NAUTILUS

Vol. 83 (2)

Table 1. Geographic distribution of nonmarine

mollusks in south-central Alaska.

Island as the most northern and westerly occurrence of this species.

W. J. Eyerdam collected this species from Unalaska in June 1932

(ANSP #177891, 2 specimens), thus extending the range 38° of

long, to the west and 3° of lat. to the north. Mr. Rae Baxter's col-

lections from Cordova further extends the range of this species

5 1/2° of latitude to die north. I suspect that search for this species

and for Vespericola would prove successful in almost any coastal

village site which was once a port of call of Seattle-based ships. We

believe that this species was introduced.

Deroceras laeve (Muller, 1774).

Five living slugs were collected from dead logs and moss in the

same area as that in which Vertigo modesta was found. Here, as

elsewhere in the region, where the senior writer has been studying

for the past 7 years, the slug occurred singly. In the interior of the

coastal plain it is a common mollusk and we were surprised at its

scarcity in the Katalla region. Suitable habitats were not wanting.

Vespericola columbiana (Lea, 1838) .

Specimens were collected at Whale Island with Haplotrema and

50 nautilus Vol. 83 (2)

at Palm Point (about H/2 miles west of the village site) in dune

grass mats. The specimens belong to the species in its strict sense

and do not have the characteristics of the subspecies. Mr. Rae Bax-

ter found this species with Haplotrema at Cordova. His collections

constitute an extension of the known range of the species of about

1 ° of latitude to the north. We would expect to find this species in

almost any coastal village or former village site which had been

supplied by west coast-based steamers.

Euconulns fulvus alaskensis (Pilsbry, 1899) .

Only three specimens of this species were found near Katalla. It

was collected from logs in alder leaf litter as described in the pre-

vious discussion of Deroceras laeve. Its rarity is surprising because

this taxon is a vigorous and abundant pioneer species of the inner

coastal plain.

Summary

Because they occur only at sites of current or former occupancy

by non-Alaskan Americans who came to the Gulf of Alaska in

relatively large numbers between 1898 and 1938, we believe that

Haplotrema sportella, Vesparicola columbiana, and Lymnaea

palustris are accidentally-introduced species. Vertigo modesta, V.

ovata, Succinea rusticana, and Oxyloma retusa have general north-

ern North American distributions which make us less convinced

that they were accidentally introduced, but their distribution

within the region is such that strong association exists between the

activity of man and the colonization patterns of the snails.

Acknowledgments

We are in the debt of other members of the 1967 field party for

helping us collect mollusks, and for being patient with us when

we delayed their progress. These persons were Dr. Leslie A. Sirkin,

Adelphi University, Miss Catherine Smith and Mr. James K. Wat-

son, Muskingum College. The National Science Foundation sup-

ported the research project of which this study was a collateral

activity. We are grateful for their support in the form of Grant

G. A. 869.

Literature cited

Baker, F. C, 1911, The Lymnaeidae of North America, Recent

and fossil: Chicago Acad. Sci., Spec. Pub'l. no. 3, 537 p., 57 pi.

Bickley, W. B., Jr., in press, The presences of gigantism in a mol-

luscan fauna from south-central Alaska.

Hubendick, Bengt, 1951, Recent Lymnaeidae: Kungl. Svenska

October, 1969

NAUTILUS

51

Table 2, Estimated population of Lymnaea

palustris in a 90 ft2 (160 ft3) pool of brackish

water on the beach east of the Katalla River

estuary, Katalla, Alaska.

Vetenskapsacademiens Handlingar, Fjarde Serien. Band 3, no.

1, 223 p., 5 pi.

Jackiewiecz, Maria, 1959, Badania nad zmienoscia i stanowiskiem

systematycznym Galba palustris O. F. Miiller: Poznan, Prace

Kom. Biol., v. 19, no. 3, 54 p.

La Rocque, Aurele, 1968, Pleistocene Mollusca of Ohio: Ohio

Dept. Nat. Resources, Div. Geol. Survey Bull., 62, pt. 3, 197 p.,

6 PI.

Pilsbry, H. A., 1940, Land Mollusca of North America (north of

Mexico) : Philadelphia Acad. Nat. Sci., Mono., no. 3, v. 1, pt.

2, 430 p.

52 nautilus Vol. 83 (2)

-, 1946, Land Mollusca of North America (north of Mexico) :

Philadelphia Acad. Nat. Sci., Mono., no. 3, v. 2, pt. 1, 520 p.

-, 1948, Land Mollusca of North America (north of Mexico) :

Philadelphia Acad. Nat. Sci., Mono., no. 3, v. 2, pt. 2, 592 p.

Taylor, D. W., 1965, The study of Pleistocene nonmarine mollusks

in North America: The Quarternary of the United States,

Princeton, Princeton Univ. Press, p. 597-611.

Tuthill, S. J., 1963, Preliminary report of the molluscan fauna of

the Martin River Glacier and associated area: Veliger, v. 6,

no. 2, p. 84-90, 1 map.

Tuthill, S. J., W. O. Field, and Lee Clayton, 1968, Postearthquake

studies at Sherman Glacier: in The Great Alaska Earthquake

of 1964 (Hydrology Volume), NAS Pub. 1603, Washington,

National Academy of Sciences, p. 318-328.

ILLUSTRATIONS OF LAMARCK'S TYPES OF

NORTH AMERICAN UNIONIDAE MOSTLY IN THE

PARIS MUSEUM

By RICHARD I. JOHNSON

Museum of Comparative Zoology

Lamarck's private collection of shells was sold, after his death

in 1829, to the Due de Rivoli (Prince Massena) . It was later pur-

chased by Baron B. Delessert, who in 1841, published a sump-

tuous iconography of the collection. This collection is now in the

Geneva Museum. Much of the remaining material on which

Lamarck based his names is in the Paris Museum (Museum

National d'Historie Naturelle) , including the collections of

Valenciennes and J. B. L. d'Audebard, Baron de Ferussac.

Most of these unionid types are illustrated here for the first

time. I am grateful to Mr. H. Chevallier who prepared the photo-

graphs and to Dr. E. Fischer-Piette whose permission made their

preparation possible.

Figured types

carinifera, Unio Figure 4

1819, An. sans Vert., 6:74, No. 16 (Habite la riviere Hudson de

l'etat de New Yorck. Cabinet de M. Valenciennes) . Holotype

Paris Museum, length 73, height 50, width 32 mm. Is Elliptio

complanatus (Lightfoot) 1786.

clava, Unio Figure 9

1819, An. sans Vert., 6:74, No. 18 (Habite dans le lac Erie

[erroneous] Michaud fils) . Holotype Paris Museum, Length 75,

height 45, width 33 mm.; also a para type, described as var. b. (la

October, 1969 nautilus 53

riviere cle la Nouvelle Ecosse [erroneous, does not occur in Nova

Scotia |) length 55, height 33, width 26 mm. The type locality is,

here, restricted, to the Ohio River, Cincinnati, Ohio. Discussed by

Ortmann and Walker (1922:25) . Is Pleurobema clava (Lamarck) .

coarctata, Unio Figure 5

1819, An. sans Vert., 6:73, No. 1 1 (Habite la riviere d'Hudson

| New York], Cabinet de M. Valenciennes). Holotype Paris

Museum, length 88, height 47, width 25 mm.: also a smaller para-

type in the Geneva Museum. Is Elliptio complanatus (Lightloot)

1786.

crassidens, Unio Figure 2

1819, An. sans Vert., 6:71, No. 3 (Habite l'Amerique septen-

trionale, dans le Mississippi, l'Ohio, et plusieurs lacs) . The speci-

men in the Paris Museum mentioned by Lamarck as var. b, is,

here selected, lectotype, length 66, height 50, width 29 mm. The

type locality is, here, restricted to the Ohio River, Cincinnati,

Ohio.

A composite species, restricted by Lea (1834: 87 [199]) to var. b.

(du lac Erie [erroneous]) on the basis of his identification of a

paratype of it, in the Massena collection, as U. cuneatus Barnes

1823. Lea also identified var. a, as U. trapezoides Lea 1831 -Plec

tomerus dombeyana Valenciennes 1827. Lamarck refers var. c to

U. crassus Say, 1817, non Retzius, 1788 Actionaias ligamentina

Lamarck 1819. Is Elliptio crassidens (Lamarck) .

georgina, Unio Figure 3

1819, An. sans Ver., 6:74, No. 17 (Habite le lac George [New

York], Cabinet de M. Valenciennes). Holotype Paris Museum,

length 59, width 29, height 15 mm. Is Elliptio complanatus (Light-

foot) 1786.

glabrata, Unio

1819, An. sans Vert., 6:75, No. 21 (Habite la rivere de l'Ohio

[erroneous], Midland) . Holotype Paris Museum, length 70,

height 32, width 20 mm. Is Elliptio complanatus (Lightfoot) 1786.

ligamentina, Unio Figure 7

1819, An. sans Vert., 6:72, No. 7 (Habite la rivere de l'Ohio,

Midland) . Holotype Paris Museum, length 73, height 50, width

32 mm. The type locality is, here, restricted to the Ohio River,

Cincinnati, Ohio.

Lea examined the type (1834: 88 [200]) and identified it as

U. crassus Say 1817, non Retzius 1788. Lamarck's name was used

consistently in the literature until 1922 when Ortmann and

Walker (1922:47) declared the species to be unrecognizable from

the description and used carinata Barnes 1823. Their action was

unwarranted in view of Lea's identification and the availability of

the type. Is Actinonaias ligamentina (Lamarck) .

54 nautilus Vol. 83 (2)

luteola, Unio Figure 8

1819, Anim. sans Vert., 6:79, No. 40 (Habite la riviere Susque-

hana [Pennsylvania and New York] et celle Mohancks [Mohawk,

New York]. Holotype Paris Museum, length 69, height 38, width

24.5 mm. The type locality is here restricted to the Susquehanna

River, Columbia, York Co., Pennsylvania.

Lea (1834:91 [203]) identified the type as U. siliquoideus

Barnes 1823, claiming the localities to be in error. Ortmann and

Walker (1922:61) declared the species to be unrecognizable from

the description, but this, as pointed out above would not be ger-

mane if Lea's identification is correct. However it is difficult to

determine the exact identity of the type, if it is from the Mohawk

River, since as pointed out by Clarke and Berg, (1959:58-62, 68-

70) Lampsilis radiata radiata (Gmelin) and the western L. r.

siliquoidea Barnes co-mingle and intergrade completely in the

lower St. Lawrence River drainage of New York.

The Mohawk River probably did not have siliquoidea in it

before the completion of the Erie Canal, some years after luteola

was described. Since only L. r. radiata occurs in the Susquehanna

River, the locality is restricted to it. This makes further attempts

(Wheeler, 1963:58) to resurrect this name unnecessary. Is Lamp-

silis radiata radiata (Gmelin, 1791) .

naviformis, Unio Figure 11

1819, An. sans Ver., 6:15, No. 20 (Habite la riviere de l'Ohio,

Michaud fils) . Holotype Paris Museum. Length 75, height 32,

width 27 mm. Type locality is here restricted to the Ohio River,

Cincinnati, Ohio. Is Quadnda cylindrica (Say) 1817.

obliqua, Unio Figure 14

1819, An. sans Vert., 6:12, No. 8 (Habite la riviere de l'Ohio, A.

Michaud). Holotype Paris Museum [lost]. The specimen in the

type collection from the Wabash | River, Indiana] from Le Sueur

Explanation to Figures

Figure 1. Unio rariplicata Lamarck. Holotype (nat. size) .

Figure 2. Unio crassidens Lamarck. Lectotype (nat. size) .

Figure 3. Unio georgina Lamarck. Holotype (nat. size) .

Figure 4. Unio carinifera Lamarck. Holotype (nat. size) .

Figure 5. Unio coarctata Lamarck. Holotype (nat. size) .

Figure 6. Unio purpurascens Lamarck. Lectotype (nat. size) .

Figure 7. Unio glabrata Lamarck. Holotype (nat. size) .

Figure 8. Unio ligamentina Lamarck. Holotype (nat. size) .

Figure 9. Unio clava Lamarck. Holotype (nat. size) .

Figure 10. Unio rotandata and U. suborbicnlata Lamarck. Holotype United

States National Museum 85760 right valve, (nat. size) .

Figure 11. Unio luteola Lamarck. Holotype (nat. size) .

Figure 12. Unio retusa Lamarck. Holotype (nat. size) .

Figure 13. Unio recta Lamarck. Holotype (nat. size) .

Figure 14. Unio naviformis Lamarck Holotype (nat. size) .

October, 1969

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60 nautilus Vol. 83 (2)

cannot be a primary type since Le Sueur was not in Indiana until

1826. Lea saw the collection in the Paris Museum in 1832 (1834;

88 1 200]) and identified it as U. undatus Barnes. Ortmann and

Walker (1922:21) declared this species to be unrecognizable from

the description, and since it is impossible to be sure it Lea saw

the actual type, or only Le Sueur's specimen, the use of the next

available taxon, Obovaria cordata Rafinesque, should be followed.

Is possibly Plearobema cordatus (Rafinesque) 1820.

purpurascens, Unio Figure 6

1819, An. sans Vert., 6:73, No. 12 (Habite les rivieres de New

Yorck, Cabinet de M. Valenciennes) . Since the type lot contains

two species, var, c, Paris Museum is, here selected as the lectotype:

length 92. height 48, width 28 mm. (du lac Champlain [New

York] Le Sueur) . Is Elliptio complanatus (Lightfoot) 1786.

rariplicata, Unio Figure I

1819, An. sans Vert., 6:71, No. 5 (Habite la riviere de l'Ohio,

Michaud) presumed holotype Paris Museum, length 70, height 50,

width 35 mm. Lamarck gives a length of 62 mm. The type locality

is here restricted to the Ohio River, Cincinnati, Ohio. Is Amb'lema

plicata (Say) 1817.

recta, Unio Figure 13

1819, An. sans Vert., 6:74, No. 19 (Habite le lac Erie, Michaud) .

Holotype Paris Museum, length 100, height 41, width 31 mm.

The original label has the additional data, "de la [vicin]ite de

Niaga [Niagara Falls, New York]." Is Lampsilis recta (Lamarck) .

retusa, Unio Figure 12

1819, An. sans Vert., 6:72, No. 9 (Habite les rivieres de la Nou-

velle Ecosse [erroneous], A. Michaud). Presumed holotype Paris

Museum, length 44, height 45, width 30 mm., Lamarck gives a

length of 47 mm. A smaller paratype, length 40, height 42, width

31 mm. The type locality is here restricted to the Ohio River,

Cincinnati, Ohio. Is Obovaria retusa (Lamarck) .

rotundata, Unio Figure 10

1819, An. sans Vert., 6:75, No. 24 (Habite . . . restricted to:

Bayou Teche, St. Mary Parish, Louisiana, by Clench and Turner,

1956, 7:192). Type in Cabinet de M. Daudebard | Baron Ferus-

sac] et celui de M. Faujas [de Saint Fond], not in Paris Museum.

Is Glebula rotundata (Lamarck) . See under next item.

suborbiculata, Unio Figure 10

1819, An. sans Vert., 6:81, No. 48 (Habite ... les eaux douces

des climats chauds? restricted to: Bayou Teche, St. Mary Parish,

Louisiana, by Clench and Turner, 1956, 7:192, Cabinet de MM.

Daudebard [Baron Ferussac] et Faujas [de Saint Fond]) . Not

in Paris Museum. Measured holotype United States National

Museum 85760 consists of a polished right valve, length 80, height

68.5, width 19 mm. Lea (1831:189 [201]) was presented this

October, 1969 nautilus 61

specimen by Baron Ferussac and believed it to be the individual

cited by Lamarck. Ferussac (Mag. de Zool., 5: cl. 5, Nos. 59, 60,

p. 31, note 10) pointed out that Lamarck based his description of

Unio rotunda and suborbicidata on the same specimen at two dif-

ferent times. There is an additional specimen of this species from

Faujas in the Paris Museum, length 92, height 92, width of the

valve 25 mm. Is Glebula rotundata (Lamarck).

Literature cited

Clarke Jr., A. H. and C. O. Berg. 1959. The freshwater mussels

of central New York. Agricultural Experiment Sta., Cornell

Univ. Ithaca, New York. Mem. 367, pp. 1-79. 7 pis.

Clench, W. J. and R. D. Turner. 1956. Freshwater Mollusks of

Alabama, Georgia, and Florida from the Escambia to the Suwan-

nee River. Bull. Florida State Mus., / (3) : 97-239, pis. 1-9.

Delessert, B. 1841. Recueil de Coquilles Decrites par Lamarck dans

son Historie Naturelle des Animaux sans Vertebres et non en-

core figurees. 40 col. pis. with letterpress, folio, Paris.

Johnson, R. I. 1952. A study of Lamarck's types of Unionidae and

Mutelidae. Nautilus, 66:63-67, 90-95.

Lamarck, J. B. P. de M. 1819. Historie Naturelle des Animaux

sans Vertebres, 6(l):68-88. Paris.

Lea, I. 1834. Observations on Lamarck's naiades. Trans. Amer.

Philos Soc, 6:86-94. Reprinted: 1834, Obs. Unio, 1: 198-206.

Ortmann, A. E. and B. Walker. 1922. On the nomenclature of

certain North American naiades. Occ. Papers, Mus. Zool., Univ.

Michigan, no. 122, pp. 1-75.

Wheeler, M. J. 1963. Type of Unio luteolus Lamarck 1819. Nau-

tilus, 77:58-61.

NOMENCLATORIAL NOTE: ARCTICIDAE

VERSUS CYPRINIDAE (MOLLUSCA; BIVALVIA)

By KENNETH J. BOSS

Museum of Comparative Zoology, Harvard University, Cambridge, Mass.

In a recent paper (Boss, 1968) I used the familial designation,

Cyprinidae, for members of a molluscan family which reached the

height of its diversity during the Jurassic and Cretaceous (Casey,

1952) and which today is represented by only one species, Arctica

islandica (Linnaeus, 1767) (Lamy, 1920) . The nomen Cyprinidae

should not be used in malacological literature and my usage was

in error.

Dr. Bruce B. Collette of the Bureau of Commercial Fisheries,

Systematics Laboratory, Washington, D.C., called my attention to

the existence of the family name Cyprinidae in Pisces for the

62 nautilus Vol. 83 (2)

important and extremely large group of freshwater fishes (about

2,000 species) commonly referred to as minnows.

Nomenclatorial comments and a clarification of generic and

familial usage of this name in malacology have been presented by

Nicol (1951) and Bowden and Heppell (1968) . For Venus

islandica Linnaeus 1767, Lamarck (1818) applied the generic name

Cyprina (type-species, Venus islandica Linnaeus, by subsequent

designation, Children 1823, p. 312) ; earlier (1812) he had used

the French vernacular "cyprine," which, of course, has no nomen-

clatorial status. Subsequently, Lamarck's Cyprina gained wide ac-

ceptance and the family-group name Cyprinidae, was introduced

for this molluscan group by Geinitz (1846). Schumacher (1817)

described Arctica with A. vulgaris Schumacher (—Venus islandica

Linnaeus) as type-species by monotypy, but this generic name was

preoccupied in Aves by Arctica Nozeman and Vosmaer, 1758. How-

ever, when the International Commission of Zoological Nomen-

clature voted to suppress the work of Nozeman and Vosmaer which

was a Dutch translation of Moehring's pre-Linnaean work, Avium

Genera (1752) (Hemming, 1944; 1954), the avian name Arctica

became invalid and the junior homonym Arctica Schumacher 1817

was available for the mollusk and became the senior homonym of

Artica Merrem 1819 (Aves).

The molluscan family-group name, Arcticidae Newton 1891 is

a junior synonym of Cyprinidae Geinitz 1846. In accordance with

priority (Article 23 d) , the senior synonym should be used. How-

ever, Geinitz's name is also a junior homonym of the piscean

Cyprinidae Rafinesque 1815, so Arcticidae Newton should be used

for the molluscan taxon.

In malacological literature, Nicol (1951) pointed out that

Arcticidae and Arctica should be used, and he has been followed

by Newell (1965) who outlined the nomenclature to be employed

in the forthcoming and authoritative section on the Bivalvia for

the Treatise of Invertebrate Paleontology. Some authors have

maintained Arcticidae but used Cyprinacea as a superfamilial

appellation (Tebble, 1966) while others have employed Cyprinidae

(e.g. Eberzin [in] Orlov, 1960; Zatsepin and Filatova, 1961;

Ziegelmeier, 1962; Davitaschvili and Merklin, 1966). According

to the Code, Article 40(a), "If a family-group name changed before

1961 because of such synonymy, has won general acceptance, it is

October, 1969 nautilus 63

to be maintained in the interests of stability" (Nicol, 1951, pointed

out the synonymy of Cyprina and Arctica and employed Arcticidae

in preference to the older Cyprinidae) . Even though there might

be some argument concerning the "general acceptance" of

the nomen Arcticidae, a strong case can be made for the use of

Arcticidae over the older Cyprinidae, primarily because the latter

is homonymous with a taxon of fishes. Further, in accordance with

Recommendation 40A, "... a family-group name . . . should be

cited with its own author and date, followed by the date of the

replaced name in parenthesis," the familial placement of Arctica

should read Arcticidae Newton 1891 (1846).

One further item concerns the nomenclature of this group of

bivalves. In 1965, Newell proposed a new suborder Arcticina for

the five superfamilies Arcticacea, Dreissenacea, Glossacea, Cor-

biculacea, and Veneracea. Alluding to the principle that ordinal-

level names be derived from the root-word of their respective

nomenclatorial types, Vokes (1967) emended Newell's name to

Venerina. However, his emendation is, by extrapolation of the

Code (Article 33 (2) (ii) ) , to be considered "unjustified" and is

to be construed as a junior objective synonym of Arcticina Newell

1965.

Literature cited

Boss, K. J. 1968. New species of Vesicomyidae from the Gulf of

Darien, Caribbean Sea (Bivalvia; Mollusca) . Bull. Mar. Sci.,

18(3) .-731-748, 28 figs.

Bowden, J. and D. Heppell. 1968. Revised list of British mollusca.

2. Unionacea-Cardiacea. J. Conch. 26:237-272.

Casey, R. 1952. Some genera and subgenera, mainly new, of

Mesozoic heterodont lamellibranchs. Proc. malac. Soc. London,

29: 121-176, pis. 7-9.

Children, J. G. 1823. Lamarck's Genera of Shells, translated from

the French. Quart. Jour. Sci., 14: 298-322.

Davitaschvili, L. Sch. and R. L. Merklin. 1966. Spravochnik po

ekologii morskikh dvustvorok. (Monograph on the ecology of

marine bivalves) . Moscow, 348 pp.

Eberzin, A. G. [in] Orlov, Y. A., ed. 1960. Osnovi Paleontologii.

Molliuski, Dvustvorchatie, Moscow, 300 pp., 44 pis.

Geinitz, H. B. 1846. Grundriss der Versteinerungskunde. Dresden

and Leipzig, 813 pp., 26 pis.

Hemming, F. 1944. Opinions and declarations rendered by the

International Commission on Zoological Nomenclature Opinion

5, The status of certain pre-Linnean names reprinted subsequent

64 nautilus Vol. 83 (2)

to 1757. London, /, pt. 14:115-126.

Hemming, 1954. [Ibid]. Opinion 241. Rejection for nomenclatorial

purposes of Nozeman and Vosmaer, 1758. Geslachten der

Vogelen, a Dutch translation of Moehring's pre-Linnaean work

entitled Avium Genera published in 1752. London, 5, pt. 2 pp.

13-22.

Lamarck, J. B. 1812. Extrait du Cours de Zoologie du Museum

d'Histoire Naturelle sur les Animaux sans Vertebres. Paris.

Lamarck, J. B. 1818. Histoire naturelle des animaux sans vertebres,

Paris, 5, 612 pp.

Lamy, E. Revision des Cypricardiacea et des Isocardiacea vivants

du Museum d'Histoire naturelle de Paris. Jour de Conch.,

64 (4) : 259-307. 1920.

Linnaeus, C. 1767. Syst. Nat., ed. 12, 1, pt. 2, pp. 533-1327.

Holmiae.

Merrem, B. 1819. [in] Ersch. J. S. and J. G. Gruber. Allgemeine

Encyclopadie Wissensch. und Kiinste. Leipzig, sect. I, 2, p. 406.

Moehring, Paul Heinrich Gerhard. 1752. Avium Genera. Bremae,

88 pp.

Newell, N. D. 1965. Classification of the Bivalvia. Am. Mus.

Novit., no. 2206, 25 pp., 3 figs.

Newton, R. B. 1891. Systematic List . . . British Oligocene and

Eocene Mollusca. London, British Museum (Natural History) ,

365 pp.

Nicol, D. 1951. Recent species of the veneroid pelecypoda Arctica.

Jour. Wash. Acad. Sci., 41 (3) : 102-106.

Nozeman, Cornells and A. Vosmaer. 1758. Geslachten der Vogelen

. . . Amsteldam: 97 pp., 2 pis.

Rafinesque, C. S. 1815. Analyse de la Nature ou Tableau de

l'Univers et des Corps Organises. Palerme, 224 pp.

Schumacher, C. F. 1817. Essai d'un nouveau systeme des habita-

tions des vers testaces. Copenhague, 287 pp., 22 pis.

