/•^/<o^?X^'X NAUTILUS

" Ime i^, ^iRift)Jr^j- July i 972

CONTENTS N. 4'

A. Byron Leonard

A New Valvata from the Pleistocene of Southern Illinois 1

William G. Lyons

New Turridae (Gastropoda: Toxoglossa) from South Florida and the Eastern Gulf of Mexico 3

Frasier O. Bingham

Several Aspects of the Reproductive Biology oi Littorina irrorata (Gastropoda) 8

David Nice!

Geologic History of Deposit- feeding Pelecypods 11

Leslie Hubricht

Two New American Pulmonata: Paravitrea seradens and Philomycus sellatus 16

Dorothy E. Beetle

A Note on Land Snails Associated with Kudzu Vine 18

William F. Gale

Seasonal Variability in CalycuUsm in Sphaerium transversum (Say) 20

SHORT PAPERS

Patrick V. Kirch

Subfossil Non-marine Gastropods from Molokai, Hawaiian Islands 23

Jon-Ame Sneli

Odostomia turrita Found on Homanis gammarus 23

James X. Corgan

Pyramidellid Genera of Pilsbry 24

Morris K. Jacobson

Observations on the Siphonal Behavior of Young Surf Clams, Spisula solidissima 25

Book Reviews 10, 19

Dates of Publication of the Nautilus, vol. 85 26

1 'X' *«^ \

Vol. 86 ( 1 ) THE NAUTILUS

A NEW VALVATA FROM THE PLEISTOCENE OF SOUTHERN ILLINO

A. Byron Leonard

Division of Biological Sciences

The University of Kansas

Lawrence, Kansas 66044

ABSTRACT

Valvata salina Leonard ( Prosobranchia: Valvatidae), a new Pleistocene, Wisconsinan

Stage species, is described from the Saline River banks near Equality, Gallatin Co.,

Illinois. It differs from V, tricarinata (Say) in being more planoid, having a low spire and a

proportionately larger umbilicus.

The Saline River and its tributaries comprise a

drainage system of limited extent in southeastern

Illinois; the river arises no farther north than

Hamilton County, and enters the Ohio River in

northeastern Hardin County, an air-line distance of

little more than 60 miles. Especially in Saline and

Gallatin counties, the Saline River and its immediate

tributaries are entrenched in sediments of late

Pleistocene Lake Saline (Willman and Frye, 1970,

Fig, 9, p. 34); artificial deepening, straightening and

cleaning of the channel by heavy earthmoving

equipment has produced many clean exposures of the

ancient lake sediments up to heights of 35 or more

feet at low water stages of the river, although the

FIG. 1, 2, 3: umbilical, apertural and spiral views ofholotype o/ Valvata salina, n. sp. Corresponding views of an

associated example ofV. tricarinata are shown in figures 4, 5, and 6. All figures approximately x 6.

THE NAUTILUS

July 1972

.Vol. 86(1)

total thickness of the lake beds ranges upward to

approximately 1 50 feet. In the course of studying the

molluscan faunas in the fossihferous clays and silts

thus made available, an hitherto unknown member of

Valvata, a genus of branchiate gastropods, was

discovered. Inasmuch as the shells do not intergrade

in their distinguishing characters with those of

Valvata tricarinata (Say) with which they occur, the

shells are described as a new species.

Valvata sahna ru sp.

Figs. 1 , 2, 3

Diagnosis: Shells possessing the general

characteristics of the genus Valvata, and resembling in

many respects Valvata tricarinata (Say) from which it

differs, however, in the following important features:

consistently greater size; more planoid form;

relatively larger umbilical opening; and lack of spiral

sculpture.

Description of holotypes: Shell forming a helicoid

spiral, whorls 3^ in number; spire depressed below

general upper surface; nucleus of VA rounded whorls

bearing granular sculpture; remaining whorls increas-

ing rapidly and uniformly in diameter toward the

aperture, where the last whorl descends slightly;

whorls robust, tricarinate as in typical V. tricarinata,

except that the peripheral carina is reduced to

nothing more than a distinct angulation, and bearing

obvious but not conspicuous diagonally transverse

striations; spiral sculpture lacking on the whorls;

aperture round, entire, and cemented to the

penultimate whorls below the angle of the peripheral

carina; operculum not known; umbilicus relatively

broad, narrowing above, but exposing all the whorls

to the nucleus. Measurements: greater diameter of

shell, 6.9 mm; lesser diamer, 5.5 mm; height of shell,

2.8 mm; diameter of umbilicus, 2.1 mm; diameter of

aperture, 1.6 nun.