Smith, E. A. 1912. On the generic name to be applied to the Venus

islandica Linnaeus. Proc. Malac. Soc. London, 10 (2) : 105-106.

Tebble, N. 1966. British bivalve seashells. London: British

Museum (Natural History), 212 pp.

Vokes, H. E. 1967. Genera of the Bivalvia, a systematic and biblio-

graphic catalogue. Bull. Amer. Paleo., 5/(232): 112-394.

Zatsepin, V. I. and Z. A. Filatova. 1961. The bivalve mollusc

Cyprina islandica (L.) , its geographic distribution and role in

the communities of benthic fauna. Trans. Inst. Ocean., Acad.

Sci. USSR, 46:201-216. (translation 74732, Dept. Northern Af-

fairs and National Resources, Ottawa, Canada) .

Ziegelmeier, E. 1962. Die Muscheln (Bivalvia) der deutschen

Meeresgebiete. Helgolander wissen. Meeresuntersuch., 6(1):

1-56, 14 pis.

October, 1969 nautilus 65

MARINE MOLLUSKS OF PORT MOLLER BAY,

ALASKA PENINSULA

By JAMES X. CORGAN

Austin Peay State University, Clarksville, Tennessee

Port Moller Bay and the confluent Herendeen Bay form the

largest indentation in the Bering Coast of the Alaska Peninsula.

Mollusks from this region were described and illustrated by Grew-

ingk in 1850 (fide Dall, 1921) . There is no other comprehensive

report on Mollusca from the Bering shore of the Alaska Penin-

sula, although the distribution of one genus was recently described

(Corgan, 1966) .

During the summer of 1965, I collected in the Port Moller-

Herendeen Bay area. Collections were made in the course of geo-

logic mapping and time did not permit intensive sampling. Still,

mollusca of the area are so little documented that presentation of

a brief list seemed desirable.

Abundance estimates given below are apparent frequencies of

beachworn, dead shells. They do not adequately reflect the abund-

ance of living specimens which could not be evaluated. Wherever

beach shells were common, a sediment sample was collected and

examined for minute species. An Odostomia was the only micro-

scopic mollusk observed.

Gastropods

Acmaea cribraria (Gould) . Common.

Acmaea digitalis (Eschscholtz) . Rare.

Acmaea pelta (Eschscholtz) . Abundant.

Margarites (Pupillaria) pupillus (Gould) . Common.

Littorina sitkana Philippi. Abundant.

Tectonatica clausa (Broderip and Sowerby) . Rare.

Tachyrhynchus erosum (Couthouy) . Common.

Thais lima (Gmelin) . Abundant.

Odostomia (Odostomia) n. sp. Rare. Comparable to O. cassandra

Bartsch, 1912.

Pelecypods

Mytilus edulis Linne. Abundant.

Pododesmus macroschismus (Deshayes) . Rare. On washed up

roots of kelp.

Clinocardium nuttalli (Conrad) . Abundant.

Tellina lutea Wood. Rare.

Gastrana irus (Hanley) . Common.

Macoma balthica (Linne). Common.

Macoma incongrua (von Martens) . Abundant.

66 nautilus Vol. 83 (2)

Macoma middendorfi Dall. Rare.

Siliqua alata (Dall) . Common.

Siliqua patula (Dixon) . Common.

Spisula polynyma (Stimpson) . Common.

Mya elegans (Eichwald) . Common.

My a japonica Jay. Common.

Mya priapus Tilesius. Common.

Hiatella arctica (Linne) . Common.

Zirfaea pilsbryi Lowe. Rare.

Literature cited

Corgan, J. X., 1966. Mya on the Alaska Penninsula: Nautilus, v.

80, p. 13-16.

Dall, W. H., 1921. Summary of the marine shell-bearing mollusks

of the northwest coast of America . . .: U. S. Nat. Mus. Bull.

112: 1-217, lbs. 1-22.

NOTES AND NEWS

The Bermuda Biological Station is now under the direction

of Dr. Wolfgang C. Sterrer, 29, graduate of the University of

Vienna and specialist in interstitial faunas of marine sands. Dr.

W. H. Sutcliffe, Jr., former Director, will now devote full time to

research at the Bedford Institute of Oceanography, Halifax, Nova

Scotia.

Johannes C. L. van der Riet. 1920-1969. We regret to announce

the untimely death of this ardent collector of Solomon Island

mollusks, on July 27 at the age of 49 in Tervuren, Belgium.

Gilbert Grau, 1906-1969 — We regret to announce the death of

our friend Gilbert Grau. He was born in Illinois, December 7,

1906, educated in music, and came to California many years ago,

the past thirty years arranging music for many outstanding motion

pictures and television. But we of the "Nautilus" world knew

Gilbert Grau as a student of the Pectinidae of the Eastern Pacific.

In 1959 he published this title in the Allan Hancock Pacific Series,

vol. 23, plates 1-59, University of Southern California Press. He

left his shell collection to the Smithsonian Institution. He was

killed in an automobile accident on May 29, 1969 — Mrs. John Q.

Burch, 1300 Mayfield Road, Apt. 61 L, Seal Beach, California

90740.

October, 1969 nautilus 67

Dr. George M. Davis will become Associate Curator of Mala-

cology at the Academy of Natural Sciences of Philadelphia (Nine-

teenth and the Parkway, Philadelphia, Pa. 19103) in the spring

or early summer of 1970. Dr. Robert Robertson is now Chairman

of the Department. Other members of the Department now in-

clude Virginia Orr Maes, Research Associate, and Nancy Wilson

Rulon, Scientific Assistant.

The American Malacological Union's 35th annual meeting was

held July 21-25, 1969 in Marinette, Wisconsin. 112 malacologists

were the guests of the University of Wisconsin-Green Bay, the

Marinette Area Chamber of Commerce and Margaret C. Teskey,

long-time AMU secretary. President Joseph Rosewater presided

over the meeting and introduced the following papers:

The molluscan fauna in North Carolina's Neuse River Estuary,

Hugh J. Porter. Mollusca of a cedar bog in Champaign County,

Ohio, Eugene P. Keferl. Donax fossor, a summer range extension

of Donax variabilis, Paul Chanley. Western Atlantic Donax, J. P.

E. Morrison. The Pleurocerid fauna of the Tennessee River,

Ralph M. Sinclair. Egg Cases of Nitidella ocellata Gmelin and an

Anachis, Dorothy Raeihle. Scandals in malacology, Morris K.

Jacobson. Patterns of oxygen consumption of the freshwater Pul-

monale snail, Lymnaea palustris, David G. S. Wright. Preliminary

report on the distribution of land snails in Northern Missouri,

Charles D. Miles and Richard L. Reeder. Some algae isolated from

Helisoma trivolvis (Say) , Richard L. Reeder and Robert G. Ander-

son. The Shells of Dioscorides of Anazarba, Kenneth Jay Boss.

Philopthalmus sp. (Trematoda) in Tarebia granifera and Mela-

noides tuberculatus in South Texas, Dr. Harold Murray and

Deborah Haines. Morphological-taxonomic notes on various fresh-

water Pulmonate snails, Harold J. Walter. (Read by title.) Prob-

lems in snail control in Hawaii, Henry van der Schalie. Problems

and techniques in experimental work with bivalve larvae, Ruth

D. Turner. Cabeza de Vaca, dealer in shells, James X. Corgan.

Histological studies of the Nephridium and Pericardial lining of

Qjiadrula nodulsta (Rafinesque) , Paul Robert Myers and Doro-

thea Franzen. Late Pleistocene nonmarine mollusks from Lake

Bretz, Lower Grand Coulee, Washington, James J. Landye. Some

possible consequences of a sea-level Panama Canal, Arthur H.

Clarke, Jr. Notes on Valvata tricarinata from Central Nebraska,

Carl W. Gugler. Contributions to the biology of New England

Nudibranch Molluscs, M. Patricia Morse. Malacology today, D. S.

Dundee. Some Techniques in studies of freshwater snails, Harold

68 nautilus Vol. 83 (2)

J. Walter. (Read by title.) Early American workers on the Naia-

des, William J. Clench. The earliest names for North American

Naiades, J. P. E. Morrison. Some basic problems in Naiad taxon-

omy, David H. Stansbery. The freshwater mussels of the Canadian

Interior Basin, Arthur H. Clarke, Jr. Where to find mussels in

creek-size streams, Herbert Athearn. Effects of pollution on the

Naiades of the Illinois River, William C. Starrett. Some research

needs and methods for protecting Naiades from extinction, Marc

J. Imlay. Seasonal variation in gonad activity in Unionid clams

and its systematic significance, William H. Heard. Functional

bilateral symmetry of the Lampsilis mantle: some problems, Louise

R. Kraemer. Studies on the structure and ultrastructure of the glo-

chidial stage of the Naiad Actinonaias ligamentina (Lamarck,

1819), Karen Heffelfinger. Life History of Pleurobema cor datum

(Rafinesque, 1820) , Paul Yokley. The life history of Truncatella

caribaeensis 'Sowerby' Reeve, Landon T. Ross. A dual behavioral

interpretation of a single environmental stimulus with freshwater

clams, Marc J. Imlay. Speciation and distribution of Arctic wedge

clams in the Western North Atlantic, John D. Davis. A revision

of the land snail genus Rabdotus in Texas, W. L. Pratt. Special

invertebrate staining techniques, Lucretia Buchanan. Paramya

stibovata, a commensal of the echiuroid Thalassema hartmani,

Charles E. Jenner and Anne B. McCrary. Changes in the Naiad

fauna of the Cumberland River at Cumberland Falls following

impoundment, David H. Stansbery. The University of Arizona

Marine Sciences Program, Albert R. Mead. Gonad development in

the three-ridge naiad, Amblema plicata (Say, 1817), Carol B. Stein.

Methods of subfamily recognition in Pacific Island endodontid

land snails, Alan Solem. The American mussel industry — eco-

nomic perspectives and ecological implications, John M. Bates.

The meeting followed the usual format: registration, papers

mornings and afternoons, shell club night, the annual dinner.

Acella haldemani was the featured mollusk; Harold Walter's beau-

tiful line drawings of the living snail and of the shell adorned the

program, and it obligingly turned up in limited quantity in the

small lake combed by 42 who attended the final day's field trip.

The 1970 meeting will be held in Key West, Florida, and the

following officers were elected to serve over the following year:

President, Alan Solem; Vice President, David H. Stansbery; 2nd

Vice President, G. Bruce Campbell; Secretary, Margaret C. Tes-

key; Treasurer, Mrs. H. B. Baker; Publications Editor, Morris K.

Jacobson; Councillors at Large, Dorothea Franzen, Dorothy

Raeihle, William E. Old, Jr., Gale G. Sphon, Jr.

October, 1969 nautilus 69

The effect of changing pH on the Unionidae. — Many

articles have appeared in the literature concerned with the calci-

phile and calciphobe characteristics of mollusks. In central New

York State, Clarke and Berg (1959) did not collect the larger

fresh-water bivalves (Unionidae) in waters with an alkalinity of

less than 47 ppm expressed as CaCO.r They stated that "Water

hardness or alkalinity varies greatly in natural waters of central

New York and exerts a strong influence on the distribution of

unionids."

Although there seems little doubt that the amount of calcium

present has profound effects on the vigor of unionid populations,

I question the absence of particular species being directly due

to lack of calcium.

On July 21, 1966, I collected mollusks in the headwaters of

Alder Creek near the town of Highmarket, Lewis County, New

York. Flowing over and between granite boulders, this stream did

not seem a likely place for mollusks. The rocks were covered with

diatomaceous algae and a few specimens of the vascular plant,

Myriophyllnm sp., were evident. The pH was 7.1; the total alkalin-

ity 47 ppm expressed as CaC03. A large population of Anondonta

cataracta was present. There was absolutely no soft substrate for

these animals to burrow in and many specimens were actually

wedged between the rocks, ventral side up.

On the 9th of August, 1966, I collected in Genegantslet Creek,

in Chenango County, near the town of Smithville Flats. This

rather large stream appeared fairly productive with sparce beds of

the aquatic vascular plants, Elodea sp. and Potamogeton sp. pres-

ent. The pH was 6.9; the alkalinity 28 ppm expressed as CaC03.

Associated with the pulmonate snail, Physa sayii, were the union-

ids, Alasmidonta undulata and Strophitus undulatus.

In the dystrophic Panther Lake in Oswego County, on July

19, 1966, I found 7 species of aquatic gastropods and dense popu-

lations of Elliptio complanata. The pH at that date was 7.1; the

total alkalinity 21 ppm expressed as CaCO.r

These few examples at least show that 4 species of unionids can

and do occur in some soft waters. Because of the negligible buffer-

ing abilities of waters low in calcium, rapid changes in pH are

practically always exhibited. It is probable that spasmodic fluc-

tuations of pH values are detrimental to fresh-water mollusks. In

70 nautilus Vol. 83 (2)

the three examples given the pH ranged only from 6.9 to 7.1. All

of these environments are in relatively undisturbed watersheds in

rather unpopulated areas. The local ecosystems are possibly stable

enough so that pH values remain fairly constant even in these

waters possessing poor buffering abilities.

These four species of Unionidae, representing two subfamilies,

may not be affected by natural variations in water hardness as

much as by the rapid changes in pH values usually associated with

this phenomenon. I hope that someone is, or will study these

relationships. Fresh-water bivalves are no longer present in many

streams where "nontoxic" chemical wastes are disposed of. It

seems probable that changes in pH may be at least partially re-

sponsible for their eradication. — Willard N. Harman.

Literature cited

Clarke, A. H., and C. O. Berg. 1959. The fresh- water mussels of

central New York. Cornell Univ. Agr. Exp. Stat. Memor.

367:1-79.

Omalogyra atomus (Philippi) from Maine. — Three speci-

mens of the minute, uncommon gastropod, Omalogyra atomus

(Philippi) were collected in tidepools at West Quoddy Head,

near Lubec, Maine. They were found in samples of algae, along

with the nest-building bivalve Musculus discors (L.) and the small

gastropods, Onoba aculeus (Gould) and Skeneopsis planorbis

(Fabricius) . Although O. atomus has rarely been mentioned in

American literature, it has been reported from Hampton Beach,

New Hampshire, and Newport and Narragansett Pier, Rhode

Island (Johnson, 1934. Proc. Boston Soc. Nat. Hist. 40(1): 100).

Whiteaves (1901. Catalogue of the Marine Invertebrata of Eastern

Canada, Ottawa, 272 pp.). Lermond (1909. Catalogue of the

Shells of Maine, 45 pp.) , and Procter (1933. Mount Desert Region

Survey, Part 5, Wistar Press, 402 pp.) make no reference to it.

Its occurrence in Maine not only constitutes a new record for that

state, but marks the first time specimens of this species have been

collected in over sixty years. O. atomus is much more abundant in

Europe and its functional morphology has been studied (Fretter

and Graham, 1962. British Prosobranch Molluscs, Ray Society,

755 pp.) . In addition, it has been reported from the east and west

coasts of Greenland (Thorson, 1944. Medd. om Gronland 121

October, 1969 nautilus 71

(13): 39, 148) and Iceland (Thorson, 1941. Zoology of Iceland

4 (60): 44) , where it has also been found living in algae with

5. planorbis and O. aculeus. Two of the three specimens collected

have been deposited in the Museum of Comparative Zoology,

Harvard University, Cambridge, Massachusetts. — Robert C. Bul-

lock, Museum of Comparative Zoology, Massachusetts.

Odostomia cassandra Bartsch. — Attempts to identify an Odos-

tomia from southwestern Alaska led to examination of the type

lot of Odostomia cassandra Bartsch, 1912. The conventional sub-

generic placement of the species is incorrect. A lectotype has never

been selected. The species was twice described as new. Authorship

of the name, date of publication, and place of publication are cur-

rently confused. An abbreviated synonymy is as follows:

Odostomia cassandra Bartsch, 1912

1912. O. (Evalea) cassandra Bartsch, Proc. U. S. Nat. Mus. 42:285,

pi. 38, fig. 5. (Skidegate, Queen Charlotte Islands, British

Columbia) .

1912. O. (Evalea) cassandra Bartsch, Bartsch, Proc. U. S. Nat. Mus.

42:341; 1921, Dall, Bull. 112 U. S. Nat. Mus.: 132.

1913. O. (Evalea) cassandra Dall and Bartsch, Victoria Mem. Mus.

Bull. 7:142, pi. 10, fig. 7.

Illustrations and descriptions in Bartsch (1912) and Dall and

Bartsch (1913) are virtually identical and are based on the same

two cotypes: USNM 220120 and a then unnumbered specimen in

the Geological Survey of Canada collection. Apparently this latter

specimen, now number 1610 in the National Museum of Canada

collection, was illustrated by Bartsch (1912). It is here selected

as the lectotype of O. cassandra Bartsch, 1912. Bartsch is clearly

author of the name, although authorship has sometimes been

attributed to Dall and Bartsch, 1913 (e.g.: MacGinitie, 1959,

p. 141).

Descriptions of O. cassandra stress an abundance of fine spiral

striations. This is the basis for assignment to the subgenus Evalea

A. Adams, 1860. As currently employed in North America, Evalea

is almost a meaningless name but it does separate some spirally

sculptured odostomids from other forms. Odostomia cassandra

should not be placed with sculptured species. Although it appears

sculptured, the surface is smooth. A sculptured appearance prob-

ably results from refraction within the thin, nearly transparent

shell. In the terms of Dall and Bartsch (1909) the species is an

72 nautilus Vol. 83 (2)

Odostomia (Odostomia).

I am indebted to Dr. Arthur H. Clarke, National Museum of

Canada, and to Dr. Joseph Rosewater, U. S. National Museum,

for access to specimens. — James X. Corgan, Austin Peay State

University, Clarksville, Tennessee, 37040.

Literature cited

Bartsch, P., 1912, Additions to the West American pyramidellid

mollusk fauna with descriptions of new species. Proc. U. S. Nat.

Mus. 42:269, pits. 35-38.

Dall, W. H., and P. Bartsch, 1909, A monograph of West Ameri-

can pyramidellid mollusks. U. S. Nat. Mus. Bull. 68: 1-258,

pits. 1-30.

, and . , 1913, New species of mollusks from the Atlantic

and Pacific coasts of Canada. Victoria Mem. Mus. Bull. 2:139-

146, pi. 10.

Mac Ginitie, N., 1959, Marine Mollusca at Point Barrow, Alaska.

Proc. U. S. Nat. Mus. 709:59-208, pis. 1-27.

Melanoides tuberculata (Muller) in Florida. — Through

the kindness of Mrs. L. E. Crovo of Miami and Mr. A. K. Anders

of Kissimmee we have received recently three lots of the intro-

duced Oriental snail, Melanoides tuberculata (Muller) . One lot,

collected in September 1966 from Lake Osceola, Main Campus,

University of Miami, Coral Gables and a second lot, collected in

November 1968 in Greynolds Park, North Miami. The third lot,

received from Mr. Anders, was collected in Hillsborough State

Park, Hillsborough County. So far as I know these are the first

records from Florida. Dr. Harold D. Murray has reported upon

the occurrence of this species in San Antonio, Bexar County, and

at New Braunfels, Comal County, Texas. (American Malacologi-

cal Union Annual Report 1964, p. 15 and Annual Report 1965,

p. 25) .

The two lots from Miami probably represent two separate intro-

ductions as they appear to be rather distinctive populations. The

series from Lake Osceola is proportionately a much stouter race

than the specimens from Greynolds Park. The morphological

differences between these two populations are well within the

variations exhibited by any large geographic series. M. tuberculata

is wide ranging, extending from north and east Africa east through

Saudi Arabia, Iran, Pakistan, India and southern China, then

southeast Asia into Indonesia as far east as Java and the Celebes.

— W. J. Clench

October, 1969 nautilus 73

The Physa Striata Complex. Three different species of P.

striata have been named. All three names have disappeared be-

cause of being synonyms or homonyms. Two of these names per-

tain to the Physidae and one to the Planorbidae. The following is

a resume of this complex.

Physidae

Physa heterostropha (Say)

Lymnaea heterostropha Say 1817, Nicholson's Encyclopedia,

article on Conchology, Vol. 2, no pagination, pi. 1, fig. 6 (Dela-

ware River) .

Physa striata Menke 1828, Synopsis Methodica Molluscorum,

Pyrmont, p. 79 (near Goshen, Massachusetts) [is Physa hetero-

stropha (Say) ].

Physa virgata Gould

Physa virgata Gould 1855, Proc. Boston Soc. Nat. Hist. 5:128

(River Gila [Arizona], and near San Diego [California]) .

Physa striata Lea 1864, Proc. Acad. Nat. Sci., Philadelphia, p.

115 (Salt Lagoon, near Monterey, California).

Physa dorbigniana Lea 1866, Jour. Acad. Nat. Sci., Philadelphia

<5:166, pi. 24, fig. 85; 1867, Observations on the Genus Unio

77:122, pi. 24, fig. 85. [New name for Physa striata Lea 1864, non

P. striata d'Orbigny 1841]. [Is Physa virgata Gould, Pilsbry 1911,

Proc. Acad. Nat. Sci., Philadelphia, p. 198].

Planorbidae

Plesiophysa pilsbryi Aguayo

Physa striata d'Orbigny 1841, [in] de la Sagra, Histoire Physique,

Politique et Naturelle de L'lle de Cuba, Mollusques 7:192, pi. 13,

fig. 14-16 (Martinique or Cuba [is Martinique]) .

Plesiophysa pilsbryi Aguayo 1935, Mem Soc. Cubana Hist. Nat.

9:121. [New name for Pfiysa striata d'Orbigny 1841, non P. striata

Menke 1828]. — W. J. Clench, Cambridge, Mass.

74 nautilus Vol. 83 (2)

Polydora infestation of Mercenaria mercenaria. — There

have been very few reports of Polydora worm infestation of Mer-

cenaria mercenaria probably because the quahog usually resides

deep enough in the substratum to avoid attack by this marine

polychaete. Landers (1967) did induce infestation of M. mer-

cenaria by Polydora ciliata in the laboratory but only in the

absence of substratum and on young specimens. Thus, it is signif-

icant to report a natural infestation of M. mercenaria brought to

my attention by Mr. Clinton Andrews, resident supervisor at the

Nantucket Research Center of the University of Massachusetts,

Nantucket Island, Massachusetts.

During November 1967, scallop dredging operations were car-

ried on in Nantucket Harbor. Several quahogs were also collected

showing extensive shell damage, and some were subsequently sent

to me by Mr. Andrews. External coloration indicated that the

posterior third of each specimen had been exposed above the

substratum. The valves had been exposed unevenly, one more

than the other, and in each case the more exposed valve exhibited

an extensive network (in the exposed part of the valve) of tubular

excavations characteristic of Polydora infestation. However, I

found no remains of this polychaete within the excavations.

Apparently, the quahogs were visually not embedded vertically in

the substratum but were tipped to one side, the infestation occur-

ring only on the upper side.

Quahogs are usually buried sufficiently to escape a scallop

dredge, but presumably recent storms had caused substratum

dislocation in the shallow harbor, and some quahogs were thus

exposed. All specimens of M. mercenaria recovered were of good

size suggesting that larger specimens may be less mobile and much

slower in re-entering the substratum. Thus exposed, they were

infested by Polydora sp.

Infestation perforations penetrated the inner surface of one

specimen within the posterior adductor muscle scar, but no exten-

sive damage appeared to have resulted. The specimens were alive

when collected. John D. Davis, Smith College, Northampton,

Massachusetts.

Reference cited

Landers, W. S. 1967. Infestation of the hard clam, Mercenaria

mercenaria by the boring polychaete worm, Polydora ciliata.

Proc. Nat. Shellfisheries Assoc. 57:63-66.

October, 1969 nautilus 75

Achatina fulica Inyades Florida — In 1966, a boy brought

home from Hawaii specimens of this Giant African Snail in his

luggage as pets. They and their offspring were released in the

neighborhood gardens of North Miami, and now, by September

1969, have become a horticultural nuisance in a 13-square block

area surrounding North Miami Avenue and N.W. 125th St. The

Florida State Agricultural Control officials are attempting to con-

fine and eliminate the infestation. — R. Tucker Abbott

JOHNSONIA: VOLUME 1

Long out of print, this useful monograph series on Western

Atlantic Conidae, Muricidae, Strombidae, Cardiidae, etc., is now

once again available. Reproduced in its original size, original

paper, and beautifully soft-bound, it may be obtained for only

$15.00 (postage and packing free) from W. and R. McCauley,

1919 Sandy Hill Road, Apt. C-12, Norristown, Pa. 19401. If you

also wish vols. 2, 3 and 4, write: Dr. W. J. Clench, Museum of

Comparative Zoology, Cambridge, Mass. 02138, U.S.A.

PUBLICATIONS RECEIVED

Powell, A. W. Baden. Sept. 9, 1969. The Subfamily Turriculinae,

Part 2 of The Family Turridae in the Indo-Pacific. Indo-Pacific

Mollusca, vol. 2, no. 10, pp. 203-416, 136 pis., 1 in color. $10.55.

This journal is now published by the Delaware Museum of

Natural History, Greenville, Delaware 19807.

Dall, William Healey, 1845-1927. The Zoological Taxa of William

Healey Dall. By K. J. Boss, Joseph Rosewater and Florence A.

Ruhoff. Bull. 287, U.S. National Museum, 427 pp. 1968. $2.50

by writing Superintendent of Documents, U.S. Gov. Printing

Office, Wash., D.C. 20402. An extremely useful and much need-

ed bibliographic work that is well done.

Fretter, Vera (editor). 1968. Studies in the Structure, Physiology

and Ecology of Molluscs. Symposia of the Zool. Soc. London.

Academic Press, N. Y. 377 pp., illus. Contains 18 excellent re-

search articles on marine, land and fresh-water mollusks, includ-

ing "A review of the bivalved gastropods and a discussion of

evolution within the Sacoglossa" by E. Alison Kay; "The biology

of interstitial [marine sand] Mollusca" by B. Swedmark; "The

Feeding Mechanism and behaviour of the opisthobranch Melibe

leonina" by Anne Hurst; "The burrowing activities of bivalves"

by E. R. Trueman; and "Habitat distribution of Solomon Island

land Mollusca" by John F. Peake.

76 nautilus Vol. 83 (2)

Nowell-Usticke, G. W. 1969. A Supplementary Listing of New

Shells. 32 pp., 6 pis. Privately printed. Price unknown.

This booklet was intended to be a supplement to Mr. Usticke's

somewhat controversial publication on the shells of St. Croix

(1959) . It is actually a summary of the author's contribution to

West Indian marine conchology and represents a serious problem

for those interested in the systematics of Caribbean mollusks. Most

taxonomists who describe new taxa make careful literature sur-

veys, study comparative material, and often try to understand the

mechanisms of speciation. Mr. Usticke, on the contrary, intro-

duces names for atypical specimens and for species which he does

not understand. In this present copyrighted "small listing," seventy

species, subspecies, forms, and varieties are described as new,

including eleven "new" Conus which he had previously described

(Nowell-Usticke, 1968; see review by Dr. R. Tucker Abbott, 1969,

Nautilus 82: 147) .

It is unfortunate that Mr. Usticke did not consult authorities

on West Indian marine mollusks, for if he had, many of the

numerous misspellings, taxonomic errors, homonyms and syno-

nyms which he has created could have been avoided. The following

examples serve to illustrate some of the errors and inconsistencies

which characterize this work: Mitra pallida Usticke, 1959 [non A.

Adams, 1851; non Pease, 1860] is listed (p. 19) as a subspecies of

M. nodulosa, yet the description begins "May be granulosa La-

marck;" Vasum aldridgei, n. sp. (p. 18) , is the muricid, Muricop-

sis philippiana Dall; Shuster and Bode (1961. Nautilus 75: 1-7)

discussed the variation of Vasum globulus nuttingi Henderson,

yet Mr. Usticke has described an additional subspecies (p. 19)

from the same general locality; Anachis multicosta, n. sp. is intro-

duced (p. 16) with the comment "Possibly pulchella Kiener;"

the author of Fenimorea ustickei Hayes is listed as "Hayes-

Usticke," leading one to wonder who actually described the spe-

cies; Triphora novem is inadequately described (p. 12) as new,

with reference made to the figure of the lectotype of T. intermedia

(C. B. Adams) ; Ischnochiton dubium, n. sp. (p. 7) , appears to be

I. bermudensis Dall and Bartsch; Brachidontes exustus var. rosa-

ceus, n. s. sp. [sic] and B. pseudorecurvus, n. sp. (p. 5) , are both

examples of the common Caribbean B. cubitus (Say) . Ringiculina

[Ringicula on pi. VI] cruzensis new sp. (p. 30) has a single denticle

October, 1969 nautilus iii

on the outer lip; to be the shell of a cephalaspid opisthobranch it

would need to lack this; many cephalaspids have columellar folds,

which Usticke's larval shell lacks; his new species is a sinusigera

of some prosobranch. Mr. Usticke continually confuses the terms

"species," "subspecies," "form," and "variety."

Abbott (1959, Nautilus 82: 147) raised the question about the

validity of this type of publication. Except for the obvious viola-

tion of Article 15 of the International Code of Zoological Nomen-

clature which eliminates the availability of approximately one

third of the proposed new names, Mr. Usticke has not directly

violated any other rule. He does, however, display his ignorance

or rejection of the principles of the Code by violating a majority

of the applicable Recommendations. Most important among the

latter are his failure to designate type specimens, deposit them in

an institution, compare his new taxa with related ones, give

adequate data concerning the types, and follow recommended spell-

ing procedures. It appears that the neglect of so many recom-

mendations and the pronounced unscientific attitude of this con-

chologist are possible grounds for declaring all three of his booklets

invalid. Works of this sort not only promote instability of mala-

cological nomenclature, but do much to hinder its progress.

The photographs are for the most part, good and, in some cases,

the only means of interpreting the inadequate descriptions.

Literature cited

Nowell-Usticke, G. W. 1959. A Check List of the Marine Shells

of St. Croix. 90 pp., 4 pis. Lane Press, Burlington.

Nowell-Usticke, G. W. 1968. Caribbean Cones from St. Croix and

the Lesser Antilles. 31 pp., 4 pis. Livingston Publ. Co., Nar-

berth, Pa.

— Robert C. Bullock, Museum of Comparative Zoology,

Cambridge, Mass.

Dance, S. Peter. 1968. Rare Shells. Univ. California Press, Berke-

ley, 128 pp., 24 pis. in color. $17.00 Foreword by R. Tucker

Abbott. Good photographs and interesting historical account of

50 presently or formerly rare shells are included.

iv NAUTILUS Vol. 83 (2)

Something New in Conchology

A professional recording of the pronunciations of 1,200

scientific species and generic names of

"Seashells of North America"

also how to enunciate — authors' names, family names, and

dozens of technical names, such as

apophysis, conchiolin, eolis, and

veliger.

Produced and published by R. Tucker Abbott

Sponsored by the Delaware Museum of Natural History

Greenville, Delaware 19807

This is a high-fidelity, long-playing (33 V3 RPM) recording

of the names and terms found in Dr. Abbott's new and

popular book, Seashells of North America (Golden Press,

N.Y.). Born in Boston, bred in Canada and a graduate of

Harvard College, Dr. Abbott has blended the customary

pronunciations of the top professional malacologists in the

United States and Canada that set a high standard of

acceptability. All names and terms are printed on the record

jacket. A unique and long-lasting aid to your study of shells

and biology, for only $4.95 (postage paid).

Obtainable from your favorite shell dealer

WILLIAM H. WEEKS SHELL COLLECTION: New price lists

of this famous collection, with full scientific data, are in prepa-

ration. Many new additions of fine and rare species are also

included. To obtain free copies write:

George E. Jacobs, 853 Riverside Drive, New York 32, N. Y.

WORLD WIDE SPECIMEN SHELLS for sale. New 1969

Price List on request. "Illustrated Catalog of Popular Mar-

ginella Species" now available also, 117 species shown with

full data and values listed with cover in color. Price $2.00.

PHILLIP W. CLOVER

Box 33, Div. 32

FPO, New York 09540

Vol. 83 JANUARY, 1970 No. 3

THK

NAUTILUS

THE PILSBRY QUARTERLY

DEVOTED TO THE INTERESTS OF CONCHOLOGISTS

EDITORS AND PUBLISHERS

Horace Burrington Baker, 11 Chelten Road, Havertown, Pa.

(Emeritus Professor of Zoology, University of Pennsylvania)

R. Tucker Abbott, duPont Chair of Malacology

Delaware Museum of Natural Historv, Greenville, Del. 19807

Charles B. Wurtz, Biology Department

La Salle College, Philadelphia, Pa. 19141

CONTENTS

Mode of life of Gonocardium, a Paleozoic pelecypod.

By David Nicol \ rv 77

The distribution of the nudibranch, Doris verrucosa Linne,

in the northwest Atlantic. By David R. Franz 80

Cymatium boschi, a new species from the Arabian Sea.

By R. T. Abbott and Hal Lewis 86

Aspella rnyrakeenae , new species from Western Mexico.

By W. K. Emerson and A. D'Attilio 88

New Mitridae and Volutomitridae. By Walter O. Cernohorsky 95

Rediscovery of the limpet, Acroloxus coloradensis,

in Colorado. By George W. Bryce, Jr 105

Longevity in Ashmunella, Moyiadenia and Sonorella.

By Munroe L. Walton 109

Notes and news 112 Publications received ... iii

$4.25 per year ($4.75 to Foreign Countries) $1.25 a copy.

Mrs. Horace B. Baker, Business Manager

11 Chelten Road, Havertown, Pennsylvania 19083

Second-Class Postage paid at Spring House, Pa.

NAUTILUS:

A Quarterly Journal devoted to the study of Mollusks, edited and published

by Horace B. Baker, Charles B. Wurtz and R. Tucker Abbott.

AUTHORS PLEASE NOTE:

Matter for publication should be sent to the acting editor. Dr. R. Tucker

Abbott, Delaware Museum of Natural History, Greenville, Delaware 19807.

Manuscripts should be typewritten and DOUBLE SPACED throughout, in-

cluding titles, footnotes and bibliographies. Please give reprint order, if any.

Please correct and return galley proof (submitted without manuscript)

quickly. Changes from manuscript, as edited, may be disregarded.

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THE NAUTILUS

Vol. 83 January, 1970 No. 3

MODE OF LIFE OF CONOCARDIUM, A PALEOZOIC

PELECYPOD

By DAVID NICOL

Department of Geology, University of Florida

Gainesville, Florida 32601

One of the most extraordinary pelecypods that ever lived was a

small family of bivalves, the Conocardiidae, typified by Conocar-

dium. Vokes (1967) , in Ins catalogue of pelecypod genera, places

this group in a separate order (Conocardioida) under the subclass

Cryptodonta. The only living family belonging to the Cryptodonta,

according to Vokes, is the Solemyidae. This allocation of the

conocardiids, as Vokes admits, can be considered only an educated

guess, and a few paleontologists have gone so far as to doubt that

the conocardiids were pelecypods or even molluscs. It is my con-

tention that these bivalved animals were molluscs and very likely

pelecypods. Therefore, I am interpreting the life habit of the

conocardiids on the assumption that they were aberrant pelecypods.

The conocardiids were among the most ancient basic stocks of

pelecypods, appearing in the Early Ordovician. The heyday of

the conocardiids seems to have been the Devonian and Mississip-

pian, although they were still common in many Pennsylvanian

faunaus. There is a clear indication of further decline of this group

in Permian marine faunas, and the species of this geologic period

are commonly of small size. There is considerable doubt that the

conocardiids survived to the very end of the Permian, and so all

reports of occurrence of conocardiids in Triassic rocks are false.

To begin with, the orientation (as to which is anterior and

which is posterior) , has long been in dispute, with opinion rather

evenly divided. There are two gapes in the shell, one at the end of

a tubular process and a much larger gape near the other end of

the shell. The tubular process varies greatly in length from almost

nonexistent to longer than the remainder of the shell. In appear-

ance it reminds one of the rostrum or tubular process of Cuspidaria.

I believe that the tubular process had to be at the posterior end

and probably contained the exhalant and possibly the inhalant

77

78 nautilus Vol. 83 (3)

siphons. The larger gape, then, would be for the foot and is

located either on the ventral margin in about the same position

as in Area, sensu stricto, or somewhat farther anteriorly near the

an tero- ventral margin (compare figs. 1 and 2) . In size and shape

but not in position, the gape in Conocardium is much like that of

Tridacna crocea Lamarck (fig. 5) . The ornamentation of Cono-

cardium also resembles that of some species of Area, sensu stricto,

(figs. 3 and 4) .

To further complicate matters, some (but most certainly not all)

species of Conocardium had a shelly structure on the posterior side

of the valves that La Rocque (1950, p. 317) called a hood. The

hood was a scooplike shelly extension that began at about the

middle of the exterior of the valves and extended posteriorly. The

possible function of this structure is unknown to me.

Another puzzling feature noticed by Chronic (1953, p. 154, in

Newell, Chronic, and Roberts) in a species of Permian Cono-

cardium from Peru is that a part of the valve margins appears to

be fused. I noticed this same condition in some well preserved

specimens from the Pennsylvanian of Texas. This would probably

make the animals nearly or completely immobile in the adult stage

because the gape for the foot near the anterior end is not large

enough to permit much movement. Most fossil pelecypods are

found with the valves separated, but most conocardiids are found

with the valves still joined, and this also adds some credence to

the observation that the valve margins may have been fused.

I believe that Conocardium was attached by a stout short byssus

to the subtrate; it was an epifaunal, not an infaunal pelecypod.

In most species of Conocardium, the size, shape, and position of

the gape is similar to Area, sensu stricto. Most species of Cono-

cardium probably lived in an attached position like Area with

beaks and dorsal side up, the valves perpendicular to the sub-

strate. In some species of Conocardium, the byssal gape is farther

forward in an antero-ventral position so that the animal would

have the posterior side somewhat tilted upward in the attached

position. Hind (1900, pp. 452-453) assumed that the large gape in

the valves of Conocardium was for the passage of a stout byssus.

My reasons for believing that Conocardium was byssally at-

tached are as follows: Conocardium had a very thick shell; modern

burrowers and borers have thin, or relatively thin, shells. The

January, 1970

NAUTILUS

79

Fig. 1. Conocardium sp., ventral view of both halves showing the byssal

gape. Fig. 2. Area ventricosa Lamarck, ventral view of both valves showing

the byssal gape. Fig. 3. Conocardium sp., exterior view of a right valve.

Fig. 4. Area ventricosa Lamarck, exterior view of a left valve. Fig. 5. Tridacna

crocea Lamarck, ventral view of both valves showing the byssal gape.

valves of Conocardium are gibbous or highly convex. Living pele-

cypods of great convexity are poor and shallow burrowers, if they

burrow at all. The anterior side of most species of Conocardium is

strongly alate or winged. The anterior margin comes to a point at

the antero-dorsal border. Many byssally attached forms have an

anterior wing-like projection, as for example Pteria; but the an-

terior side of burrowers and borers is commonly gently rounded

80 nautilus Vol. 83 (3)

or even truncated. If the valves were fused in the adult stage of

some species, how could burrowing or boring be accomplished

easily? Moreover, the gape is not as large or round as it is in

modern pholads, which use a sucker-like action of the large foot

to aid in boring.

The only weakness to my argument that I can see is why did

these animals have a posterior tube or rostrum? There is no adap-

tive significance to this structure for a byssally attached pelecypod.

Perhaps there was no adaptive significance to the posterior tube,

but we know almost nothing about the anatomy of the animal.

Furthermore, what is the adaptive significance for the hood which

occurs in some species of conocardiids? The difficulty in interpret-

ing the mode of life of Conocardium is that even its external form

is so unlike any other pelecypod, living or extinct, that compari-

sons are exceedingly difficult to make.

I am convinced, however, that on the basis of the overall ap-

pearance of the shell, Conocardium was a byssally-attached epi-

faunal pelecypod. The likelihood that Conocardium was an in-

faunal burrower or borer is indeed remote.

I am indebted to Miss Marsha E. Jessup of the U. S. National

Museum for making the drawings for this paper.

Literature cited

Hind, W., 1900. A monograph of the British Carboniferous

Lamellibranchiata. Part V. Coelonotidae, Solenomyidae, Cono-

cardiidae, Cardiidae: Palaeontographical Soc. London, Vol. for

1900, pp. 361-436, pis. XL-LIV.

La Rocque, A., 1950. Pre-Traverse Devonian pelecypods of Michi-

gan: Mus. Paleo., Univ. Michigan, 7(10) : pp. 271-366.

Newell, N. D., J. Chronic, and T. G. Roberts, 1953. Upper Paleo-

zoic of Peru: Geol. Soc. Am. Mem. 58, 276 pp.

Vokes, H. E., 1967. Genera of the Bivalvia: A systematic and bibli-

ographic catalogue: Bull. Am. Paleo., 51 (232) : pp. 111-394.

THE DISTRIBUTION OF THE NUDIBRANCH DORIS

VERRUCOSA LINNE IN THE NORTHWEST ATLANTIC1

By DAVID R. FRANZ

Systematic & Environmental Biology, Biological Sciences

University of Connecticut, Storrs, Connecticut 06268

The nudibranch Doris verrucosa Linne is a warm-water

1 Contribution No. 61 of the Marine Research Laboratory, University of

Connecticut.

January, 1970 nautilus 81

amphi-atlantic species which is distributed, in the Eastern Atlantic,

from the coast of England and Ireland, the Mediterranean, the

Adriatic and the west coast of Africa. Prior to the new distribu-

tional records noted below, the species in the West Atlantic was

reported from Sao Sebastiao, Brazil (Marcus, 1955) , the west coast

of Florida (Bergh, 1894), South Carolina (in Eliot, 1910, p. 96)

and Sapelo Island, Georgia (Marcus & Marcus, 1967) . The pur-

pose of this report is to note the occurrence and confirm the identi-

fication of Doris verrucosa on the Atlantic coast of North America

northward to Buzzards Bay, Massachusetts. The new records re-

ported here are based on collections from Beaufort, North Carolina,

the York River (Chesapeake Bay, Virginia) , the mouth of the

Mystic River, Connecticut, and Sugar Loaf Spring, Buzzards Bay,

Massachusetts. The Beaufort specimens were graciously provided

bv Mrs. Ann McCrary, University of North Carolina, and the York

River specimens by Dr. Alex Marsh, Virginia Institute of Marine

Sciences, Gloucester Point, Va. The preserved Connecticut slugs

were collected by Dr. Robert Cerwonka, Potsdam State College,

Potsdam, New York and the single living animal from the Mystic

River was collected by Mr. Gordon Hendler of the University of

Connecticut. The Buzzards Bay specimens, collected in 1902 by

U. N. Edwards, came from the collection of the Academy of

Natural Sciences of Philadelphia (ANSP No. 85618) and the

author is indebted to Dr. Robert Robertson and Kaniaulono

Bailey for the loan of this material. It is a pleasure to acknowledge

the generosity of all of these workers.

Since the records reported here represent a significant northward

extension of the known range of this species, selected descriptive

and anatomical data supporting the identification are provided.

Doris verrucosa Linne\ 1758

Altogether, 13 West Atlantic specimens have been examined,

4 each from Beaufort and Virginia, 3 from Connecticut and 2

from Buzzards Bay. The smallest animal (preserved) was 7.5 mm.

in length and 5.3 mm. wide. Other specimens ranged upward to a

maximum of 45 mm. by 28 mm. for the living slug from the Mystic

River (Fig. 2) . The length varies from 1.4 to 1.7 times the width

among the preserved animals. The preserved slugs are opaque

white, with a slight hint of yellow in one specimen. The living

animal was a dingy orange color, somewhat paler ventrally and

82 nautilus Vol. 83 (3)

on the dorsal surface of the foot (which was visable when the

animal moved about) . The notal papillae are moderately dense

but do not touch each other. The larger are rounded, the smaller

either rounded or conical. The oral tentacles are smooth and not

well developed. The rhinophores are perfoliate and the rhino-

phoral pits are surrounded by a pair of large papillae, one on

each side. In some specimens, a smaller papilla may occur on the

posterior and/or the anterior margin of the pit. The branchial pit

is surrounded by a mixture of larger and smaller papillae. There

are 12 to 15 unipinnate branchiae.

The labial cuticle is smooth and colorless. In the four specimens

dissected, the radula formulae were as follows: (25-26) -0- (25-26) ,

(31-35) -0- (31-35) , (35-36) -0- (35-36) , (27-47) -0- (27-47) . The total

number of rows ranged from 24 to 42. The teeth are hamate, but

variable in size and appearance depending on position within a

row (Fig. 1) . The extreme inner and outer teeth are smallest and

the outer two or three may be rudimentary.

The ampulla of the hermaphroditic gonoduct is thick, with a

muscular coat, and enters the anterior gland mass far forward and

ventrally. The bifurcation of the hermaphroditic gonoduct into

male and female systems occurs within the anterior gland mass.

The prostatic portion of the vas deferens leaves the gland mass

anteriorly near the point of confluence with the ampulla and con-

tinues along the anterior margin of the gland mass for a short

distance before diverging from it. The sharp division between

prostatic and muscular portions of the vas deferens is marked by

a 1 80 degree reflection and at this point, a penial retractor muscle

inserts on the vas deferens. Distally, the vas deferens is visably

coiled within its muscular sheath. The vagina begins in the copula-

tory atrium and joins the duct of the spermatheca near its origin.

Just distal to this, the convoluted uterine duct and the duct of the

spermatocyst diverge, the former entering the anterior gland mass.

A non-glandular oviduct connecting the gland mass and the atrium

of oviposition is not present. The relationship of the various parts

of the anterior reproductive complex are shown diagrammatically

in Fig. 3.

The living animal from Beebe Cove was collected sublittorally

from a drifted frond of Lnminaria in very shallow water (water

temp. 24.5°C at bottom) .

January, 1970 nautilus 83

Discussion

These specimens differ only slightly from Brazilian animals

described by Marcus (1955), and specimens from Sapelo Island,

Georgia, reported by Marcus and Marcus (1967) . In a recent

paper, Schmekel (1968) has drawn a distinction between D. ver-

rucosa and the very similar but smaller species D. ocelligera Bergh,

1881, based predominantly on size relationships, the size and shape

of the mantle papillae, the size and number of rhinophoral and

branchial papillae and, internally, the absence of a penial retractor

(ocelligera). The general body proportions (length less than twice

the width) , the existence in the present animals of both small and

large branchial papillae and a penial retractor, all of which are

absent in D. ocelligera, eliminate the latter species. Doris verrucosa

must also be compared with D. bovena Marcus, 1955. In this

species, however, the rhinophoral pits are guarded by 8-10 papillae

whereas the branchial margin is smooth. Furthermore, bifurcation

of the hermaphroditic gonoduct occurs outside of the anterior

gland mass.

In her recent discussion of D. verrucosa from the Gulf of Naples,

Schmekel (1968) emphasizes the importance of the height of the

larger notal papillae, some of which in the figure provided, are

about twice as high as wide. However, these elongate papillae were

not present in two of three specimens from the Gulf of Naples

examined by the author (ANSP 84454) . In comparing the maxi-

mum diameter and density of notal papillae of all specimens in-

cluding the three Naples animals noted above it is evident that

these characters are very variable and tend to form a continuous

intergradation which does not appear to be correlated either with

geography or size of animal.

In all other aspects of internal and external anatomy, the West

Atlantic specimens are very similar those from the Gulf of Naples.

Zoogeography: The relatively large size of the Connecticut slugs

suggests that these were older animals which had survived at least

one winter in Beebe Cove. Winter conditions in the cove are

severe, and temperatures may remain below 5°C for extended

periods. Frequently an ice sheet covers most of the cove and may

persist several weeks, depending on weather conditions. Two of the

four York River specimens were collected in December suggesting

the possibility that the species is also permanent resident in Chesa-

84

NAUTILUS

Vol. 83 (3)

34

29

17

Fig. 1. Radula of Doris verrucosa. Teeth are numbered beginning medially

;ind increasing laterally. No's 1 and 17 are lateral views, 29 to 34 frontal views.

Line equals 0.1 mm.

peake Bay. In Beebe Cove, intensive collecting by the author and

his students during the summers of 1967 and 1968 failed to turn

up a single specimen. Since the single animal from the cove col-

lected in July of 1969 was in the process of oviposition, the thermal

requirements for reproduction were obviously present at that time.

However, it is possible that successful recruitment of D. verrucosa

in New England is dependent on the seasonal incursion of larvae

from the Gulf Stream. The influence of the Gulf Stream is com-

monly indicated by the presence of Sargassum in Block Island

and Fishers Island Sounds. In any case, the shallow waters of the

Mystic River as well as other local areas in southern New England

do attain summer temperatures high enough to provide the thermal

requirements for the survival and reproduction of this species.

D. verrucosa is the only warm-water amphiatlantic nudibranch

known to extend north of Cape Hatteras, N. C.

References cited

Bergh, R., 1894. Die Opisthobranchien. Reports on the dredging

operations ... by the U. S. Fish Commission Steamer "Alba-

tross." Bull. Mus. Comp. Zool. Harvard Univ. 25 (10) : 125-233.

Eliot, C, 1910. The British Nudibranchiate Mollusca, Part VIII

(Supple.) Ray Society, London.

January, 1970

NAUTILUS

85

Marcus, E., 1955. Opisthobranchia from Brazil. Univ. Sao Paulo,

Bol. 207: 89-200.

Marcus, E. and E. Marcus, 19f>7. Some opisthobranchs of Sapelo

Island, Georgia, U.S.A. Malacologia 5(1-2): 199-222.

Schmekel, L., 1968. Die Gattung Doris (Gastropoda, Nudi-

branchia) im Golf von Neapel. Publ. Staz. Zool. Napoli 36 (2) :

167-197.

Fig. 2. Doris verrucosa Linne collected from the mouth of the Mystic River

(Beebe Cove) Connecticut in July, 1969. Maximum length of the living

animal was 45 mm.

MVd

Fig. 3. Anterior genital complex of Doris verrucosa (diagrammatic) .

Am, ampulla; GM, anterior genital mass; MVd, muscular vas deferens;

Pr, prostatic vas deferens; Spc, spermatocyst; Spt, spermatheca; UD, uterine

duct; Vg, vagina.

8(i

NAUTILUS

Vol. 83 (3)

CYMATIUM BOSCHI, NEW SPECIES FROM

THE ARABIAN SEA

By R. TUCKER ABBOTT1 and HAL LEWIS-

Cymatium (Ranularia) boschi new species. Figs. 1-4 holotype. Figs. 5 and

paratype, A.M.N.H. no. 13920(5.

1 Delaware Museum of Natural History, Greenville, Del. 19807.

'-' Wynnewood, Pennsylvania.

January, 1970 nautilus 87

Dr. Donald Bosch, a physician at the American Mission Hospi-

tal in Muscat, Oman, recently sent us a Cymatium for identifica-

tion. It appears to be undescribed, and in our careful search of the

literature we could find only one illustration of it (fig. 0, pi. 10,

second edition, 1757, Argenville's L'Histoire Naturelle . . . Con-

chyliologie) which, to our knowledge, has never had a binomial

name applied to it. The new species is in the subgenus Ranularia

Schumacher, 1817 (type by Herrmannsen's 1847, p. 388 designa-

tion: Tritonium clavator Chemnitz, 1780, which is now gutturniurn

(Roding, 1787)).

Cymatium (Ranularia) boschi new species Figs- 1"6

Description: Shell reaching about 78 mm. (almost 3 inches) in

length, subtrigonal, spirally corded, widi tabulate whorls, and

colored whitish with a suffusion of soft light-brown. Nuclear

whorls (total number unknown) , fragile, smooth, horn-brown.

Postnuclear whorls 4; rather squarish or tabulate in the spire, the

shoulder being either squarely or roundly angulate. Suture mi-

nutely impressed. Below it, the top of the whorl is flattish and with

9 or 10 retractively slanting, narrow, small, axial folds or ridges.

Body whorl with 11 or 12 spiral cords, the upper one or two bear-

ing 10 or 11 small nodes. The cords below this becoming increas-

ingly small and smooth. Siphonal canal relatively broad and short.

Aperture elongate, broad at both ends. Outer lip thick, with 3 to 5

poorly developed broad clumps of twin lirae or teeth. Parietal wall

raised, white-glazed, glossy, with weak, short spiral lirae well within

the aperture. Outer lip with a large, rounded, nodular varix. Up-

per or posterior end of aperture broadly V-shaped and not bounded

by large teedi. Periostracum light-brown, moderately thick over

the entire shell, deciduous when dry, axial fimbriated in short,

thin blades. Soft parts and operculum unknown.

Measurements (mm.):

Holotype; Delaware Mus. Nat. Hist.

Figured paratype AM NH 139206

Paratype. Donald Bosch collection

Paratype; ANSP 188844

Types: The type locality is Al Masirah Island, Oman Protec-

88 nautilus Vol. 83 (3)

torate, southeast Arabia. The holotype is in the Delaware Museum

of Natural History (Greenville, Delaware) no. 22862. Figured

paratype and 3 worn and partially broken paratypes in the Ameri-

can Museum of Natural History, no. 139206. One paratype in the

Donald Bosch private collection. One paratype from Muscat, Gulf

of Oman, in the Academy of Natural Sciences of Philadelphia,

no. 188844. This species is named for Dr. Donald Bosch, a physi-

cian who has added greatly to our knowledge of south Arabian

marine mollusks.

Comparative Remarks: Cymatium boschi differs from other

members of the subgenus Ranularia in having the upper part of

the whorls tabulate. The flat top of the shoulder is about at right

angle to the wall of the whorl above. It also has a broad and rather

short siphonal canal, a feature which gives the whole shell a some-

what trigonal shape. The upper or posterior end of the white aper-

ture is broadly V-shaped and not constricted into a pseudo-posterior

canal as that in such species as trilineatum (Reeve, 1844), mori-

tinctum (Reeve, 1844) and sinense (Reeve, 1844). The parietal

wall is white, a feature found in caudatum (Gmelin, 1791) which

has a deep sutural channel, and in sinense, which has 6 or 7 very

large teeth on the inside of the outer lip. C. boschi differs from

gutturnium (Roding, 1789) in that the latter has a globular body

whorl with beaded spiral cords and a peach to reddish oval peri-

stome. The periostracum of boschi appears to lack the occasional,

long, dark-brown setae seen in gutturnium and sarcostorna (Reeve,

1844).

ASPELLA MYRAKEENAE, NEW SPECIES FROM

WESTERN MEXICO

11y WILLIAM K. EMERSON1 and ANTHONY D'ATTILIO2

Recent collecting along the central coast of western Mexico has

brought to light specimens of a distinctive new species of aspellid

gastropod. We take extreme pleasure in describing this interesting

discovery in honor of Dr. A. Myra Keen, distinguished paleontolo-

gist of Stanford University, California.

1 American Museum of Natural History, Central Park West, New York, N. Y.

10024.

2 San Diego Natural History Museum, California 92001

January, 1970 nautilus 89

Aspclla (? Dermomurex) myrakeenae Emerson and D'Attilio, new

species Figures 1-11

Description of holotype: Shell is nearly 25 mm. in height, broadly

ovate, widest below the median area, with six convex whorls; sur-

face is chalky white; the worn nucleus consists of I1/9 whorls. There

are 6 broad, strong, obliquely descending axial ridges on the spire

and the body whorl which continues over the shoulder. The varices

are attached buttress-like to the preceding whorl where they are

excavated on the receding side. The excavations continue as mod-

erately deep basins on the shoulder. The suture is deeply im-

pressed. The spiral sculpture is mostly diminished or obsolete in

the areas between the varical sculpture. On the varices the spiral

sculpture consists of strong, large cords of which there are 2 on

the spire and 5 on the body whorl. Where the spiral sculpture on

the varices reaches die surface of the body whorl, the deeply

carved troughs between the spiral cords terminate abruptly. The

polished, ovate aperture is widely flaring, and it has a white

border. More deeply within the aperture there are visible 6 spiral

bands of brown. The outer lip is crenulated by the spiral cords

over-running the axial sculpture; 6 denticles are present within the

outer lip; anteriorly the first 2 are paired, followed by 3 evenly

spaced, with the 6th and most posteriorly situated one weakly

developed. The siphonal canal is moderately short, open, and re-

curved. The fasciole retains a strong ridge consisting of the blunt

ends of the axial varices. A slight umbilical excavation occurs at

the upper portion of the attachment of the fasciole to the canal.

The white chalky layer of the shell surface is finely, axially striated.

Where the outer chalky layer of the shell is abraded off the sur-

face, especially on beach specimens, the harder shelly matter below

is revealed to be incised by close, regularly spaced, sharp edged,

spiral channels. If the chalky layer is completely removed, the

areas between the spiral cords on the varices and the body whorl

are found to be colored a rusty brown.

The holotype, which is the largest specimen we have seen, is

not complete, being holed in the apertural side of the body whorl

(figs. 1,2) . Specimens with mature apertures have a white porcel-

laneous layer that covers and completely masks the brown bands

in the aperture. The apertural bands are most prominent on speci-

90 nautilus Vol. 83 (3)

mens with newly formed outer lips (fig. 6) . The exterior color

bands under the chalky layer are commonly poorly developed or

they may be lacking in some specimens (figs. 4, 5) . Not all

5 varices are equally well-developed on the body whorl of some

specimens; one or more varices may be reduced to raised ridges

which remain only at the suture as buttresses that are attached to

the previous whorl.

The operculum has a basal nucleus with the inner surface well

marked (figure 10) .

The radular dentition (figure 11) is similar to that of Aspella

obeliscus (A. Adams) , and in major characters recalls those of the

Typhinae and the Muricinae.

Measurements: Holotype, 24.8 mm. in length, 14.5 mm. in

width; largest paratype: 20.8 mm. in length, 13 mm. in width;

smallest paratype: 9.9 mm. in length, 7.1 mm. in width.

Type locality: Banderas Bay, Nayarit, Mexico; intertidal, under

rocks.

Type specimens: holotype, A.M.N.H. No. 153298; 9 paratypes,

A.M.N.H. No. 155887; collected by Joseph and Helen DuShane,

January 12-18, 1969. Additional paratypic specimens will be de-

posited in the following collections: San Diego Natural History

Museum, Los Angeles County Museum of Natural History, Cali-

fornia Academy of Sciences, Department of Geology of Stanford

University, Department of Geology of Tulane University, and

that of Joseph and Helen DuShane of Whittier, California.

Other specimens examined:

La Paz, Baja California, Mexico, A.M.N.H. No. 155905, 1 speci-

men, intertidal, under rocks, Twila Bratcher, April, 1965, ex. -

Ben and Ruth Purdy collection. Mariner's Point, Mazatlan,

Sinaloa, Mexico, A.M.N.H. No. 152592, 2 specimens, intertidal,

Tom Rice, February 12, 1968. Off Mazatlan, Mexico, A.M.N.H.

No. 152600, 2 specimens, by SCUBA diving, December, 1968, ex. -

Richard Petit collection. Barra de Navidad, Jalisco, Mexico,

A.M.N.H. No. 153353, 12 specimens, intertidal, under rocks, low

tide, Joseph and Helen DuShane, February 1, 1969.

Remarks: Specimens of this new species are currently masquer-

ading in collections under the name Aspella indentata (Carpenter,

1 857, p. 527) , a poorly known taxon originally described from

Mazatlan, Sinaloa, Mexico (Keen, 1968, p. 425, pi. 58, fig. 64,

January, 1970 nautilus 91

holotype illustrated) . Carpenter's species, which is rare in collec-

tions, has an elongate shell with a thick, denticular outer lip

formed in the comparatively small aperture of the mature speci-

mens. We have examined a beach-worn specimen with a mature

aperture (see fig. 3, herein) , which is in the collection of the Los

Angeles County Museum of Natural History and it is accompanied

by a label reading "Santelmo Bay, West Mexico, Capt. Fred

Lewis." According to Dr. James H. McLean (in litteris), Capt.

Lewis collected in the Bay of Panama as well as along the Mexican

coast. Therefore, this specimen may well have been obtained by

him at Bahia Santelmo, Isla del Ray of the archipelago of the

Perlas Islands, Panama, the type locality of the holotype of Aspella

perplexa Keen, 1958.

Keen (1958a, pi. 30, fig. 13) illustrated what appears to be a

specimen of this new species from Zihuatanejo, Guerrero, Mexico,

as a questionable hypotype of Aspella perplexa Keen (1958a,

p. 248, pi. 30, fig. 1 1, holotype) . Keen (1958b, p. 365, fig. 378) has

referred the holotype of her A. perplexa to the synonymy of A. in-

dentata Carpenter, but she subsequently expressed doubt regard-

ing this action after examining Carpenter's type specimen (Keen,

1968, p. 425). The illustration of the holotype of A. perplexa,

suggests a well-preserved specimen that appears to have an im-

mature outer lip. Additional Panamanian material is needed before

the mature apertural characters of specimens from this region can

be determined with certainty. The limited data, however, suggests

to us that the holotype of A. perplexa probably falls within the

expected morphological range of variability of Carpenter's A. in-

dentata and that this species apparently ranges from Panama to

Mexico.

The new species also has been confused with the Panamic As-

pella obeliscus (A. Adams, 1853, p. 269; Sowerby, 1879, Murex sp.

90b, pi. 402, fig. 233, not Murex obeliscus Dillwyn, 1817, which is

Cerithium obeliscus Bruguiere, 1792 [= Rhinoclavis sinensis

(Gmelin, 1791)]). "Murex" obeliscus A. Adams was originally

described without locality data, but this taxon was subsequently

cited from St. Thomas Island in the Caribbean (Sowerby, 1879) .

This citation led to the supposition on the part of students that

A. obeliscus was a junior synonym of the Caribbean A. pauper Tula

(C. B. Adams, 1850, p. 60; Clench and Turner, 1950, p. 323, pi. 39,

92 nautilus Vol. 83 (3)

fig. 16, lectotype, Jamaica, type locality) . Specimens from western

Mexico (figure 7) , however, appear to be close to Sowerby's

figured specimen. At our request, Dr. John Peake of the British

Museum (Natural History) kindly provided us with photographs

of three syntypes of A. obeliscus, two of which are figured herein

(figs. 8,9) . None of the syntypes of A. obeliscus now possesses the

rich coloring as depicted by Sowerby's illustration, although one

of the syntypic specimens (fig. 9) closely approaches it in general

form and proportions. Sowerby's drawing may represent a com-

posite of the three syntypes, or it may have been based on non-

typological material.

As pointed out by Keen (1968, p. 425) , the New World repre-

sentatives of Aspella (sensu lato) are in dire need of study. Since

the last review of this muricid group appeared (Hertlein and

Strong, 1951, pp. 77-79) , much new information has accumulated.

However, critical data on the radular and opercular characters of

the type species of the supraspecific taxa that have been placed in

this complex must be obtained before generic assignments can be

undertaken with confidence.

Acknowledgements: We are grateful to the following friends and

colleagues for valuable assistance in providing us with specimens

and for extending courtesies of various kinds: Mrs. Twila Bratcher

of Hollywood, California, Mr. and Mrs. Joseph DuShane of Whit-

tier, California, Dr. A. Myra Keen of Stanford University, Stan-

ford, California, Dr. James H. McLean of the Los Angeles County

EXPLANATION TO FIGURES (Opposite Page)

Figures 1,2,4,5,6, Aspella (?Dermomurex) myrakeenae, n.sp. Figs. 1,3, holo-

type. Figs. 4,5, La Paz, Baja California, Mexico; specimen with mature aper-

ture; ex-Purdy collection. Fig. 6, Paratype, showing immature outer lip with

color banding. All X2. Fig. 3, Aspella (?Dermomurex) indentata (Carpenter,

1857) , a beach-worn specimen from "Santelmo Bay," [?Isla del Ray, Panama],

L.A.C.M.N.H. No. A5498, courtesy of Dr. J. H. McLean; specimen with mature

aperture. X2. Figures 7-9, Aspella (?Dermomurex) obeliscus (A. Adams, 1853) .

Fig. 7, Masachapa, Nicaragua, beach specimen, courtesy of Dr. E. H. Vokes.

X2. Figs. 8-9, 2 of 3 syntypes in British Museum (Natural History) , teste

Peake; photographs courtesy of Dr. John Peake (enlarged) .

January, 1970

NAiniLUS

8

94

NAUTILUS

Vol. 83 (3)

Figure 10- Inner and outer surfaces of the operculum of Aspella (TDertno-

murex) myrakeenae, new sp.; greatly enlarged.

Museum of Natural History, Los Angeles, California, Dr. John

Peake of the British Museum (Natural History) , London, Mr.

Richard Petit of Ocean Drive Beach, South Carolina, Mr. and Mrs.

Ben Purdy of San Diego, California, Dr. George E. Radwin of the

Natural History Museum, San Diego, Mr. Thomas C. Rice of

Poulsbo, Washington, Mr. and Mrs. Carl Shy of Westminister,

California, and Dr. Emily H. Vokes of Tulane University, New

Orleans, Louisiana.

Literature cited

Adams, Arthur, 1853. Proc. Zool. Soc. London, for 1851, pp. 267-

272 (December, 7) .

Adams, C. B., 1850. Contributions to Conchology, no. 4, pp. 56-68

(January) .

Carpenter, P. P., 1857. Catalogue of the collection of Mazatlan

shells in the British Museum collected by Frederick Reigen.

London, i-iv -f ix-xvi + 552 pp.

Clench, W. J., and R. D. Turner, 1950. Occas. Papers on Mollusks,

Harvard Univ., 7 (15) : 233-404, pis. 29-403.

Hertlein, L. G., and A. M. Strong, 1951. Bull. So. Calif. Acad. Sci.,

50 (2) : 76-80, pi. 26.

Keen, A. M., 1958a. Bull. Anier. Paleont., 38 (172) : 236-255, pis.

30-31 (May, 23) .

, 1958b. Sea shells of tropical West America; marine mollusks

from Lower California to Colombia, i-ix + 624 pp., 1700 text

figs. Stanford Univ. Press. (December, 5) .

-, 1968. Veliger, 10 (4): 389-439, pis. 55-59, 171 text figs.

(April, 1).

January. 1970 nautilus 95

Sowerby, G. B., Jr. (Second of Name), [Editor], 1879. Thesaurus

conchyliorum, or Monographs of genera of shells. London.

4, Murex pp. 1-55, pis. 380-405.

Figure 11. Radular dentition showing central tooth and one lateral tooth

of Aspella (TDermomurex) myrakeenae, n. sp., from Coastocomate, Jalisco.

Mexico, C. and L. Shy, leg.; greatly enlarged; (radular mount courtesy of

Dr. (i. E. Radwin) .

NEW MITRIDAE AND VOLUTOMITRIDAE

By WALTER O. C HRNOHORSkY

Auckland Institute & Museum, Auckland, New Zealand

Family Mitridae

Genus Concilia Swainson, 1840

Type species by subsequent designation (Herrmannsen, 1846)

Tiara isabella Swainson, 1831.

Cancilla scrobiculata crosnieri new subspecies. Figs. 1 and 2

Description: Shell moderately large, fusiform-elongate, thick and

solid, uniformly creamy-white under a thin, black periostracum.

Teleconch of 9 convex whorls, protoconch missing, sutures distinct

but not deep. Sculptured with irregular and unequal spiral threads,

about 42 on the body whorl and 19 on the penultimate whorl in

the type; some spiral threads are broader and flatter than others in

the same specimen. Interstices of spirals almost smooth in parts,

but finely axially striate or even pitted in some portion of the

shell; fine, close-set and somewhat obsolete longitudinal growth

striae override spirals cords and interstices. Aperture equal in

height to the spire, narrow and elongate, smooth within, edge of

labial lip thickened and simple; columella with 5 oblique folds,

siphonal fasciole calloused, moderately straight or slightly recurved.

All specimens were collected devoid of animal. The new sub-

96

NAUTILUS

Vol. 83 (3)

Fig. 1. Concilia scrobiculata crosnieri Cernohorsky. Holotype DMNH No.

22392. Point Noire, Congo, West Africa, 135 metres. Length 73.1 mm.

Fig. 2. C. scrobiculata crosnieri Cernohorsky. Paratype ANSP No. 316133.

Point Noire, Congo, West Africa, 135 metres. Length 89.8 mm. Fig. 3. Mitra

pele Cernohorsky. Holotype USNM No. 679608. 1 mile NW of Pitcairn Id.,

Pacific Ocean, 65-70 fathoms. Length 41.1 mm. Fig. 4. M. pele Cernohorsky.

Paratype BPBM. Off Waikiki, Oahu, Hawaii, 100 feet. Length ca. 49.0 mm.

(photo Dr. A. Kay, Honolulu) . Fig. 5. M. pele Cernohorsky. Paratype USNM

No. 679610. 1 mile NW of Pitcairn Id., Pacific Ocean. Length 24.9 mm.

January, 1970

NAUTILUS

97

Fig. 6- Vexillum (Costellaria) nodospiculum Cernohorsky. Holotype USNM

No. 274716. Off Ft. Duniurug, Masbate, Philippines, 153 fathoms. Length

6.0 mm. Fig. 7. V. (C.) nodospiculum Cernohorsky. Faratype USNM No.

279049. Off Silungan Id., Borneo, 305 fathoms. Length 6.3 mm. Fig. 8- Micro-

voluta joloensis Cernohorsky. Holotype USNM No. 288396. Off Cagayan Id.,

folo Sea, Philippines, 508 fathoms. Length 9.5 mm. (slender, costate speci-

men) . Fig. 9- M. joloensis Cernohorsky. Faratype USNM No. 290450. Iligan

Bay, Nth. Mindanao, Philippines, 445 fathoms. Length 8.6 mm. (slender,

smooth specimen) . Fig. 10. M. joloensis Cernohorsky. Faratype USNM

290189. Off Apo Id., Sth. Negros, Philippines, 256 fathoms. Length 9.3 mm.

(broad, obscurely costate, internally Urate specimen) .

98

NAUTILUS

Vol. 83 (3)

Fig. 11. Early whorls of Vexillum (Costellaria) nodospiculum Cernohorsky.

Fig. 12. Early whorls of Microvoluta joloensis Cernohorsky.

species exhibits the usual amount of variation one has come to

expect in Mitridae, some specimens being more slender than others,

spiral cords are either elevated and rounded or depressed and flat.

There are 35-45 spiral threads on the body whorl and 1 3-20 on the

penultimate whorl; interstices may be moderately smooth, finely

axially striate or even prominently pitted. The number of adult

whorls will vary from 8-10 and columellar folds from 4-5.

Holotype measurements: Length 73.1 mm.; max. diam. 18.5 mm.;

height of aperture 37.0 mm.

Type locality: Point Noire, Congo, West Africa, Lat. 4°57/S,

Long. 11°22'E, in 135 metres (4-V-1962) .

Distribution and types: Holotype No. 22392 in the Delaware

Museum of Natural History, Greenville; paratype No. 1 (89.8 x

21.0 x 43.7 mm.) in the Academy of Natural Sciences of Phila-

delphia ANSP 316133; from the type locality; paratype No. 2

(85.3 x 23.2 x 43.7 mm.) in the Laboratoire de Malacologie,

Museum d'Histoire Naturelle, Paris, from Point Noire, Congo,

160 metres (leg. J. Marteau, 12-VIII-1966) ; paratype No. 3 (76.8 x

21.0 x 38.6 mm.) in author's collection, Auckland Institute & Mu-

seum, Auckland, from coast of Dahomey, Lat. 6°05'N, Long.

2°37'E, in 150-200 metres (leg. A. Crosnier, 24-X-1963) ; paratype

No. 4 (71.4 x 18.8 x 36.6 mm.) and paratype No. 5 (56.4 x 14.6 x

28.8 mm. - juvenile specimen) in the Delaware Museum of Natural

History, Greenville, from coast of Dahomey, 150-200 metres.

The West African subspecies is the living representative of the

European Mio-Pliocene Cancilla scrobiculata (Brocchi, 1814) , and

has close relatives in Miocene deposits of the Caribbean and North-

ern South America: C. symmetrica (Gabb, 1873) from Dominica,

C. limonensis (Olsson, 1922) from Costa Rica and C. sieversi

January, 1970 nautilu3 99

(Rutsch, 1934) from Venezuela. The remaining two surviving

members of the scrobiculata complex, are geographically widely

separated: Cancilla larranagai (Carcelles, 1947) , a somewhat

stunted, less prominently sculptured form from temperate waters

olf Uruguay (58-65 fathoms) , and the tropical C. Isabella

(Swainson, 1831) from the Sino- Japanese region.

The new subspecies is named for Prof. A. Crosnier, Point Noire,

Congo, who first brought the species to our attention, and for his

work on West African molluscs.

Genus Mitra Roding, 1798

Type species by subsequent designation (Montfort, 1810) Valuta

episcopalis Linnaeus, 1758 =z Vohita mitra Linnaeus, 1758.

Mitra pele new species. Figs. 3 and 4

Description: Shell medium in size, ovate-fusiform, thick and

solid, tan in colour under a thin dark brown periostracum. Tele-

conch of 4i/> almost flat-sided whorls, portion of spire and proto

conch missing, sutures distinct but not deep. Sculptured with

smooth and thin spiral grooves, about 30 on the body whorl and

9 on the penultimate whorl; under magnification, close-set, fine

longitudinal striae are discernible on the shell. Aperture about

equal in height to the spire, moderately narrow, white in colour

and smooth within; labial lip thickened and simple, columella with

a white prominent callus and 7 oblique, close-set folds, siphonal

canal straight.

In the paratype specimens, the spiral grooves on the body whorl

number from 25-30 and columellar folds from 5-7. Juvenile speci-

mens are light orange-brown on the lower half of the body whorl

while remaining whorls are dark brown.

Holotype measurements: Length 41.1 mm. -(-; max. diam. 14.5

mm.; height of aperture 22.7 mm.

Type locality: Station PIT VI, Haul 20, 1 mile NW of Pitcairn

Island, Pacific Ocean, in 65-70 fathoms, stones and coral rubble

(Pele Expedition, 20-X-1967) .

Distribution and types: Holotype No. USNM 679608 in the U.S.

National Museum, Washington; paratype No. 1, USNM 679609

(39.4 x 14.5 x 21.8 mm.) , from the type locality, in the U.S. Na-

tional Museum, Washington; paratype No. 2, USNM 679610 (24.9

100 nautilus Vol. 83 (3)

x 9.2 x 13.2 mm. - juvenile specimen) , Sta. PIT, Haul 19, 1 mile

NW of Pitcairn Island, in 55-65 fathoms, in U.S. National Mu-

seum; paratype No. 3 (ca. 49.0 mm.) off Waikiki, Oahu, Hawaii,

in 100 feet, in Bernice P. Bishop Museum, Honolulu.

Mitra pele is the same species as M. nigra J. Cate, 1960 (non

Voluta nigra Gmelin, 1791) , from 75 fathoms, Keehi Lagoon, Oahu

Hawaii. Mitra nigra (Gmelin) , based on an illustration in Chem-

nitz (1788, p. 168, pi. 151, figs. 1430, 1431) from "Guinea," West

Africa, is the same species as Mitra fusca Reeve, 1844, and of

authors, but not of Swainson, 1824. Mitra castanea Roding, 1798,

and M. melaniana Lamarck, 1811, are objective synonyms, M. adan-

sonii Philippi, 1849, a subjective synonym of M. nigra (Gmelin) ;

the species is restricted to the North-eastern Atlantic and West

Africa. Chemnitz (loc. cit.) remarked that he received the "genus"

(not species) from Tranquebar and Greenland; from India he

most probably received Mitra chinensis Griffith & Pidgeon, 1834,

and from Greenland very likely Volutomitra groenlandica (Beck

in Moller, 1842).

The taxon Mitra nigra (Gmelin) , has also been erroneously

associated with the Austral-New Zealand species Vici mitra conter-

mina Iredale, 1936 (non Mitra contermina Bellardi, 1887). The

Austral-Neozelanic species is Mitra carbonaria Swainson, 1822, and

the distributional range of the species is south from New South

Wales to Tasmania, across the Tasman to New Zealand (synonym

M. maoria Finlay, 1927) and northward to the Kermadec Islands.

Mitra carbonaria is thinner and less solid than the Polynesian

M. pele, has finer, punctate spirals and a brown aperture; the an-

terior of the parietal wall is slightly concave, dark brown in colour,

and lacks the white columellar callosity of M. pele. The spiral

striae become basal cords on the siphonal fasciole in M. carbonaria,

whereas in M. pele the spiral grooves continue to the base, with

intervening flat spiral cords. Juvenile specimens of the two species

differ quite prominently.

Mitra fusca Swainson, 1824, described from the Indian Ocean,

was subsequently well illustrated by Swainson (1831) , but wrongly

figured by Reeve (1844) and erroneously attributed to the Atlantic

Ocean by subsequent authors. Scabricola (Siuainsonia) fusca (Swain-

son) , is the earliest name for the Indo-Pacific species subsequently

described as Mitra limata Reeve, 1845, Cylindra formosa Pease,

January, 1970 nautilus 101

1868, and Mitra zephyrina Sowerby, 1874.

Mitra pele has not been recorded outside the Hawaiian-Poly-

nesian region.

Genus Vexillum Roding, 1798

Subgenus Costellaria Swainson, 1840

Type species by monotypy Mitra rigida Swainson, 1821 =

M. semifasciata Lamarck, 1811.

Vexillum (Costellaria) yiodospiculum new species Figs. 6, 7, 11

Description: Shell very small, fusiform, moderately thin, white

in colour throughout. Teleconch of 5i/2 concave and nodulose

whorls, part of protoconch missing but remaining whorls smooth

and milky-white; whorls concavely excavated centrally, sutures

rather deep. Sculptured with prominent axial ribs, about 17 on the

body whorl and 13 on the penultimate whorl; axial ribs form coarse

nodes anteriorly and posteriorly to the sutures. Interstices of axial

ribs smooth, except on the centre of the body whorl where 4

prominent spiral cords are visible. Siphonal fasciole with 6 dis-

tinct and oblique cords, 4 of which are extensions of the columel-

lar folds. Aperture narrow, shorter than spire, lirate within, labial

lip constricted anteriorly; columella with 4 oblique folds, anterior

fold the smallest, parietal wall with a callus pad.

The range of variation in 49 specimens of the type series is as

follows: length 3.2-6.6 mm.; width 34%-41% of length; height of

aperture 36-45% of length; teleconch 4-5i/£ whorls; protoconch

2i/9-3i/2 whorls; axial ribs on body whorl 8-17; on penultimate

whorl 9-14; spiral striae on body whorl 0-12; on penultimate whorl

1-4; columellar folds 3-4. In some specimens the spiral striae are

obsolete on the body whorl, while in others spiral sculpture is

prominent.

Holotype measurements: Length 6.0 mm.; max. diam. 2.2 mm.;

height of aperture 2.7 mm.

Type locality: Station 5394, off Pt. Dumurug, Masbate, Philip-

pine Islands, in 153 fathoms, green mud (U.S. Bureau of Fish-

eries) .

Distribution and types: Holotype USNM 274716a in U.S. Na-

tional Museum, Washington; paratypes No. 1-12, USNM 274716,

from the type locality, in the U.S. National Museum; paratype

102 nautilus Vol. 83 (3)

No. 13 from the type locality in the author's collection, Auckland

Institute & Museum; paratypes No. 14-15, USNM 259640, Sta.

5194, off Chocolate Id., Nth. Cebu, Philippines, 148 fathoms, green

mud, 56.5°F; paratype No. 16, USNM 259766, Sta. 5197, off

Cabilao Id., W. Bohol, Philippines, 174 fathoms, green mud,

54.3°F; paratype No. 17, USNM 259755, same data as preceding;

paratype No. 18, USNM 277531, Sta. 5212, off Sibugay Id., E. of

Masbate, Philippines, 108 fathoms, grey mud and sand, 59.9°F;

paratype No. 19, USNM 288875, Sta. 5407, off Ponson Id., Camotes

Islds., Philippines, 350 fathoms, green mud; paratype No. 20,

USNM 287908, Sta. 5408, off Gapitancillo Id., N. Cebu, Philip-

pines, 159 fathoms, green mud, 55.4°F; paratypes No. 21-24,

USNM 289179, Sta. 5409, same as preceding, but in 189 fathoms;

paratypes No. 25-32, USNM 289146, Sta. 5419, off Lauis Pt.,

E. Cebu, Philippines, 175 fathoms, green mud, 54.5°F; paratype

No. 33, USNM 290285, Sta. 5513, Iligan Bay, Nth. Mindanao,

Philippines, 505 fathoms, grey mud and fine sand, 52.8°F; para-

type No. 34, USNM 286714, Sta. 5541, off Tagolo Pt., Nth. Min-

danao, Philippines, 219 fathoms, fine sand and broken shale,

53.3°F; paratypes No. 35-36, USNM 278819, Sta. 5589, Sibuko

Bay, off Mabul Id., Borneo, Indonesia, 260 fathoms, fine grey sand

and mud, 45.7°F; paratypes No. 37-46, USNM 279049, Sta. 5592,

off Silungan Id., Borneo, Indonesia, 305 fathoms, green mud,

43.3°F; paratype No. 47, Sta. 5592, same data as preceding, in

author's collection, Auckland Institute & Museum; paratype No.

48, USNM 235018, Cebu, Philippines, coll. Bartsch. (all types in

the U.S. National Museum, collected by the U.S. Bureau of

Fisheries, except as listed) .

This minute, spinose deep-water species is one of the smallest

Vexilline species recorded. It bears no real resemblance to any of

the Recent species, but is a distant relative of the common, inter-

tidal species Vexillum (Costellaria) coronatum (Helbling, 1779),

V. concentricum (Reeve, 1844) [=r Mitra mucronata Broderip,

1836 = M. echinata A. Adams, 1853], and V. stephanucha (Mel-

vill, 1897) . These three species greatly exceed V. nodospiculum in

size (20-45 mm.) , are brightly coloured and of different form and

sculptural arrangement.

January, 1970 nautilus 103

Family Volutomitridae

Genus Microvoluta Angas, 1877

Type species by monotypy Microvoluta australis Angas, 1877.

Microvoluta joloensis new species Figs. 8-10

Description: Shell small, fusiformly-elongate, livid in colour.

Teleconch of 61/, convex whorls, protoconch of 2 smooth, globose

nuclear whorls. Sculptured with angulate and arcuate axial ribs,

about 13 on the body whorl and 18 on the penultimate whorl, and

obsolete spiral striae; sutures well impressed and with a faint indi-

cation of a sutural girdle. Aperture narrow, equal in height to

spire, smooth within (in holotype) , labial lip thin and simple,

constricted towards the base; columella concave, with 4 thin, dis-

tant folds, posterior fold shorter than succeeding one. Siphonal

canal spout-shaped, notch lacking, siphonal fasciole slightly twisted

towards the aperture.

The species is very variable in shape, colour and sculpture. Some

specimens are fusiformly-elongate, others more ovate; some indi-

viduals are off-white, banded with livid, or are livid in colour and

have a white sutural band. Many specimens have a well-formed

sutural girdle, and axial ribs tend to become obsolete on the last

2 whorls; axials number up to 18 on both whorls, and spirals are

either distinct or obscure. From the 51 specimens examined, 38

specimens were smooth within the aperture, while 13 specimens

were lirate; this feature is typical of Volutomitridae. The length

of the type specimens ranged from 3.6 - 10.4 mm., the width from

34%-43% of length and the height of aperture from 42%-52%

of length.

Holotype measurements: Length 9.5 mm.; max. diam. 3.4 mm.;

height of aperture 4.6 mm.

Type locality: Station 5423, off Cagayan Island, Jolo Sea, Philip-

pine Islands, in 508 fathoms, grey mud and compact sand, 49.8°F

(U.S. Bureau of Fisheries) .

Distribution and types: Holotype No. USNM 288396 is in the

U.S. National Museum, Washington; paratypes No. 1-11, USNM

288396 and paratype No. 12, USNM 288475, are from the type

locality; paratype No. 13, USNM 288686, Sta. 5424, off Cagayan

Id., Jolo Sea, 340 fathoms, 50.4°F; paratypes No. 14-28, USNM

288133, Sta. 5425, off Cagayan Id., Jolo Sea, 495 fathoms, 49.4°F;

104 NAUTILUS Vol. 83 (3)

paratypes No. 29-38, USNM 290425, and paratype No. 39, USNM

290450, Sta. 5512, Iligan Bay, Nth. Mindanao, 445 fathoms, grey

mud and fine sand, 52.8 °F; paratypes No. 40-41, USNM 290260

and paratype No. 42, USNM 290260a, Sta. 5513, Iligan Bay, Nth.

Mindanao, 505 fathoms; paratypes No. 43-49, USNM 290134,

Sta. 5529, off Balicasag Id., Sth. Bohol, 441 fathoms, grey mud,

53 °F; paratype No. 50, USNM 290189, Sta. 5538, off Apo Id.,

Sth. Negros, 256 fathoms, green mud and sand, 53.3°F (all Philip-

pine Islands, U.S. Bureau of Fisheries) . All paratypes are in the

U.S. National Museum, with the exception of paratypes No. 11

and No. 38, which are in the author's collection, Auckland In-

stitute &: Museum, Auckland.

This is the first Microvoluta reported from the Philippine

Islands. The species is superficially similar to Microvoluta inter-

media (Dall, 1890), from the West Indies, but in that species

the whorls are angulate, axial ribs thin, straight and obscure below

the presutural ramp. With other Microvoluta species it shares fea-

tures of thin shell, bulbous protoconch, spout-shaped siphonal

canal, absence of siphonal notch, smooth or lirate labrum and a

recurved beak.

Acknowledgements: I would like to thank Dr. R. T. Abbott,

Delaware Museum of Natural History, Greenville, and Dr. Harald

A. Rehder, Smithsonian Institution, U.S. National Museum, Wash-

ington, for giving me the opportunity to describe these new mol-

luscan species.

Literature cited

Cate, J. M., 1960. Range extension and synonymy for Mitra nigra

(Schroter, 1788). The Veliger, 3(2): 49-51.

Chemnitz, J. H., 1788. Neues systematisches Conchylien-Cabinet.

Nurnberg, 10: 1-376; pits. 137-173.

Reeve, L. A., 1844-45. Conchologia Iconica. Monograph of the

genus Mitra. London, Vol. 2, pits. 1-39.

Swainson, W., 1824. The characters of several new shells, belonging

to the Linnaean Volutae, with a few observations on the present

state of Conchology. Quart. Journ. Sci. Lit. & Arts, 17 (33) :

28-38.

, 1829-33. Zoological Illustrations. London, 2nd series, Vols.

1-3, pits. 1-136.

January, 1970 nautilus 105

REDISCOVERY OF THE LIMPET, ACROLOXUS

COLORADENSIS (BASOMMATOPHORA:

ACROLOXIDAE), IN COLORADO1

By GEORGE W. BRYCE, JR.

Department of Biology, University of Colorado, Boulder, Colorado

Professor Junius Henderson of the University of Colorado on

two different occasions in the 1920's collected specimens of Acro-

loxus coloradensis (Henderson) , a small limpet, from a mountain

lake in Boulder County, Colorado. Since that time the species has

been reported alive only once (Russell and Brunson, 1967) from

the United States, although many others (Basch, 1963) have

searched specifically for it on various occasions at its type locality.

In October 1966 at Peterson Lake I found two living specimens

of A. coloradensis. A subsequent careful search by me and Scott J.

Herrmann, of the Department of Biology at Southern Colorado

College, revealed a small population in shallow littoral regions

of the lake.

Henderson first found this elusive species on 28 June 1920, and

sent specimens to Bryant Walker for identification. In the Cata-

logue of Mollusca of the University of Colorado Museum, entry

number 10,113 in Henderson's own hand quotes Walker as writ-

ing, "Most remarkable. Notice the spine-like apex. 'For the love

of Mike' get more."

Despite Walker's enthusiastic imploration for more specimens,

the species was not again collected until August 1924 by Junius

and F. G. Henderson. Soon thereafter, Walker (1925) described

the species, naming it Ancylus hendersoni after its discoverer. Five

years later Henderson (1930) found that Walker's specific desig-

nation was preoccupied, having been previously used in the same

genus by Walker in 1908. Henderson renamed the limpet Ancylus

coloradensis. Subsequently the species has been assigned to the

genus Acroloxus (Taylor, 1960) , since it is closely related to the

European Acroloxus lacustris (Linnaeus) . Cytological studies of

A. lacustris (Burch, 1962) suggest that it merits separate familial

status, if not indeed separate ordinal status.

Peterson Lake, the type locality misidentified by Henderson as

"Eldora Lake," is located in Boulder County, Colorado, about 5.5

1 Contribution No. 53, Limnology Laboratory, University of Colorado.

106 nautilus Vol. 83 (3)

kilometers WSW of Nederland and about 25 kilometers WSW of

Boulder (35 road-kilometers from Boulder) . Its exact location is

T. 1 S., R. 73 W., at adjacent corners of sections 20, 21, 28, and

29 (39°56'N, 105°34'W). A smaller lake lying about 0.1 kilometer

to the southeast and about 30 feet higher in elevation is the true

"Lake Eldora." Commercial developments at Peterson Lake have

markedly changed the character of its swamps in the past several

years.

Peterson Lake is situated in the Montane Zone of the Southern

Rocky Mountains at an elevation of 2818 m (9245 feet) and in the

coniferous forest characteristic of that zone. It is about 0.15 X 0.65

km in size. Total residues of its water ranged from 32.4 to 63.1 mg

per 1 from October 1966 to August 1967, and bound carbonates,

as determined by methyl orange titrations, ranged from 12.83 to

19.30 ppm over the same period. Its waters are derived from

swampy areas and are brown-stained, probably by humic extrac-

tives. Pennak (1945) found a water color of 30 on the platinum-

cobalt scale and a pH of 7.7. Water temperatures at the time of

limpet collections ranged from 11.1 to 1.8° C in October 1966 and

up to 18.8° C in August 1967. A complete ice cover (up to 36 cm

thick in 1966-67) is present for about four months each year.

The present population of limpets is concentrated at the west

or head end of the lake and especially along the southwest shore.

Population densities at favorable sites were found to range up to

72 individuals per square meter. Dorsal shell color of individual

limpets varies from light tan flecked with dark brown to almost

solid dark-brown with occasional spots of light brown. A random

sample of 42 specimens ranged in width and length from 0.64 X

1.04 mm to 2.97 X 4.88 mm, with a mean of 1.66 X 2.69 mm. The

small, flattened shells of the animals blend almost exactly with

the dark rocks on which they live, and their observation is ex-

ceedingly difficult, especially in the shade.

The southwest side of the lake which harbors the main popula-

tion center is thickly strewn with the small to large granitic rocks

on which the limpets live. On this shore the coniferous forest

crowds up to the edge of the lake, producing almost day-long

shade in many areas and resulting in cooler average temperatures,

as evidenced by the much longer period of ice cover on this shore

than on the rest of the lake.

January, 1970 nautilus 107

The limpets have since been collected during different seasons

almost invariably on the unclersuriaces of the rocks, apparently

a Hording a shady, algal-free niche for the limpets. Thus, it is

indicated that they may feed on the film of bacteria and detritus.

The limpets have been collected from a water depth of a few centi-

meters to about 40 cm. Dr. Robert W. Pennak (personal com-

munication) suggests that their population center may lie here

where the moderate wave action would keep the rocks free of

the dense brown silt which is deposited over most of the lake

bottom.

The small planorbid snail Promenetus coloradoensis (Baker) is

almost invariably associated with the limpets. Taylor (Hibbard

and Taylor, I960) synonymizes this species with the more com-

mon P. exactions (Say) . Both the limpest and the snail are fre-

quently found on the same rock, but the planorbid ranges more

widely over the surfaces of individual rocks than does the limpet.

Since the collections of Henderson, living Acroloxus coloraden-

sis has been reported once from the United States, and the only

other reports known to me are from Canada. In the summer of

1925 Alan Mo/ley (1926) reported the species from Jasper

National Park, Alberta. In a later report (1930) Mozley clarified

the localities in which A. coloradensis were collected as Lake No. 1

(not Iris Lake as reported in his 1926 paper) and Lake No. 5,

both in an area of about 53° N latitude in Jasper Park. Iris Lake,

only about 0.4 km east of Mozley 's Lake No. 1, lies at an altitude

of 1306 m (4285 feet) in the Canadian Rocky Mountains. Mozley

sent specimens to Walker who identified them as the species under

discussion.

In the United States, Russell and Brunson (1967) have reported

this species from Lost Lake, Glacier National Park, Montana. This

lake lies at about 49° N latitude at an elevation of 1433 m (4700

feet) . These authors also found the limpet "on the undersurfaces

of rocks in shallow water near the shore."

It is significant that Acroloxus coloradensis has also been re-

cently reported by Dwight W. Taylor (1954, 19(50) as a fossil

from the High Plains. It has been found in the Berends fauna, a

Pleistocene assemblage of southwestern Kansas and adjacent Okla-

homa (Taylor, 1954) and also in the Dixon local fauna of south-

ern Kansas and the Sand Draw local fauna of northcentral Neb-

108 nautilus Vol. 83 (3)

raska (Taylor, 1960) , both the latter of early Pleistocene age. Evi-

dently A. coloradensis is a relict species once widely distributed

over North America, but now with a distinctly disjunct distribu-

tion. Its present altitudinal and latitudinal distribution indicates

that it is a cold-water stenothermal species. It is likely that living

populations of A. coloradensis will be found to occur in other in-

frequent and scattered Rocky Mountain lakes in which the con-

tinued existence of the species has been fortuitously favored.

I wish to express appreciation to Dr. Robert W. Pennak (De-

partment of Biology, University of Colorado) and Dr. Hugo G.

Rodeck (Director, University of Colorado Museum) for their aid

and cooperation in this investigation.

Literature cited

Basch, P. F. 1963. A review of the recent freshwater limpet snails

of North America (Mollusca: Pulmonata) . Bull. Mus. Comp.

Zool., 129: 399-461.

Burch, J. B. 1962. Cytotaxonomic studies of freshwater limpets

(Gastropoda: Basommatophora) . I. The European lake limpet,

Acroloxus lacustris. Malacologia, 1: 55-72.

Henderson, J. 1930. Ancylus coloradensis, new name for A. hen-

dersoni Walker, 1925, not 1908. Nautilus, 44: 31.

Hibbard, C. W., & D. W. Taylor. 1960: Two late Pleistocene

faunas from southwestern Kansas. Contrib. Mus. Paleontol.,

Univ. Michigan, 16: 1-223.

Mozley, A. 1926. Preliminary list of the Mollusca of Jasper Park,

Alberta. Nautilus, 40: 53-56.

Mozley, A. 1930. Reports of the Jasper Park Lakes investigations,

1925-26. The Mollusca of Jasper Park. Trans. Roy. Soc. Edin-

burgh, 56: 647-669.

Pennak, R. W. 1945. Some aspects of the regional limnology of

northern Colorado. Univ. Colorado Studies, Ser. D, 2: 263-293.

Russell, R. H., & R. B. Brunson. 1967. Acroloxus coloradensis

from Montana. Nautilus, 81: 33.

Taylor, D. W. 1954. A new Pleistocene fauna and new species of

fossil snails from the High Plains. Michigan Univ. Mus. Zool.

Occ. Papers, No. 557, 16 p.

Taylor, D. W. 1960. Late Cenozoic molluscan faunas from the

High Plains. Geol. Surv. Prof. Paper 337, 94 p.

Walker, B. 1925. New species of North American Ancylidae and

Lancidae. Michigan Univ. Mus. Zool. Occ. Papers, No. 165,

13 p.

January, 1970 nautiluj 109

LONGEVITY IN ASHMUNELLA, MONADENIA AND

SONORELLA

By MUNROE L. WALTON

1 108 North Central Ave., Glendale, California 91202

An earlier contribution (Walton, 1963) recorded data concern-

ing the length of life in west American land snails kept in my

basement cellar. Information concerning the cages, feeding and

methods of observation is presented there and is not repeated. Ad-

ditional records have been accumulated and death claimed several

individuals reported as still alive in 1963. The opportunity is

taken to provide data on several species and to summarize longev-

ity data on three genera. For assistance in organizing this data and

interpreting the results, I am indebted to Alan Solem, Curator of

Lower Invertebrates, Field Museum of Natural History. Chicago.

The species order follows Walton (1963) .

Ashmunella kochi amblya Pilsbry: Collected li/2 miles up Pine

Springs Canon, Guadalupe Mts., Culberson Co., Texas on March

19, 1952. Two adults were found dead in January 1957 and August

1958. They had produced two young that were first observed in

June 1954. In October 1956, at 21 mm. in diameter, the young

formed a reflected lip. The second generation shells were observed

in copulation on June 25, 1959 and a single juvenile was discovered

on August 6, 1959. Although the second generation shells were

observed in copulation again on August 7, 1960, no additional

young were produced prior to their deaths in August and Novem-

ber 1962. The single third generation specimen reached 18 mm. in

diameter, but did not have a reflected lip, at its death about July

30, 1962, although reaching within a week of three years. In con-

trast, the second generation specimens reached maximum shell

growth in 2 14 years, mated at age 5 and 6, and survived to 814

and 814 years.

Ashmunella townsendi Bartsch: Collected on Hwy. 380, 5.5 miles

west of Lincoln, Lincoln Co., New Mexico on October 19, 1951.

Ten adults were caged and subsequently found dead on March 10,

1960 (4 specimens) ; June 8, 1960; September 4, 1960; September

21, 1960; March 16, 1963; April 4, 1963; May 24, 1963; January

20, 1964 February 25, 1964; and September 18, 1965. Survival time

ranged from 8i/2 to just under 14 years for individuals that had

110 nautilus Vol. 83 (3)

already completed shell growth. If we assume that 2]/4 to 2i/2

years is required for completion of shell growth as in A. kochi

amblya, then a life span of I614 years is quite possible.

Sonorella sabinoensis dispar Pilsbry: Collected from east side of

Pima Canyon, Santa Catalina Mts., Pima Co., Arizona on Novem-

ber 4, 1965. Three caged juveniles reached adulthood and repro-

duced before being found dead on May 25, 1967; November 8,

1967; and December 4, 1967. Twelve second generation young were

hatched February 26, 1966. Four died when very small; two were

found dead May 8, 1966 at 4 mm. in diameter; one on November

7, 1966 at 9 mm. in diameter; one on January 16, 1967 at 15 mm.

in diameter; one on March 12, 1967 at 17 mm. in diameter; while

three developed a reflected lip by December 15, 1967 and are still

alive in April 1969. Fourteen third generation young appeared

July 15, 1968 and remain under observation. An additional eleven

young were noticed first as an egg mass on September 28, 1968 and

hatched October 3, 1968. Less than two years were required from

hatching to adult size and young were produced in less than 214

years. This contrasts greatly with S. virilis (see Walton, 1963,

p. 128) , where two juveniles failed to reach adult size in seven

years of captivity.

Monadenia fidelis beryllica Chace: Collected four miles north

of Gold Beach along U. S. Hwy. 101, Curry Co., Oregon on Sep-

tember 13, 1954. Data on size and death dates for 19 of 22 exam-

ples born in July 1956 were given by Walton (1963, p. 130) . The

last three shells died about August 12, 1963 at 25 mm. in diam-

eter, July 6, 1964 at 29 mm. in diameter and July 22, 1964 at

29 mm. in diameter. Adult size was reached in 214 to almost 3

years, while maximum survivorship was 8 years for two of the

twenty-two specimens.

Monadenia fidelis klamathica Berry: Collected 5 miles upstream

from forks of Salmon River, Siskiyou Co., California on August

27, 1959. Nine of ten caged adults died between February 8th and

September 30, 1960 (see Walton, 1963, p. 130) . The tenth speci-

men died February 14, 1963, having produced ten young about

August 1, 1961. Three examples, at 11, 13 and 15 mm. in diameter,

died in October 1962; the remaining seven died in 1963 on about

March 4th (23 mm.) , July 8th (21 mm. and 15 mm.) , July 25th

(20 mm.), August 15th (19 mm.), August 28th (21 mm.), and

January, 1970 nautilus 111

October 13th (21 mm.) . Adult size is about 30 mm., so that these

juveniles reached two-thirds of adult size in 19-26 months.

Monadenia churchi Hanna and Smith: Collected 1.3 miles west

of Junction City, Trinity Co., California on December 28, 1959.

One 8 mm. juvenile died October 20, 1960 at a size of 11 mm. in

diameter. A 13 mm. juvenile became adult at 26 mm. diameter in

1963 and is still alive. Over the years it had been kept caged with

an Ashmunella. No other snails had been kept in the same cage.

Hence the appearance of a young Monadenia churchi on June 6,

1968, nearly eight years after die death of the other juvenile

M. churchi, was most unexpected. Probably this is the result of

self-fertilization, since eight year storage of sperm would be ex-

ceedingly unlikely, even if an 11 mm. specimen could produce

sperm, which is very unlikely. No other Monadenia had been kept

in the same cage and cross-family fertilization with the Ashmunella

can be dismissed as a meaningful possibility. The only logical

remaining possibility is self-fertilization, since no shifting of speci-

mens between cages was undertaken.

Monadenia troglodytes Hanna and Smith: Collected in a ravine

near Sam well Cave, Shasta Co., California on September 20, 1963.

A 27 mm. adult remains alive in April 1969. A 13 mm. juvenile

reached 19 mm. in diameter on September 19, 1964 and 22 mm.

on July 9, 1965 when it was accidentally dropped, broke and died.

Conclusions: Data presented above and in Walton (1963) indi-

cate that species of Ashmunella may have a life span of up to 814

to I61/2 years. Sonorella sabinoensis dispar reproduced within 214

years of birth, while S. virilis collected as juveniles about two thirds

adult size did not develop adult shell characters in over 7 years.

Since the range of moisture conditions inhabitated by different

species of Sonorella is so large, considerable variation in length of

life is not unexpected. Monadenia fidelis has a life span of at

least 8 years; M. churchi lives for more than 10 years; and M. troglo-

dytes lived for over 5 1/2 years as an adult. The latter two species

are still under observation and these are not maximum figures.

These genera are easily maintained and show considerable varia-

tion in growth rates and length of life. While too long lived for

use in thesis problems, amateur collectors could accumulate very

important data on their life histories and reproductive patterns

with minimum equipment and effort.

112 nautilus Vol. 83 (3)

Reference

Walton, M. L. 1963. Length of life in west American land snails.

Nautilus 75(4) : 127-131.

NOTES

Northward range extension of Cyclinella tenuis Recluz.1

— The venerid bivalve, C. tenuis, was described by Recluz (1852.

Jour, de Conch., 3: 250) from Baie de la Pointe-a-Pitre, Guade-

loupe in the Lesser Antilles. The northernmost occurrence of

C. tenuis was reported as Cape Hatteras, North Carolina, by Dall

(1889. U.S. Nat. Mus. Bull. 37, p. 56) and Johnson (1934. Proc.

Boston Soc. Nat. Hist., 40: 48) . Hackney (1944. Nautilus, 58: 58)

found 3 single valves at Beaufort, North Carolina, during the sum-

mer of 1943. Others, including Dall (1902. Proc. U.S. Nat. Mus.,

26: 373) and Palmer (1927. Paleo. Amer., 1 (5) : 201) stated that

the species lives from as far north as the Cedar Keys, Florida, in

the Gulf of Mexico through the West Indies to Sao Paulo, Brazil.

Recently, Abbott (1968. Seashells of North America, Golden Press,.

N.Y., p. 234) gave the range as southern Florida and the Caribbean.

Cyclinella tenuis was found by M. L. Wass in the York River at

Gloucester Point, Virginia, on August 30, 1961, when a single

specimen was taken at a depth of 26 feet. In June 1962, a second

specimen was dredged in Chesapeake Bay off the mouth of the

Rappahannock River at a depth of 52 feet. Bottom salinity ranges

as low as 18 o/oo at the York River site. Both specimens, each

about 1 inch long, fragile and seemingly distorted, were originally

mistakenly referred to Dosinia discus. Subsequently, adult speci-

mens have been taken by Paul Chanley and Michael Castagna near

the VIMS Eastern Shore Laboratory at Wachapreague, Virginia.

The general scarcity of the species in Chesapeake Bay may be in-

dicated by the fact that the first two specimens were found in a

series of samples containing over 50 square meters of substrate.

Turgeon figured a specimen from Willis Wharf, Virginia, in her

unpublished M.Sc. thesis (1968. Guide to estuarine and inshore

bivalves of Virginia. 126 pp., College of William and Mary).

Further, specimens in the U.S. National Museum were collected by

N. W. Moncure at Buckroe Beach, Virginia, in December 1957,

1 Contribution No. 329 from VIMS, Gloucester Point, Virginia.

January, 1970 nautilus 113

so C. tenuis has been in the area for several years. The occurrence

of this species in Virginia extends its known range northward by

about 180 miles. Apparently, it lives in sandy substrates in shallow

water to 36 fathoms.

Specimens examined: (abbreviations: MCZ, Museum of Com-

parative Zoology; USNM, United States National Museum; VIMS,

Virginia Institute of Marine Science) . VIRGINIA: off mouth of

Rappahannock River; York River, off VIMS pier in 8 m.; off

Yorktown; Wachapreague; off Willis Wharf, Eastern Shore (all

VIMS) ; Buckroe Beach (USNM) . SOUTH CAROLINA: Mag-

nolia, Charleston Co. (USNM) . FLORIDA: off Palm Beach, in

12-14 fms.; Sombrero Key; Pirates Cove, Sugarloaf Key; Sand Id.

(all MCZ) ; west channel entrance to Key West; Dry Tortugas

(both USNM) ; Fort Myers Beach; Sanibel Id. (both MCZ) ; Mul-

let Key; Dunedin; Cedar Key (all USNM) . TEXAS: Matagorda

Bay (USNM) ; Port Aransas; Port Isabel (both MCZ) . MEXICO:

Tuxpan, Veracruz (MCZ) . CUBA: Barrera Station 203, Cabanas

Harbor, in 3-12 fms.; Guantanomo, Cuba (both USNM). JA-

MAICA: Green Id., Hanover; Kingston (both USNM). HIS-

PANIOLA: Puerto Plata, Santo Domingo (MCZ). PUERTO

RICO: Mayagiiez; Playa de Ponce (both USNM: MCZ) . VIRGIN

IDS.: St. Thomas (USNM) . LESSER ANTILLES: 2 m. S. of Fort

George, Scarborough, Tobago, in 36 fms. (MCZ) . COLOMBIA:

Cartagena (USNM) . VENEZUELA: La Orchila (USNM) . BRA-

ZIL: Bahia de Todos os Santos, Bahia; Victoria, Espirito Santo,

(both MCZ) ; Rio de Janiero (USNM) ; Praia da Urea, Districto

Federal (MCZ) ; Tabatinga, Sao Paulo (USNM) . — K. J. Boss,

Museum of Comparative Zoology, and M. L. Wass, Virginia

Institute of Marine Science, Gloucester Point, Virginia 23062.

Hermaphroditism in Margaritifera falcata (Gould)

(Pelecypoda: Margaritiferidae). — Twenty-six specimens of this

species were collected on 4 October 1966 from the Blackfoot River

at Idaho Hwy. -34, 13 miles north of Soda Springs, Caribou

County, Idaho. The animals displayed the anatomical features of

the Margaritiferidae Haas (see Ortmann, 1912, Ann. Carnegie

Mus., 8: 223) , and the morphology of the demibranchs showed

M. falcata to be a member of the subfamily Margaritiferinae

Modell (1942, Arch. Moll., 74: 184). Histological examination of

114 NAUTILUS Vol. 83 (3)

the visceral mass of each animal (10 slides per specimen) revealed

that M. falcata consists only of hermaphrodites; true males and

females were lacking in the collection. These monoecious animals

contained proportionately much more ovarian tissue than testicu-

lar tissue which was irregularly scattered in patches ventro-medially

to the digestive gland. Very little gametogenesis was present and

few sperm morulae were found, and the great bulk of the gonads

was filled with mature gametes. Van der Schalie (1966, Mala-

cologia, 5: 77) stated that M. margaritifera and Cumberlandia

monodonta have "occasional hermaphrodites" and that no mar-

garitiferid is "usually monoecious." Margaritifera falcata of west-

ern North America proves to be an unexpected exception. Among

other unionaceans, only the unionids Lasmigona compressa, L. sub-

viridis (Anodontinae) and Carunculina parva (Lampsilinae) are

uniformly monoecious (van der Schalie, Op. cit.) .

None of the 26 individuals of M. falcata in this October collec-

tion from Idaho were gravid. Murphy (1942, Calif. Fish Game,

28: 89-102) demonstrated that in California this species is a short-

term breeder (i.e., tachytictic) , being gravid only between mid-

May and late June. Ortmann (Op. cit.) previously reported that

the Holarctic M. margaritifera is gravid in the summer (p. 232) ,

and that M. hembeli of the southern United States is not gravid in

February (p. 236) . It is peculiar that in M. falcata large quantities

of mature gametes are present more than seven months before

ova can be expected to be released to the demibranchs. — William

H. Heard, Florida State University, Tallahassee, Florida, 32306.

Anodontoides ferussacianus (Lea) in the Susequehanna River

watershed in New York State. — In the summer of 1969 a biologi-

cal survey at the source of the Susquehanna River (Otsego Lake,

Otsego County, New York) was undertaken. This study was part

of an integrated research program initiated at the Stephen C.

Clark Biological Field Station at Cooperstown, a facility of the

Biology Department at the State University College at Oneonta.

Six species of Unionidae have been collected in the Lake. They

are, in order of abundance, Lampsilis radiata (Gmelin) , Anodonta

cataracta (Say) , Elliptio complanata (Solander) , Alasmidonta un-

dulata (Say) and Anodontoides ferussacianus (Lea) . Two living

specimens of the latter have been found. The valves are in good

January, 1970 nautilus 115

condition and the definitive beak sculpture characteristic of the

species shows clearly.

Except for A. ferussacianns all species collected are commonly

found in die Susquehanna watershed. In the most recent authori-

tative work on New York State mollusks Clarke and Berg (1959)

gave the distribution of A. feriissaciamis as follows "Mississippi and

Ohio drainages, and St. Lawrence drainage from Lake Michigan

to the Ottawa River. Lake Champlain, (H. D. Athearn, personal

communication)." Those authors felt the records of A. ferussaci-

anus in the Susquehanna watershed given by Marshall (1895)

were probably incorrect.

In light of this new record Marshall's data should be re-evalu-

ated. On the other hand, the specimens were found only at the

mouth of one stream flowing into Otsego Lake. This inlet finds its

source in a swampy area to the north which also drains to the

Mohawk River.

Because the Erie Canal allows for mixing of the St. Lawrence

and Mohawk River faunas it is possible that A. feritssacianus has

recently been introduced into the Susquehanna watershed via these

streams. — Willard N. Harman, New York State University Col-

lege, Oneonta.

References

Clarke, A. H. and C. O. Berg., 1959. The freshwater mussels of

central New York. Cornell Univ. Agr. Exp. Sta. Mem. 367:

1-79.

Marshall, W. B., 1895. Geographical distribution of New York

Unionidae. N. Y. S. Mus. 48th Ann. Rept. 47-99.

NEWS

The 36th annual meeting of the American Malacological Union

will be held from July 16 through July 20, 1970. Winter address

for the secretary is: Mrs. Margaret C. Teskey, P.O. Box 239, Big

Pine Key, Florida 33043. The meetings will be in Key West,

Florida.

The Third Annual meeting of the Western Society of Malacolo-

gists is scheduled for June 24 to 27, 1970, at Stanford University.

Officers of the society are: President: Myra Keen. Vice-President:

Eugene V. Coan. Second Vice-President: Beatrice Burch. Secretary:

Carol Skoglund. Treasurer: Barry Roth. Councillors-at-large:

Twila Bratcher and Robert Talmadge.

116 nautilus Vol. 83 (3)

A symposium on advances and new techniques in molluscan

taxonomy is being organized by Eugene Coan, and other topics

for symopsia are under consideration. Inquiries about the meeting

and applications for membership should be sent to the secretary,

Mrs. Paul Skoglund, 3846 East Highland Avenue, Phoenix, Ari-

zona 85018.

Percy A. Morris, well-known writer of popular books on con-

chology, and Chief Preparator at the Yale Peabody Museum of

Natural History, died on December 14, 1969, after a short illness,

at New Haven, Connecticut, at the age of 70. He was born January

2, 1899, in Seymour, Connecticut. During his 41 years at Yale he

curated the important A. E. Verrill U.S. Fish Commission collec-

tions. Among the ten books he authored in natural history, his

best known were the "Field Guides to the Shells," published by

Houghton Mifflin Co. in 1947 and 1952. He founded the Connec-

ticut Shell Club in 1946 and was its President for 23 years. Mr.

Morris is survived by his wife, nee Violet French, and by a

daughter, Mrs. Marion Rasey.

January, 1970 nautilus hi

PUBLICATIONS RECEIVED

Okutani, Takashi and John A. McGowan. 1969. Systematics, Dis-

tribution and Abundance of the Epiplanktonic Squid (Cephalo-

poda. Decapoda) Larvae of the Californian Current April, 1954

- March, 1957. Bull. 14 Scripps Inst. Oceanography, Univ. Calif.

Press, 90 pp., illus.

Adam, W. and J. Knudsen. 1969. Quelques Genres de Mollusques

Prosobranches Marins Inconnu ou peu Connus de lAfrique

Occidentale. Bull. Inst, royal Sci. nat. Belgique, 44, no. 27: 1-69.

New species of Cir cuius, Cynisca, Tornus, Macromphalina, Coch-

liolepis, and Megalomphalus , all beautifully illustrated.

Malacological Review, vol. 1. 1968. 187 pp. Editor unknown; pub-

lisher unknown. Address: P.O. Box 801, Whitmore Lake, Michi-

gan 48189. $5.00. Contains 7 excellent original articles on land,

freshwater, marine mollusks and cephalopods. Reproduces "List

of Contents" for 13 other molluscan journals published in 1967.

Haas, Fritz. 1969. Superfamilia Unionacea. Das Tierreich, Berlin,

Lief. 88, pp. i-x, 1-663. Walter de Gruyter and Co., 13 Genthiner

Strasse, Berlin 30, West Germany. (DM 380, which is about

$103.00!) . An exhaustive compendium of the freshwater mussels

of the world, giving keys to families, genera and subgenera;

synonymies, distributions and remarks for all species, and con-

taining an accounting for about 6,000 names. No illustrations

of species.

Young, Richard E. and Clyde F. E. Roper. 1969. A Monograph of

the Cephalopoda of the North Atlantic: the Family Cyclo-

teuthidae. Smithsonian Contributions to Zoology, no. 5, pp. 1-24.

Includes DiscoteutJiis new genus, and D. discus and laciniosa

new species.

Parodiz, Juan J. 1969. The Teritary Non-Marine Mollusca of

South America. Annals of Carnegie Museum, vol. 40, 242 pp.,

numerous photographs, maps, covering 22 families and 48

genera, including 10 new species of freshwater and land gastro-

pods, and Paleoanculosa, new genus, in the family Pleuroceridae.

iv nautilus Vol. 83 (3)

Something New in Conchology

A professional recording of the pronunciations of 1,200

scientific species and generic names of

"Seashells of North America"

also how to enunciate — authors' names, family names, and

dozens of technical names, such as

apophysis, conchiolin, eolis, and

veliger.

Produced and published by R. Tucker Abbott

Sponsored by the Delaware Museum of Natural History

Greenville, Delaware 19807

This is a high-fidelity, long-playing (331/3 RPM) recording

of the names and terms found in Dr. Abbott's new and

popular book, Seashells of North America (Golden Press,

N.Y.). Born in Boston, bred in Canada and a graduate of

Harvard College, Dr. Abbott has blended the customary

pronunciations of the top professional malacologists in the

United States and Canada that set a high standard of

acceptability. All names and terms are printed on the record

jacket. A unique and long-lasting aid to your study of shells

and biology, for only $4.95 (postage paid).

Obtainable from your favorite shell dealer

WILLIAM H. WEEKS SHELL COLLECTION: New price lists

of this famous collection, with full scientific data, are in prepa-

ration. Many new additions of fine and rare species are also

included. To obtain free copies write:

George E. Jacobs, 853 Riverside Drive, New York 32, N. Y.

WORLD WIDE SPECIMEN SHELLS for sale. New 1970

Price List on request. "Illustrated Catalog of Popular Mar-

ginella Species" now available also, 117 species shown with

full data and values listed with cover in color. Price $2.00.

PHILLIP W. CLOVER

Apartado de Correos 22

Rota Cadiz, Spain

r

Vol. 83 APRIL, 1970 No. 4

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EDITORS AND PUBLISHERS

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Charles B. Wurtz, Biology Department

La Salle College, Philadelphia, Pa. 19141

Marine Biological Laboratory

CONTENTS MAY U1970

Fritz Haas, 1886-1969. By Alan Solem

woods rule; m/K2.

Shell variability in Polygyra dorfeuilliant

By Branley A. Branson 120

Latirus varai, a new fasciolariid gastropod from the Caribbean.

By Robert C. Bullock 133

Maintenance of land mollusks in the laboratory.

By W. Patrick Carney 1 36

Histological studies of the nephridium and pericardial lining

of Quadrula nodulata. By P. R. Myers and D. S. Franzen 139

Another record of insect dispersal of an Ancylid snail.

By Joseph Rosewater 144

Notes 145 News 146

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THE NAUTILUS

Vol. 83 April, 1970 No. 4

FRITZ HAAS, 1886-1969

By ALAN SOLEM

Field Museum of Natural History

Chicago, Illinois 60605

The death of Fritz Haas on December 26, 1969, ended a malaco-

logical career of sixty-one years that spanned two continents. From

1911 until mid- 1936 he was at the Natur-Museum Senckenberg,

Frankfurt-am-Main, Germany, first as Assistant Keeper, then as

Keeper of Invertebrate Zoology. Forced to leave by Nazi perse-

cution, it was fortunate that on August 1, 1938 he could start a

second career as Curator of Lower Invertebrates at Field Museum

of Natural History, Chicago. Although he became Curator Emeri-

tus on January 1, 1959, he spent every working day at the Museum

until a stroke in December 1965 slowed him considerably. Even

after moving to Florida in October 1968, a continual stream of

letters and notes kept him in contact with Museum affairs.

Fritz Haas was born January 4, 1886 in Frankfurt-am-Main and

received his biological training and outlook through the herpe-

tologist and malacologist Oscar Bottger. His life long interest in

and love of the unionid clams came from the influence of Wilhelm

Kobelt, who he succeeded as Keeper at the Natur-Museum Senc-

kenberg. During his years at Frankfurt he enjoyed use of match-

less library facilities and unrivaled collections. Even an unexpected

four year exile in Spain during World War I was turned to

scientific use. Almost 2,000 published pages on the mollusks of

Catalonia resulted from cooperative work with Arturo Bofill dur-

ing this exile. In the 1920's he began his classic section "Bivalvia"

for Bronn's Klassen und Ordnungen des Tierreichs. Eventually

this was completed in 1956 after reaching 2,400 published pages.

Numerous descriptive and faunistic papers on unionids, plus de-

scriptive reports on expedition materials were written during his

Frankfurt years together with "Bau und Bildung des Perlen"

(1931) and "Fauna Malacologica Terrestre y de Agua Dulce de

Cataluna" (1929).

117

118 nautilus Vol. 83 (4)

On arriving in Chicago he had to establish a research library

and reference collections where none had existed. Until 1956 he

had only casual student help or a summer assistant, with all

routine work being done personally. By 1954 die collection con-

tained 54,000 catalogued sets. With the addition of myself as

Assistant Curator in 1956 and a full-time assistant in 1959, growth

became much more rapid. Even in his retirement years, Fritz

Haas spent half-time in routine identifications and cataloguing.

The current 165,000 catalogued sets of mollusks and fine library

in Field Museum of Natural History stand as a monument to

his years of labor. His first years in Chicago saw comparatively

little finished research, but growing collections from South Amer-

ica led to numerous descriptive studies. Short field trips to various

North American localities also resulted in faunistic publications.

As a climax to his career, the editors of "Das Tierreich" asked

him to prepare a species level monograph of the freshwater mus-

sels. Manuscript for this was completed a few weeks before his

stroke, and the 663 page monograph was issued in January 1969.

Later the same year saw publication in the Treatise on Inverte-

brate Paleontology of his generic revision of the unionids. This

had been submitted for publication in 1950, which surely is close

to a record delay.

October 20, 1967 marked the beginning of his sixtieth year

as a publishing scientist. This was celebrated by issuing a review

of his career, together with a list of his publications and new taxa

(Fieldiana: Zoology, 53, pp. 69-144), plus presentation of con-

gratulatory letters from 125 malacologists and colleagues. His

productivity totaled more than 300 articles and 385 new mollus-

can taxa.

Outlines of a career tell little of a man. Fritz was a kindly and

gentle person, with a keen wit and quiet humor. He was raised in

a different tradition from that common in American scientific

circles. The sharp exchanges and testings of ideas and theories

that are part of our scientific meetings and discussions were alien

to his pattern. Work was a personal search for truth. The test of

time and subsequent work would judge the success of this effort.

His manuscripts were not shown to others prior to publication

and rarely would he comment on new publications. During twenty-

five years of association, he would make only gender or diacritical

April. 1970

NAUTILUS

19

mark corrections on my manuscripts. After seeing an absymally

poor paper on unionids, he once made the worst criticism I ever

heard from him — "He gives some data that will be useful some-

day."

This seeming aloofness from his colleagues and profession often

led to misunderstandings and problems. After having unrivaled

collections at Frankfurt, it was hard for him to adjust to the

total lack of such at Chicago. His work pattern seldom involved

consulting collections at other museums and some of his papers

on Nearctic material were unfortunate. Unionid and South Amer-

120 nautilus Vol. 83 (4)

ican studies were on much firmer ground and will stand the test

of time far better.

In later years his work became more introspective and repre-

sented advances from his previous studies rather than from the

general state of knowledge in a field. Thus, his "Das Tierreich"

monograph was a development from his "A Tentative Classifica-

tion of the Palearctic Unionids" (1940) and not a completely

modern synthesis. It presents the ideas developed in fifty years of

work on a group and remains an invaluable landmark.

Much more than in Europe, museum malacology in America has

been bound with amateur shell collectors. Fritz never understood

the collector's outlook and brief contacts were frequent mutual

disasters. Yet a generation of volunteer workers at Field Museum

passed from initial fright to delight, affection and respect for him

after their initial shocks.

Fritz Haas was a scholar and a gentleman, whose legacy is in

knowledge, research facilities and enriched lives.

SHELL VARIABILITY IN POLYGYRA DORFEUILLIANA

By BRANLEY A. BRANSON

Eastern Kentucky University

Richmond, Kentucky 40475

Polygyra dorfeuilliana Lea, probably a trans-Mississippian deriv-

ative of P. troostiana Lea (Tennessee River Drainage) , ranges

through Missouri, southeastern Kansas, the eastern two-thirds of

Texas and Oklahoma, into the northwestern portion of Louisiana.

Pilsbry and Ferriss (1906) compared shells from widespread locali-

ties in the snail's range, and Pilsbry (1940) expanded their com-

ments and somewhat re-evaluated concepts of the species. Pilsbry

and Ferriss (loc. cit.) indicated that the subspecies P. dorfeuilliana

sampsoni Wetherby was restricted to the Ozarkian portion of

Kansas, Missouri, and Oklahoma, stating that there was, however,

"no line to be drawn between them (referring to sampsoni and

typical dorfeuilliana), as in many places both forms and the inter-

grades occur together" (parentheses mine) . They restricted P. d.

percostata Pilsbry to the Red River area of southwestern Arkansas,

whereas P. d. perstriata Pilsbry and Ferriss was "another incipient

April, 1970

NAUTILUS

121

race . . . from Tushkahoma and Potcau, Oklahoma." Polygyra

dorfeuilliana dorfeuilliana Lea supposedly occupied die remainder

of die general range.

According to Pilsbry (1940) , the holotype of P. dorfeuilliana

^r-\

Figure 1. Population sampling areas in Arkansas, Missouri and Oklahoma

in Polygyra dorfeuilliana. See text for explanation.

122

NAUTILUS

Vol. 83 (4)

has nearly a smooth lower surface, the growth striae being very

fine and crossed by microscopic, impressed spiral lines. Through-

out the range, however, both striate and smooth shells were found

within single lots. In the typical race, the umbilicus varied by

imperceptible stages from moderately narrow to the wide-open

one in the nominate form sampsoni. Pilsbry (1940) allowed Poly-

gyra dorfeuilliana sampsoni to stand as a subspecies, although the

form admittedly did not have a definite range exclusive of P. d.

dorfeuilliana. In sampsoni, the shell is supposedly more depressed

than in dorfeuilliana, and the umbilicus is broader in the last

whorl; striation on the base is weak to almost obsolete. According

to Pilsbry (1940) , shells from northeastern Oklahoma possess

heavier striations than ones from Eureka Springs, Arkansas (type

locality) .

The two remaining "races" were deleted by Pilsbry (1940) as

recognizable subspecies. In P. d. perstriata, the umbilicus was sup

posed to be widely open, as in sampsoni, and the base was finely

and densely striate; perstriata was considered as a locally abundant

"mutation" which occurred sporadically throughout the species'

range. In percostata, the riblets of the upper surface continued

onto the base, which was strongly rib-striate.

Branson (1959) , in a general discussion of the biotic divisions

of Oklahoma, stated that P. dorfeuilliana tended to break into

Figure 2. Shell dimensions selected for evaluation in Polygyra dorfeuilliana.

A, basal view: a, diameter aperture; b, width parietal tooth; c, length parietal

tooth; d, shell diameter; B, shell height.

April, 1970 nautilus 123

many local populations, and he considered the form sampsoni to

be an adaptive response to habitats in wooded ravines along

stream margins; percostata was more or less typical of the Quachita

Mountains (Lower Carolinian); and perstriata ranged along the

western Ozarks. Three years later, after studying large series of

shells, Branson (1962) wrote: "Specimens with the characters of

the various "subspecies" can be found throughout the range of

the species, and thus are largely imaginary. Polygyra d. sampsoni

and P. d. perstriata should be relegated to the synonymy of P.

dorferilliana."

Thus, this is the state of knowledge in Polygyra dor] cuilliana ,

and most of it is based upon judgments rather than actual inves-

tigation. Over the years, many arguments have been advanced con-

cerning local molluscan populations ("microgeographical races,"

or "infraspecies," auct) . Since certain aspects of these arguments

are germain to die present work, they are briefly reviewed here.

Some time ago, working with insular (Hawaii, Societies, etc.)

representatives of the Achatinellidas and Partulidae, Clench (1954)

noted that minor changes in shell morphology and coloration

occur from one mountain ridge and valley to another, and he

considered such variation as developed clines. This same type of

variation can be noted in polygyrid snails of die Rocky Mountains

in the United States (See Pilsbry's 1940-1946 monographs) . It is

these local "microgeographical races" which many authors in the

past have described as species or subspecies. In this regard, Burch

and Thompson (1958) were of the opinion that many mollusks

tend to form large numbers of local populations isolated because

of sedentary habits and wide distribution. Such populations may

be morphologically distinguished from one another, and their

characters either genetically determined or ecophenotypically con-

ditioned. Moreover, genetically independent characters do not nec-

essarily vary gradually from one extreme to another at opposite

ends of clines, although definite steps may be obvious along the

variation gradient (Edwards, 1954; 1956) . This morphological

variation may be correlated with environmental influence upon

genetically fixed characters. In the opinion of Hubendick (1951),

all of these populations and microgeographical races comprise the

entity tenned "species," and it is absurd to favor each of them

with subspecific epithets.

124 nautilus Vol. 83 (4)

Clarification of the systematics at the subspecific level depends

upon understanding the type and extent of variation which occurs

in the characteristics normally utilized to recognize and describe

the so-called races. In the Polygyridae, these characteristics are at

present conchological. It is the purpose of this investigation to

demonstrate multi-directional variation in the shell of Polygyra

dorfeuilliana correlated with geography, and to show the difficulty

of recognizing definitive races within the species based upon such

characters.

Methods

Shells from live and dead specimens were secured from 40

localities in five population centers (Fig. 1), representing a large

number of local conditions. Six mensurable conchological features

(Figs. 2,3) and four indices (figs. 9,10,11,12) were compared.

This technique was utilized by Richards and Ferguson (1965)

in Australorbis.

Measurements were made by means of needle-nosed calipers

and a steel metric scale calibrated to 0.5 mm, and estimations were

made to 0.1 mm. Whorl number was determined by making cam-

era lucida sketches of shells, following which measurements were

made by means of a polarizing planimeter.

Statistical evaluation of data, range, mean, standard deviation,

standard error, and coefficient of variation, follows Snedecor

(1956) , and data-graphing follows Hubbs and Hubbs (1953) . In

general, it is probably safe to assume that when the black bars

derived from the data of various populations do not overlap that

the difference between means is not due to chance alone but

represents actual differences between populations.

Population Sampling Areas

For the purpose of analysis (variation associated with geog-

raphy) eastern Oklahoma was divided into four regions (Fig. 1),

with some samples coming from adjacent Arkansas and Missouri,

based upon natural geographic phenomena. These are listed be-

low with the collecting data for specimens secured in each region.

Area A. The Arbuckle Mountains fonn a low dissected plateau

sloping gently toward the southeast (Weidman, 1922) . The some-

what triangular plateau, from 1300 feet elevation in the west to

750 feet in the east, encompasses about 800 square miles. Usually,

April, 1970 nautilus 125

only the highest part west of the Washita River in Murray county,

Oklahoma, is referred to as the Arbuckles, and it was from this

region that my samples were collected. Surrounding the older rocks

of the plateau are Pennsylvania and Permian strata some 100 to

200 feet lower. The mountains are more or less surrounded by

plains.

Collecting sites: (a.) 29 February 1964; 31 specimens, 0.5 mile

north of main gate, Piatt National Park, Murray county, Okla-

homa, (b.) 29 February 1964; 69 specimens, Ordovician talus,

6.1 miles south of Turner Falls, U.S. Highway 77, Carter County,

Oklahoma. N = 100 utilized.

Area B. The so-called Osage Biotic District of Blair and Hub-

bell (1938) is a rather distinct subdivision of Dice's (1943) Texan

Province. The physiography is uneven, with low rounded hills

and some eroded ridges. The dominant vegetation is tall grasses

and oak-hickory associations.

Collecting sites: (a.) 28 August 1963; 67 specimens; 0.8 mile

north of Childers, U.S. Highway 60, Nowata County, Oklahoma.

(b.) 28 August 1968; 193 specimens; 4 miles east of Nowata, U.S.

Highway 60, Nowata County, Oklahoma, (c.) 13 September 1963;

44 specimens; 0.5 mile east of Keystone Dam, Arkansas River

Bluffs, old Oklahoma State Highway 51, Tulsa County, (d.) 27

August 1963; 5 specimens; Sand Creek Bluffs, Osage Hills State

Park, Osage County, Oklahoma; (e.) 25 August 1963; 12 speci-

mens; bluffs overlooking a small pond near main entrance, Osage

Hills State Park, Osage County, Oklahoma. N = 321 utilized.

Area C. The Ozark District, sharply separated from the Osage

by the Cherokee Plains, and the flood plains of the Neosho River,

is a more or less well-known uplift of mainly Mississippian and

Pennsylvanian rocks. To the south these mountains are bordered

by plains scarcely fOO feet above sea level, and are separated from

Area D. by the broad valley of the Arkansas River.

Collecting sites: (a.) 24 August 1963; 1 specimen; Neosho River

bluffs, Cherokee Recreation Area #1, Langley, Mayes County,

Oklahoma, (b.) 17 May 1964; 5 specimens; 3 miles east of

Pineville, U.S. Highway 71, McDonald County, Missouri, (c.)

26 August 1963; 8 specimens; 5 miles east of Pineville; (d.) 26

August 1963; 35 specimens; near Disney Dam, Oklahoma High-

way 20, Mayes County, (e.) 4 April 1963; 7 specimens; 7.4 miles

126

NAUTILUS

Vol. 83 (4)

S HELL DIAMETER

6 0 6-5 70 75 8.0 85 9.0 9.5 10.0 10.5 1 1.0

C=0.040

C = 0-072

C = 0.094

C = 0.I02

C = 0.06l

Figure 3. Shell diameter in five populations of Polygyra dorfeuilliana. Let-

ters on vertical axis correspond to populations delimited in text, c = coefficient

of variation. Measurements in mm. Black bar = two standard errors on each

side of mean; hollow bar = one standard deviation on each side of mean;

shell height

Figure 4. Shell height in five populations of Polygyra dorfeuilliana. Ab-

mean — vertical line; range = horizontal line,

breviations and incidentals as in figure 3.

2 5

A

B

C

D

E

DIAMETER OF APERTURE

30 3-5

C=0065

4;0

C = 0.79

C = 0.084

C=0.I03

C = 0.050

Figure 5. Apertural diameter in five populations of Polygyra dorfeuilliana.

Abbreviations and incidentals as in figure 3.

April, 1970

NAUTILUS

127

3:0

C=0.0I5

Figure 6. Height of parietal tooth in five populations of Polygyra dorfeuil-

liana. Abbreviations and incidentals as in figure 3.

50

5-25

WHORL NUMBER

5 5

575

__i

60

00.036

C=0.034

C= 0.041

C = 0.036

C'0.039

Figure 7. Width of parietal tooth in five populations of Polygyra dorfeuil-

liana. Abbreviations and incidentals as in figure 3.

i.o

WIDTH OF TOOTH

15 2.0

rfc

00 045

C= 0.132

C= 0 141

C-= 0.075

2 5

C= 0.169

Figure 8. Whorl number in five populations of Polygyra dorfeuilliana. Ab-

breviations and incidentals as in figure 3.

128

NAUTILUS

Vol. 83 (4)

0 5

0.6

WIDTH OF TOOTH/HEIGHT OF TOOTH

0.7 0.8 0.9 1.0 12

1,3

1:4

C = 0.070

C = OIIO

C = 0.I45

O0.I55

2 C = 0.260

Figure 9. Width of parietal tooth/Height of parietal tooth in five popula-

tions of Polygyra dorfeuilliana. Abbreviations and incidentals as in figure 3.

HEIGHT OF SHELL/DIAMETER OF SHELL

0.3 04 0.5 0.6 0.7

C=0205

C-0.064

C= 0.117

C» 0.144

C = 0.085

Figure 10. Height of shell/Diameter of shell in five populations of Polygyra

dorfeuilliana. Abbreviations and incidentals as in figure 3.

WIDTH OF TOOTH/HEIGHT OF TOOTH/DIAMETER OF APERTURE

0.1 0.2 0.3 0.4 0.5

O0.I02

C = O.I37

C= 0.179

C=0.I96

E Ural C=0.09l

Figure 11. Width of parietal tooth/Height of parietal tooth/Diameter of

aperture in five populations of Polygyra dorfeuilliana. Abbreviations and

incidentals as in figure 3.

J°ii_

WHORL NUMBER/SHELL DIAMETER

0.5 0.6 0 7

0.8

09

C*0.03l

C = 0.054

=Z1

C = 0 104

C=0 089

C = 0.04l

Figure 12. Whorl number /Shell diameter in five populations of Polygyra

dorfeuilliana. Abbreviations and incidentals as in figure 3.

April, 1970 nautilus 129

east of junction of Oklahoma Highways 259 and 59, Adair County,

(f.) 25 August 1963; 11 specimens; 5.5 miles east of Salina, Okla-

homa Highway 82, Mayes County, (g.) 26 June 1963; 7 specimens;

1 mile south of Lanagan, U.S. Highway 71, McDonald County,

Missouri, (h.) 24 August 1963; 40 specimens; east end Tenkiller

Dam, Sequoyah County, Oklahoma, (i.) 26 August 1963; 10

specimens; hills above Spavinaw Creek, Spavinaw, Mayes County,

Oklahoma, (j.) 21 September 1963; 25 specimens; 0.1 mile north,

2 miles east of Turkey Ford, Delaware County, Oklahoma, (k.)

24 August 1963; 66 specimens; west side of Tenkiller Dam,

Sequoyah County, Oklahoma. (1.) 13 May 1963; 28 specimens;

Lanagan, McDonald County, Missouri, (m.) 20 August 1963; 114

specimens; 15 miles east of Cassville, Missouri, (n.) 4 June 1963;

3 specimens; 7.4 miles east of Jet, Adair County, Oklahoma, (o.)

25 August 1963; 4 specimens; 1.2 miles east of Flint, Delaware

County, Oklahoma, (p.) 16 May 1963; 3 specimens; Lost Creek,

15 miles southeast of Miami, State Highway 10, Ottawa County,

Oklahoma. N = 246 utilized.

Area D. Quachita Mountains. As indicated, these mountains

are separated from the Ozarks by the Arkansas River valley to

the north, and they are bordered southward by the Austroriparian

lowlands, and in the west by plains.

Collecting sites: (a.) 31 May 1963; 12 specimens; 3 miles south

of Foreman, Sevier County, Arkansas, (b.) 22 August 1963; 9

specimens; 0.5 mile east, 1.5 miles north of Muse, Oklahoma High-

way 63, LeFlore County, Oklahoma ("Billy Creek Recreation

Area") . (c.) 23 August 1963; 3 specimens; Robbers Cave State

Park, 5 miles north of Willburton, Latimer County, Oklahoma,

(d.) 18 August 1963, Windingstair Mountain, 0.5 mile from junc-

tion of Oklahoma Highways 59 and 259, LeFlore County, Okla-

homa, (e.) 21 August 1965; 91 specimens; Lake Talihina, 2 miles

west of Talihina, Latimer County, Oklahoma, (f.) 21 August

1963; 7 specimens; Kiamichi Mountain, 0.9 mile north of Hono-

bia, Pushmataha County, Oklahoma, (g.) 22 August 1963; 11

specimens; 26.6 miles south of Hartshorn, Jackfork Mountain,

Pittsburg County, Oklahoma; (h.) 19 August 1963; 24 specimens;

1.4 miles southeast of Monroe, Oklahoma Highway 83, Poteau

Mountain, LeFlore County, (i.) 21 August 1963; 45 specimens;

1 mile west of Kiamichi fire tower, Kiamichi Mountain, LeFlore

130 nautilus Vol. 83 (4)

County, Oklahoma, (j.) 21 August 1963; 79 specimens; Potatoe

Hills (Flying "W" Ranch) , 5 miles west of Talihina, Oklahoma

Highway 83, Latimer County, Oklahoma, (k.) 22 August 1963;

55 specimens; Blue Mountain, Holson Valley Road (Branch off

State 271), LeFlore County, Oklahoma. (1.) 20 August 1963; 5

specimens; 0.7 mile south of Page, Oklahoma Highway 259, Le-

Flore County, Oklahoma, (m.) 23 August 1963; 63 specimens;

Cavanal Mountains, 2 miles west of Poteau, LeFlore County,

Oklahoma, (n.) 23 August 1963; 46 specimens; 5.3 miles north

of Wilburton, Oklahoma Highway 2, Latimer County, (o.) 19

August 1963; 35 specimens; 1.4 miles southeast of Monroe, Okla-

homa Highway 83, Poteau Mountain, LeFlore County, (p.) 19

August 1963; 15 specimens; Walker Mountain, 11.1 miles east,

2.8 miles south of Page, LeFlore County, Oklahoma. N = 618

utilized.

Area E. Plains between Quachitas and Arbuckles.

Collecting sites: (a.) 27 August 1963; 63 specimens; Rock Creek

Bluffs, 2 miles south of Eufala, U.S. Highway 69, Mcintosh

County, Oklahoma, (b.) 27 August 1963; 133 specimens; 19 miles

north of Atoka, U.S. Highway 69, Atoka County, Oklahoma.

N = 196 utilized

Results

Comparison of shell diameters in die series of P. dorfeuilliana

demonstrated considerable variation (Fig. 3) , with the smallest

shells coming from the Arbuckles and the largest ones from the

Ozarks and Quachitas. Comparison of the means possibly indi-

cates the presence of a weakly developed cline extending from

the more arid regions toward the east (particularly if the data

gleaned from population E is lumped with that from D) : A . . .

B . . . C . . . D (E) . In the field, specimens from area A appeared

smaller than those secured elsewhere. Moreover, shell-height

differences were obvious by inspection.

Shell heights (Fig. 4) varied rather greatly, but there is only

slight to no overlap in the means of populations A and E with

the means of shells taken from other regions, thus indicating a

tendency for shells to become more elevated in the southern part

of the range. The ratio height/diameter (Fig. 10) reflects the

flatter shells from areas B and C and a trend toward intermediacy

in ones from area D.

April, 1970 nautilus 131

In snails with indeterminate growth patterns, as in Polygyra, total

diameter is obviously a function of whorl number. However, the

thickened flange (lip) produced at the aperture of most poly-

gyrids also influences the diameter. Whorl number varies within

a rather narrow spectrum (Fig. 7) after the lip is produced. It

will be noted that western specimens (with no overlap) tend to

produce from a quarter to one-half less whorl than more easterly

populations. When whorl number is viewed as a percentage of

the total diameter (Fig. 12) a similar picture emerges, but the

populations tend to smooth-out somewhat.

In general, although the range of variation was considerable,

there seemed to be little difference between the means of apertural

diameters (Fig. 5) , except in specimens from area E. Deviations

in this population were significant at the 0.01 level (student "t"

distribution) . Within the aperture are two deeply immersed lip

denticles ("teeth") , and standing in front of the aperture, often

nearly blocking it, is a tongue-like parietal tooth (Fig. 2) which

varied from 1.0 mm to 3.0 mm in height, being significantly

shorter in shells secured in region D (Fig. 6) . The width of this

denticle (Fig. 8) is also variable, but there does not appear to

be much difference between the means of the various groups in-

vestigated. The ratio width of tooth/height of tooth (Fig. 9)

demonstrates the same type of variation in range, but no significant

differences among means; the variation is concordant. On the

width of tooth/height of tooth

other hand, the ratio again indi-

diameter of aperture

cates a significant difference in specimens from area D. In other

words, consistent differences in the form of the parietal tooth seem

to be correlated with those observed in the aperture.

Discussion

Most of the variable characteristics described above are con-

sidered parameters of microgeographical races, and many of them

are probably genetically determined, although some traits may

be ecophenotypic responses to local conditions. It is possible, when

observing only one or a few mensurable characters, to show non-

overlap in mean differences, and hence one might be tempted to

assign a given population a subspecific epithet. This is apparent

132 nautilus Vol. 83 (4)

in shells collected from area D. However, more characteristics over-

lap than not. Furthermore, when features other than those dis-

cussed above are observed, such as external sculpture and colora-

tion, specimens from area A (especially from the Osage region)

are more strikingly different than ones from area D. In my esti-

mation, then, to reiterate a previous statement, the subspecies of

Polygyra dorfeuilliana, as such, are fictional. Looking at samples

secured from widely separated regions may seem to indicate real

differences, but when these differences are subjected to scrutiny,

they are seen to be only discordant variation associated with the

sendentary habits of the snail.

Summary

Detailed morphologic analyses were performed on 1,481 shells of

Polygyra dorfeuilliana secured from 40 localities in five major pop-

ulation centers in Oklahoma, Missouri and Arkansas. Such charac-

ters show considerable variation, some of them (shell diameter and

height) seem to vary clinally from west to east and north to south,

respectively. Other characteristics, whorl number, apertural diam-

width of parietal tooth/height of parietal tooth

eter, and the ratio

diameter of aperture

show consistent differences between some populations. These are

considered to be examples of discordant variation.

Literature cited

Blair, W. F. and T. H. Hubbell. 1938. The biotic districts of

Oklahoma. Amer. Midi. Nat. 20: 425-454.

Branson, B. A. 1959. The Recent Gastropoda of Oklahoma. Part

1, historical review, general comments and higher taxonomic

categories. Proc. Oklahoma Acad. Sci. 39: 21-37.

Branson, B. A. 1962. The Recent Gastropoda of Oklahoma, Part

IV. Terrestrial species, families Polygyridae and Bulimulidae.

Proc. Oklahoma Acad. Sci. 42: 60-80.

Burch, J. B. and F. G. Thompson. 1958. Taxonomic characters

and infraspecific variation in Mollusks. Syst. Zool. 7: 48.

Clench, W. J. 1954. The occurrence of clines in molluscan popu-

lations. Syst. Zool. 3: 122-125.

Dice, L. R. 1943. The Biotic Provinces of North America. Univ.

Mich. Press, Ann Arbor, 78p.

Edwards, J. G. 1954. A new approach to infraspecific categories.

Syst. Zool. 3: 1-20.

April, 1970 nautilus 133

Edwards, J. G. 1956. Clarification of certain aspects of infraspecific

systematics. Syst. Zool. 5: 92-94.

Hubbs, C. L. and Clark Hubbs. 1953. An improved graphical

analysis and comparison of series of samples. Syst. Zool. 2: 49-

56; 92.

Hubendick, B. 1951. Recent Lymnaeidae, their variation, mor-

phology, taxonomy, nomenclature, and distribution. Kungl.

Svenska Vetenskad. Handl. 3: 1-223; 5 pi.

Pilsbry, H. A. 1940. Land Mollusca of North America (North of

Mexico) . Acad. Nat. Sci. Philadelphia Mongr. 3 (2) : i-viii;

575-994; i-ix.

Pilsbry, H. A. and J. H. Ferriss. 1906. Mollusca of the Ozarkian

fauna. Proc. Acad. Nat. Sci. Philadelphia 1906: 529-567.

Richards, C. S. and F. T. Ferguson. 1965. Variability in Austra-

lorbis glabratus (Say) . Trans. Amer. Micros. Soc. 84: 580-587.

Snedecor, G. W. 1956. Statistical Methods. Iowa State College

Press, Ames.

Weidman, S. 1922. Physiographic history of the Arbuckle Moun-

tians. Proc. Okla. Acad. Sci. 2: 74-17.

LATIRUS VARAI, A NEW FASCIOLARIID

GASTROPOD FROM THE CARIBBEAN

By ROBERT C. BULLOCK

Museum of Comparative Zoology, Harvard University

Cambridge, Massachusetts 02138

Species of the genus Latirus (Gastropoda: Fasciolariidae), though

well represented in the West Indian faunal province, are not

commonly collected alive. The numerous taxonomic problems that

exist within the group are related to the considerable intraspecific

variation of each species. In a recent study of the genus Latirus

of the western Atlantic (Bullock, 1968) , a new species was noted

and is herein described:

Latirus (Latirus) varai, new species (Figure 1)

Description: Shell large, up to 70 mm in length and 27.8 mm in

greatest diameter. Spire slightly higher than half the length of

the shell. Whorls 10, including 2 whorls of the protoconch. Aper-

ture squarish in the mature specimen examined and about the

same length as the siphonal canal. Axial sculpture of 7-8 ribs

per whorl and numerous fine growth lines. Spiral sculpture of

134 nautilus Vol. 83 (4)

13-15 cords which are more pronounced as they pass over the axial

ribs. 9-1 1 spiral cords on the upper whorls. Columella with 4

folds, the upper one weaker than the others. Outer lip slightly

crenulate, with about 15 irregular, sometimes beaded, lirae run-

ning into the aperture. Fasciole well developed, pseudo-umbilicus

slit-like. Periostracum unknown.

Shell white to tawny brown in the areas of the sutural ramp

and the neck. Axial ribs light chestnut brown and crossed by

white spiral cords. Aperture white.

Measurements: Holotype 70.0 mm in length, 27.8 mm in great-

est diameter; Paratype (also from the type locality) 52.4 mm in

length.

Type locality: Off Gibara, Oriente Province, Cuba in 100 fath-

oms.

Type depositories: Holotype, Museum of Comparative Zoology,

Harvard University 262589; paratype in the collection of Mr. John

Finlay of Wilmington, Delaware (both types collected dead) .

Remarks: This species is unlike any other West Indian Latirus.

The only other member of Latirus s.s. from the Caribbean is the

highly variable Latirus distinctus A. Adams, 1855 (= Turbinella

trochlearis Kobelt, 1874; L. mcgintyi Pilsbry, 1939; and possibly

Fusus cariniferus Lamarck, 1816). L. varai may easily be dis-

tinguished from L. distinctus by its stronger spiral sculpture,

more rounded whorls, and especially by its possession of the

chestnut brown coloration on the axial ribs, not between them.

In comparison with other species, L. varai appears to be related

to Latirus kandai Kuroda, known only from Kii Peninsula, Hon-

shu, Japan (Habe, 1964) .

This species is named in honor of Senor Joaquin de la Vara

of Gibara, Oriente Province, Cuba, who collected the specimens.

Acknowledgments

Both specimens of L. varai belonged to Mr. John Finlay of

Wilmington, Delaware, who kindly donated the holotype to the

Museum of Comparative Zoology. My dissertation of western

Atlantic Latirus was done under the direction of Dr. John H.

Dearborn and presented to the Zoology Department of the Uni-

versity of Maine at Orono, for partial fulfillment of the require-

ments of the Master of Science degree.

April, 1970

NAUTILUS

1.85

Figure 1. Latirus (Latirus) varai Bullock, new species. Holotype, off Cuba.

Length: 70 mm.

References cited

Adams, A. 1855. Descriptions of twenty-seven new species of shells

from the collection of Hugh Cuming, Esq. Proc. Zool. Soc.

London for 1854: 311-317.

Bullock, R. C. 1968. The fasciolariid genera Latirus, Dolicho-

latirus, and Teralatirus (Mollusca: Gastropoda) in the western

Atlantic. Unpublished M.S. thesis, University of Maine, 109 p.,

8 pis., 2 figs.

Habe, T. 1964. Shells of the Western Pacific in Color, 2. Osaka,

233 p., 66 pis.

Kobelt, W. 1874. Turbinella and Fasciolaria [in] Martini-Chem-

nitz, Systematiscb.es Conchylien-Cabinet, 3 (3A): 65-88, pis. 8,

figs. 17-21.

Lamarck, J. 1816. Tableau encyclopedique et methodique des

trois regnes de la nature. Paris, 97 pis.

Pilsbry, H. A. 1939. A triad of umbilicate Latirus, Recent and

Pliocene. Nautilus, 52 (3) : 84-86, 1 pi.

136

NAUTILUS

Vol. 83 (4)

MAINTENANCE OF LAND MOLLUSKS IN THE

LABORATORY

By W. PATRICK CARNEY

Department of Zoology, University of Montana

Missoula, Montana 59801

Since molluscan hosts of medically important trematodes are

predominantly aquatic, most accounts of snail raising relate to

aquatic snails. Only recently has the importance of land snails as

vectors for human or domestic animal helminths been realized.

Terraria and techniques for the cultivation of land snails have

been developed by numerous workers (Carmichael, 1937; Archer,

1937; Krull, 1937; Denton, 1944; Peterson, 1947; Webb, 1947;

Tang, 1950; Mapes and Krull, 1951; Ulmer, 1951; Patten, 1952;

Rowan, 1955; Pillmore, 1958; Hohorst and Lammler, 1962; For-

rester, 1962; Kingston, 1965; Kingston, 1966) . Because of the

growing recognition of zoonotic diseases involving land snail hosts,

additional techniques for laboratory maintenance of land molluscs

outfit- i'

Figure 1. Basic water table construction. Each level was lined with plastic.

Water was then allowed to flow onto the table to a depth of approximately 2".

The siphon apparatus in Figure 2 maintained a constant water level.

April, 1970

NAUTILUS

137

should be of interest. The methods described below for raising a

variety of land mollusks in large numbers have the advantages of

being inexpensive and requiring a minimum of maintenance.

Colonies were maintained in a basement laboratory provided with

a floor drain. Temperature control was achieved by utilizing a

shallow, plastic-lined water table of approximately 150 square feet

of surface and one to two inches deep (Figure 1) . City water, nor-

mally varying in temperature from 9°C to 15°C, was allowed to

flow slowly onto the water table. A glass siphon apparatus (Fig-

ure 2) kept the water level constant barring extreme variations

in pressure.

Snails were bred and raised in various types of plastic containers

available in local stores for domestic use. Most of these were either

round, shallow tubs approximately 12 inches in diameter or rec-

tangular and square boxes 6-12 inches on a side. Each container

held one inch of moist sand, serving as a source of minerals and as

a site for egg laying. One or two layers of damp laboratory paper

towelling covered the sand. The containers were immersed to a

depth of one-two inches in the water bath. For breeding and cul-

Figure 2. Glass siphon apparatus which maintains a constant water level

on water table at a level parallel with point (A) . The plastic lining is

indicated at (B) .

138 nautilus Vol. 83 (4)

ture of Allogona ptychophora and Triodopsis mullani, two previ-

ously soaked pumice bricks were placed on the paper towelling.

The container was then closed with a one-ply glass lid. For smaller

snails, Discus cronkhitei, Zonitoides arboreus, and Z. nitidus and

the slugs Deroceras laeve and D. reticulatum decaying wood was

substituted for pumice bricks. All species were fed lettuce and

carrots sparingly. Occasionally a piece of chalk was added for

calcium supply. Containers were inspected at least once each week

at which time the paper towelling was changed, old food removed,

and fresh food provided.

Although their shells never attained the thickness of those col-

lected in the field. Laboratory raised Allogona ptychophora reached

maturity and attained normal size in one year. The thin shells of

laboratory raised specimens, although quite fragile, proved an

advantage in studies of trematode infections since the hepatopan-

creas could be observed through the shell. Laboratory and natural

raised specimens of T. mulloni, D. cronkhitei , Z. arboreus and

Z. nitidus showed no significant difference in shell thickness.

Acknowledgements: Appreciation is extended to Dr. Glen R.

Webb for his comments, and to Richard Russell and Sharon

Carney for their technical assistance.

References

Archer, A. F., 1937. Civilization and land mollusks in Ohio. Nau-

tilus 50: 117-121.

Carmichael, E. B., 1937. Culture methods for Limax flavus. pp.

529-531. In J. G. Needham (ed.) , Culture methods for inverte-

brate animals. Dover Publications, Inc., New York.

Denton, }. F., 1944. Studies on the life history of Eurytrema

procyonis Denton, 1942. J. Parasitol. 30: 277-286.

Forrester, D. J., 1962. Land mollusca as possible intermediate hosts

of Protostrongylus stilesi, a lungworm of bighorn sheep in west-

ern Montana. Proc. Mont. Acad. Sci. 22: 82-92.

Hohorst, W. and G. Lammler, 1962. Experimentelle Dicrocoeliose-

Studien. Z. Tropenmed. Parasit. 13: 377-397.

Kingston, N., 1965. On the life cycle of Brachylecithum orfi King-

ston and Freeman, 1959 (Trematoda: Dicrocoeliich.e) , from the

liver of the ruffed grouse, Bonasa umbellus L. Infections in the

vertebrate and molluscan hosts. Can. J. Zool. 43: 745-766.

, 1966. Observations on the laboratory rearing of terrestrial

molluscs. Am. Midi. Natur. 76: 528-532.

Krull, W. H., 1937. Rearing terrestrial snails, pp. 526-527. In J. G.

April, 1970 nautilus 139

Needham (ed.) , Culture methods for invertebrate animals.

Dover Publications, Inc., New York.

Mapes, C. R. and W. H. Krull, 1951. Collection of the snail,

Cionella lubrica, and its maintenance in the laboratory. Cornell

Vet. 41: 433-444.

Patten, J. A., 1952. The life cycle of Conspicuum icteridorum

Denton and Byrd, 1951 (Trematoda: Docrocoeliidae) . }. Para-

sitol. 38: 165-182.

Peterson, A., 1947. A manual of entomological equipment and

methods. Edwards Brothers Inc., Ann Arbor, Mich. (c.f. Rowan,

1955).

Pillmore, R. E., 1958. Life cycles of the lungworm genus Proto-

strongylus in Colorado. J. Colo. - Wyo. Acad. Sci. 4: 44-45.

Rowan, W. B., 1955. The life cycle and epizootiology of the rabbit

trematode, Hasstilesia tricolor (Stiles and Hassall, 1894) Hall,

1916 (Trematoda: Brachylaemidae) . Trans. Am. Microscop.

Soc. 74: 1-21.

Tang, C. C, 1950. Studies on the life history of Eurytrema pancrea-

ticum Janson, 1889. J. Parasitol. 36: 559-573.

Ulmer, M. J., 1951. Posthamiostomum helicis (Leidy, 1847) Rob-

inson, 1949 (Trematoda) , its life history and a revision of the

subfamily Brachylaeminae. Part I. Trans. Am. Microscop. Soc.

70: 189-238.

Webb, G. R., 1947. The mating-anatomy technique as applied to

polygyrid landsnails. Amer. Naturalist 81: 134-147.

HISTOLOGICAL STUDIES OF THE NEPHRIDIUM AND

PERICARDIAL LINING OF QUADRULA NODULATA

By PAUL ROBERT MYERS and DOROTHEA S. FRANZEN

Illinois Wesleyan University, Bloomington, Illinois 61701

The mechanism of excretion of the pelecypod excretory system

is not yet fully understood. It is the intention of the authors of

this paper to contribute to the knowledge of this system. Alhough

data concerning the organic and inorganic nature of the excretory

products are available (Picken, 1936) , the actual cellular processes

are still seemingly unexplored. Duwe (1948) has contributed a

limited histological description of the nephridium. Through the

employment of histological and histochemical techniques an at-

tempt has been made here to provide information regarding the

processes involved in excretion.

140 NAUTILUS Vol. 83 (4)

Materials and Methods

Quadrula nodalata (Rafinesque, 1820) is a small freshwater

clam averaging 3.2 cm. in length, inhabiting a sandy to rocky

bottom in shallow water. The specimens for this study were col-

lected in the Mississippi River eight miles north of Quincy, Illi-

nois. Specimens were removed from their shells by severing the

adductor muscles and gently removing the viscera from the shell.

Immediately the clams were fixed in Zenker's or Heidenhein's

Susa fixatives to minimize histological changes. Chemical fixation,

embedding by the paraffin method, and sectioning were by stand-

ard procedures. Histological sections were cut six microns in diick-

ness and stained with Mallory's Triple Stain, Heidenhein's Azan

Stain or Alum Hematoxylin and Eosin Stain according to methods

outlined by Galigher (1964).

Observations

Each of the laterally paired nephridia of the clam is shaped

like a U lying on one side with the open end facing anteriorly.

The ventral stem of the U opens anteriorly into the pericardial

cavity to form the ciliated nephrostome. Laterally and slightly pos-

terior to the nephrostome, the dorsal portion of the U opens into

the suprabranchial chamber to form the nephridiopore (Fig. 1) .

The ventral portion of the nephridium is characterized by many

involuting diverticula, thus accounting for the glandular, spongy

appearance seen in gross dissection. Its epithelium is of the simple

columnar type. The dorsal portion lacks the diverticula and forms

a sac-like structure whose epithelium is of a more cuboidal type.

This sac is histologically and morphologically quite unlike the

ventral, glandular portion, suggests the function of a bladder, and

opens anteriorly into the suprabranchial chamber via the ciliated

nephridiopore. Dorsal to the bladder lies the reduced coelom, or

pericardial cavity, enclosing the heart which is folded around the

intestine. The pericardial lining is comprised of simple columnar

epithelium. Surrounding the nephridium and pericardial cavity

is a large blood sinus, the pericardial sinus, which contains loose

connective tissue, tissue fluid, and blood cells (Fig. 1) . Closely

applied to the basal portion of the columnar epithelium are two

layers of tissue, the innermost a simple squamous epithelium and

the outermost a connective tissue layer. These two layers of tissue

April, 1970

NAUTILUS

141

separate the nephridium and pericardium from the pericardial

sinus.

Results

Histological study of the pericardial and nephridial tissues dis-

closed that three types of epithelial cells are apparently involved

in excretion: (A) Most abundant are ciliated simple columnar

us;> X.' -

Fig. 1. Dorsal x-sec. through pericardial cavity and nephridium. a. auricle;

b. bladder; i. intestine; n. nephridium; n.I. nephridial lumen; p.c. pericardial

cavity; p.s. pericardial sinus; v. ventricle. Fig. 2. Fining of pericardial cavity.

A. cell type A; b.c. blood cell; C. cell type C; c.t. connective tissue; s.e.

squamous epithelium; s.v. secondary vesicle; v. vacuole. Fig. 3. Cells of

nephridium. A. cell type A; B. cell type B; nil. nucleus. Fig. 4. Ventral portion

of pericardial cavity, c.d. cellular debris.

142 nautilus Vol. 83 (4)

cells which are noted as type A (Figs. 2 & 3) . The cells contain

a large, granular, centrally located nucleus with one or two nu-

cleoli. These cells are interspersed at random in the nephridial

epithelial tissue and pericardial lining where they, evidently, func-

tion in movement of material from the pericardial cavity or the

lumen of the nephridium. This type cell lines also the nephridio-

pore and the nephrostome. (B) Another type cell also observed,

noted as type B (Fig. 3) , is columnar and contains a large, non-

granular vacuole which displaces the cell's nucleus laterally and

ventrally to a basal position. The cell appears mainly in the pos-

terior region of the nephridium and occurs less frequently in the

lining of the pericardial cavity. Further observation revealed con-

striction of secondary vesicles into the lumen of the nephridium

or pericardial cavitv. (C) Most relevant to the study is the pres-

ence of a non-ciliated, nucleate, columnar cell abundant both in

the nephridial tissue and lining of the pericardium (Figs. 2 & 4) .

The cell is randomly interspersed between the other two cell types

and, as mentioned earlier, is in close contact with the blood in the

pericardial sinus. Constriction of secondary vesicles was observed

similar to that observed in the type B cell.

Discussion

On the basis of the above observations and results, three types of

simple columnar epithelial cells are apparently involved in an ex-

cretory function. The very nature of the constriction of secondary

vesicles suggests the mechanism of excretion. The cell types have

been presented above as types A, B, and C.

Cell type A is ciliated. Its function is evidently movement of

material from the pericardial cavity toward the lumen of the

nephridium or from the lumen of the nephridium to the supra-

branchial chamber.

Cell type B presented an enigma in that observations could not

confirm its function, but they did suggest a plausible explanation.

The majority of the secondary vesicles are non-granular and very

sensitive to osmotic changes as confirmed by many broken vesicles.

Therefore, the cell is actively enveloping excrement in a mem-

brane and pinching it off in the form of secondary vesicles which

suggests an excretory activity. Differential staining with Heiden-

hein's Azan Stain indicates that the material in the vacuoles is

chemically basic.

April, 1970 nautilus 143

Cell type C did not suggest a glandular function as in type B,

but strongly indicated excretory activity. Fretter and Graham

(1962) reported actual migration of blood cells into the nephri-

dium of certain gastropods, and this, in part, is what was observed.

Blood cells and cellular debris appeared as though they were being

ingested by the cell. Whether the blood cells actually migrate or

whether the excretory cell engulfs it by phagocytosis could not be

definitely determined. There appeared to be various stages of

breakdown of ingested material. Some of the ingested blood cells

were still intact while some were broken down to the extent that it

could not be determined whether this material was partially di-

gested blood cells or some other kind of cellular debris. In later or

more advanced stages of breakdown of the ingested material, the

morphological characteristics of the normal columnar cell changed.

It became more club shaped as though digested material were

being concentrated at one end of the cell. In cases where there was

a large amount of ingested material, a membrane could be ob-

served surrounding a vacuole. Material in the vacuoles stained

blue with Mallory's Triple Stain. In various cells of type C, sec-

ondary vesicles were being constricted and released into the peri-

cardial cavity or the nephridial lumen. These vesicles contained

granular material and were enclosed in a definite membrane. The

change in staining properties of the ingested material as it is

broken down indicates digestive activity by the excretory cell.

Type B cells may also be functional in excretion, but results indi-

cated that the nature of the material excreted is different from that

excreted by type C. It was concluded that the type B cell has a

more glandular-like function. Its resemblance to the familiar gob-

let cell of the vertebrate alimentary tract and also the observance

of the presence of a similar type cell in the mantle prompted

this conclusion.

Summary

Histological examination of the epithelium of the nephridium

and lining of the pericardial cavity of Quadrula nodulata (Rafin-

esque, 1820) revealed excretory activity on the cellular level. Re-

sults indicated that material is ingested by certain cells, enzymati-

cally broken down, and excreted by way of constriction of secondary

vesicles into the nephidial lumen or pericardial cavity. Three

types of cells were observed to comprise the tissue of the nephridium

144 nautilus Vol. 83 (4)

and lining of the pericardial cavity, and the nature of the activity

observed in these cells suggests a mechanism of excretion on the

cellular level.

References

Chandler, A. B., and R. A. Hand, 1961. Phagocytized Platelets: A

Source of Lipids in Human Thrombi and Atherosclerotic

Plaques. Science 134: 946-947.

Duwe, A. E., 1948. The Microscopic Structure of the Kidney of

Freshwater Clams in the Genus Unio. Turtox News 36 (7) : 161-

163, 3 figs.

Fretter, V., and A. Graham, 1962. British Prosobranch Molluscs.

Printed for the Ray Society. Adlard 8c Son Ltd., Bartholomew

Press, Dorking, pp. v-xvi and 2-755, figs. 1-317.

Galigher, A., and E. Kosloff, 1964. Essentials of Practical Micro-

technique. Lea and Febiger, Philadelphia. 484 p.

Gans, H., V. Subramanian, and B. H. Tan, 1969. Selective Phago-

cytosis: A New Concept in Protein Catabolism. Science 159:

107-109.

Picken, L. E. R., 1937. The Mechanism of Urine Formation in

Invertebrates. Jour. Exp. Biol. 14 (1) : 20-34.

ANOTHER RECORD OF INSECT DISPERSAL

OF AN ANCYLID SNAIL

By DR. JOSEPH ROSEWATER

Division of Mollusks

National Museum of Natural History

Washington, D. C. 20560

Two living specimens of Lavapex fusciis (C. B. Adams) (nomen-

clature of Basch, 1959 and 1963) were found attached to the

outer surface of the elytra of a large (33 mm) dytiscid water

beetle, Cybister occidentalis Aub£, taken in a light-trap at the

Archbold Biological Station, Highlands County, Florida, March

7, 1967. The snails are catalogued under USNM 680346; speci-

men lengths are: 6.4 and 6.2 mm; widths: 4.3, 4.0 mm. According

to Dr. S. W. Frost of The Pennsylvania State University, who

sent the snails for identification, the light-trap was close to a

drainage ditch, within a half mile of a swampy area, and about

six miles from a small lake. As shown by Rees (1965) Ancylidae

are the only fresh-water snails known to be transported by insects,

and L. fuscus has been found previously on the elytra of Dinutes

sp. in Wellesley, Massachusetts (Johnson, 1904) . Although Rees

April, 1970 nautilus 145

(ibid.) suggested that insects probably play a role only in the

local dispersal of mollusks, a species could cross a continent or

traverse narrow water barriers between islands by such "puddle-

jumping." Such classical associations are utilized frequently in

explaining the zoogeography and dispersal of species, and it is

refreshing occasionally to witness one of these phenomena in

action.

Literature cited

Basch, P. F. 1959. The anatomy of Lccvapex fuscus, a freshwater

limpet (Gastropoda: Pulmonata) . Miscellaneous Publications,

Museum of Zoology, University of Michigan, no. 108, pp. 1-56,

figs., tbl.

. 1963. A review of the Recent freshwater limpet snails of

North America. Bulletin of the Museum of Comparative Zool-

ogy Harvard, 129: no. 8, pp. 399-461, figs.

Johnson, C. W. 1904. Ancyli adhering to water beetles. The Nau-

tilus, 17: no. 10, p. 120.

Rees, W. J. 1965. Presidential Address. The Aerial Dispersal of

Mollusca. Proceedings of the Malacological Society of London,

36: part 5, pp. 269-282.

NOTES

Corbicula in Baja California — Two staff members of the

Academy, Stanley C. Williams and V. F. Lee, collected a series

of live Corbicula manilensis (Philippi) from an irrigation canal,

one-half mile north of Cerro Prieto, just south of Mexicali,

California, at an elevation of 100 feet. While abundant in the

Salton Sea Basin, Colorado, it is believed that this is the first

record for Baja California. — Ralph Olen Fox, Department of

Invertebrate Zoology, California Academy of Sciences, San Fran-

cisco, California 94118.

WILLIAM H. WEEKS SHELL COLLECTION: New price lists

of this famous collection, with full scientific data, are in prepa-

ration. Many new additions of fine and rare species are also

included. To obtain free copies write:

George E. Jacobs, 853 Riverside Drive, New York 32, N. Y.

146 nautilus Vol. 83 (4)

NEWS

A Symposium on the Indian Ocean and Adjacent Seas will

be held at Cochin, India, January 12-18, 1971. It is sponsored

by the Marine Biological Association of India. Contributions of

papers and participation are being requested. Registration forms

and details may be obtained airmail from D. E. G. Silas, Marine

Biol. Assoc., Jyothi Buildings, Gopalaprabhu Cross Road, Cochin

11, India.

d'Alte Aldridge Welch — We regret to announce the death

of our longtime friend and colleague on January 4, 1970, in

Cleveland, Ohio. He was shot in a holdup attempt outside the

Racquet Club on December 10, 1969. Dr. Welch published 10

papers on land mollusks, mainly on the Hawaiian Achatinella,

in the Bulletin of the B. P. Bishop Museum, the Proceedings of

the Academy of Natural Sciences of Philadelphia, and The

Nautilus between 1929 and 1958.

He was born in New York City on April 9, 1907. His parents

were Charles James and Elizabeth (Livingston) Welch. He re-

ceived his Ph.D. from Johns Hopkins University in 1937. From

1942 to the time of his death he was a professor of biology at

John Carroll University in Cleveland. He was a Research Fellow

of the Department of Mollusks, Academy of Natural Sciences of

Philadelphia. As a hobby, he studied American children's books

printed prior to 1821, and published an extensive bibliography

in the Proceedings of the American Antiquarian Society from

1963 to 1968. Dr. Welch is survived by his wife, Ann Frances

(Goddard) , and five children, of 2298 Coventry Road, Cleveland

Heights, Ohio 44118 — editors.

WORLD WIDE SPECIMEN SHELLS for sale. New 1970

Price List on request. "Illustrated Catalog of Popular Mar-

ginella Species" now available also, 117 species shown with

full data and values listed with cover in color. Price $2.00.

PHILLIP W. CLOVER

Apartado de Correos 22

Rota Cadiz, Spain

April, 1970 nautilus \^\ (Index) iii

CONTENTS \- y

Names of new genera, species, etc. in italics

Acella haldemani 68

Achatina fulica invades Florida 75

"Achatina", new publication 146

Acroloxus coloradensis 105

Alaska, marines of Port Moller Bay 65

Alaska, nonmarine mollusks 44

American Malacological Union 67

Ancylid dispersal by beetle 144

Anodontoides ferussacianus 114

Artica islandica, nomenclature 61

Articidae versus Cyprinidae 61

Ashmunella, longevity 109

Aspella myrakeenae Emerson and D'Attilio 88

Bermuda Biological Station 66

Bryce, George W., Jr 105

Bursatella leachii pleii, biology 40

Caecum condylum D. R. Moore 26

Caecum insularum D. R. Moore 28

Cancellomorum 19

Concilia scrobiculata crosnieri Cernohorsky 95

Colorado 105

Conocardium of the Paleozoic 77

Corbicula in Baja California 145

Coryphella verrucosa rufibranchialis 37

Cyclinella tenuis 113

Cymatium boschi Abbott and Lewis 86

Cyprinidae 61

Dates of the Nautilus issue 35

Davis, George M 67

Donax fossor, variabilis 1

Doris verrucoa, biology 80

Galapagos Islands marines 19

Gallagher, J. L. and Harry W. Wells 22

Gilbertson, Lance H 29

Grau, Gilbert (1906-1969) 66

Haas, Fritz (1886-1969) 117

iv nautilus Vol. 83 (Index)

Histology of Quadrula nephridium 139

Indian Ocean Symposium 146

Laevapex fuscus, dispersal by insect 144

Lamarck's unionid types illustrated 52

Land mollusks, indoor raising 136

Latirus varai Bullock 133

Margaritifera falcata, hermaphroditism 113

Melanoides tuberculata in Florida 72

in Texas 67

Mercenaria, with Polydora infestation 74

Microvoluta joloensis Cernohorsky 103

Mitra pele Cernohorsky 99

Mitridae, new species 95

Monodenia, longevity 109

Moore, Donald R 26

Moore, George Mitchell (1906-1968) No. 1, iii

Morris, Percy A. (1899-1969) 116

Morum veleroae, from Galapagos 19

North Carolina 40

Nudibranch feeding 37

Odostomia cassandra Bartsch, synonym 71

Oklahoma, Polygyra 120

Oliva incrassata forma burchorum Zeigler 15

Oliva tremulina forma oldi Zeigler 16

Omalogyra atomus from Maine 70

Physa of Rawson 36

Physa striata complex 73

Plesiophysa pilsbryi 73

Polygyra dorfeuilliana, variability 120

Quadrula, histology of nephridium 139

Rangia cuneata, range extension 22

Sonorella, longevity 109

Sonorella odorata in Arizona 29

Succinea bakeri 42

Unionidae, effect of change pH 69

Unionidae, Lamarck's types 52

Unionids of Alabama, Georgia and Florida 34

van der Riet, J. C. L. (1920-1969) 66

Vexillum nodospiculum Cernohorsky 101

April, 1970

NAl 1 II. I ->

(Index) v

Volutomitridae, new species 95

Welch. d'Alte Aldridge (1907-1970) 146

vi nautilus Vol. 83 (Index)

INDEX TO AUTHORS

Abbott, R. Tucker 75

Abbott, R. Tucker and Hal Lewis 86

Boss, Kenneth J 61, 112

Branson, Branley A 35, 120

Bullock, Robert C 71, 76, 133

Carney, W. Patrick 136

Cernohorsky, Walter 0 95

Chanley, Paul 1

Clench, W. J 72, 73

Corgan, James X 65, 71

D'Attilio, Anthony (William K. Emerson) 88

Davis, John D 74

Emerson, William K 19

Emerson, William K. and Anthony D'Attilio 88

Fox, Ralph Olen 145

Franz, David R 80

Franzen, Dorothea S. (P. R. Myers) 139

Harman, Willard N 70, 115

Heard, William H 113

Hubricht, Leslie 42

Johnson, Richard 1 34, 52

Johnson, Robert L. (Samuel J. Tuthill) 44

Kruczynski, W. L. and Hugh J. Porter 40

Lewis, Hal (R. Tucker Abbott) 86

Michelson, Edward H 36

Morse, M. Patricia 37

Myers, Paul Robert and D. S. Franzen 139

Nicol, David 77

Porter, Hugh J. (W. L. Kruczynski) 40

Rosewater, Joseph No. 1, iii, 144

Tuthill, Samuel J. and Robert L. Johnson 44

Walton, Munroe L 109

Wells, Harry W. (John L. Gallagher) 22

Zeigler, R. F «. 14

i

Vol. 83 JULY, 1969 No. 1

THE ^

NAUTILUS &

THE PILSBRY QUARTERLY V£

DEVOTED TO THE INTERESTS OF CONCHOLOGISTS \&

EDITORS AND PUBLISHERS

Horace Burrington Baker, 11 Chelten Road, Havertown, Pa.

(Emeritus Professor of Zoology, University of Pennsylvania)

R. Tucker Abbott, duPont Chair of Malacology

Delaware Museum of Natural History, Greenville, Del. 19807

Charles B. Wi'rtz, Biology Department

La Salle College, Philadelphia, Pa. 19141

CONTENTS

Donax fossor: a summer range extension of Donax

variabilis. By Paul Chanley 1

Two infrasubspecific forms in Oliva. By Rowland F. Ziegler. . 14

Galapagan records for Morum veleroae (Gastropoda:

Tonnacea) . by William K. Emerson 19

Northern range extension and winter mortality of Rangia

cuneata. By John L. Gallagher and Harry W. Wells 22

A new Caecum from the tropical Western Atlantic.

By Donald R. Moore 20

Notes on the biology of the snail Sonorella odoral.a in

Arizona. By Lance H. Gilbertson 29

Further Additions to the unionid fauna of the Gull

drainage of Alabama, Georgia and Florida.

By Richard I. Johnson 34

Notes and News 35 Publications received iv

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Matter for publication should be sent to the acting editor, Dr. R. Tucker

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