Type locality: Equality Northeast Section, in the

NW 1/4, NE 1/4, NW 1/4 sec. 16, T 9 S, R 8 E, Gallatin

County, Illinois. The fossiferous exposure is in the

right bank of the Sahne River, about a mile northeast

of the village of Equality.

Geological horizon: Equality Formation (WiUman

and Frye, 1970, p. 72), Woodfordian substage of the

Wisconsinan Stage of the Pleistocene Series.

Types: The holotype, and paratypes from the type

locahty and from two other localities exposed in

Lake Sahne sediments, are on deposit in the

paleontologjcal collections of the Illinois State

Geological Survey.

Comparisons: At each of the three locaUties from

which Valvata salina was recovered, it occurred with

V. tricarinata, as well as with about 20 other

molluscan species, several kinds of ostracods, seeds of

vascular plants and nucules of Chara and Nitella. The

greater diameter of the shells of V. salina varies

between 6.0 and 7,5 mm, while that of the associated

shells of V. tricarinata ranges from 4.6 mm to 5.0

mm. The planoid form of V. salina is exemplified by

the relation between greater diameter and the height

of the shells; the height comparises only 40 to 45 per

cent of the greater diameter, while in V. tricarinata

from the same deposits the corresponding range is

between 75 and 82 per cent. In shells of V. salina, the

diameter of the umbiMcus comprises from 31 to 34

per cent of the greater diameter of the shells, while in

V. tricarinata the diameter of the umbilicus

represents only 17 to 19 per cent of the greater

diameter. These data confirm the earlier statement

that Valvata salina is not only consistently larger than

examples of V. tricarinata which occur v^th it, but

also that it also differs from the latter in being much

more planoid, and in having a relatively broader

umbilicus.

LITERATURE CITED

Willman, H. B. and J. C. Frye 1970. Pleistocene

stratigraphy of Illinois. Illinois State Geol, Surv.

Bull. 94. 204 pp. 14 figs. 7 tables. 3 pis.

(in pocket).

Vol. 86 (1)

THE NAUTILUS

NEW TURRIDAE (GASTROPODA: TOXOGLOSSA) FROM SOUTH FLORIDA

AND THE EASTERN GULF OF MEXICO

William G. Lyons

Florida Department of Natural Resources

Marine Research Laboratory'

St. Petersburg, Florida 33731

ABSTRACT

Three new species, Cerodrillia gjrardi, Brachycythara barbarae, and Granoturris

presleyi are described primarily from collections taken in the eastern Gulf of Mexico, and

are compared with other related species. Daphnella margaretae is described from south

Florida. Daphnella retifera Dall, D, margaretae, and D. bartschi Dall, a closely related

species from the tropical eastern Pacific, are assigned to the subgenus Paradaphne

Laseron, previously known only from the Indo-Pacific.

Several undescribed turrids were found during

examination of mollusks collected in Project Hour-

glass, a systematic benthic sampling program con-

ducted in the eastern Gulf of Mexico by the Florida

Department of Natural Resources Marine Research

Laboratory (Lyons, 1968; Joyce & Williams, 1969).

New species of Cerodrillia, Brachycythara, and

Granoturris from these collections are described

herein. A previously unknown species of Daphnella

(Paradaphne) from south Florida is also described.

Collections of Recent Turridae of the Academy of

Natural Sciences of Philadelphia, Pennsylvania

(ANSP), the Museum of Comparative Zoology, Cam-

bridge, Massachusetts (MCZ), and the National Mu-

seum of Natural History, Smithsonian Institution,

Washington, D. C. (USNM) were examined during this

investigation. Additional specimens for study were

provided by Mrs. Margaret Kennedy, Delray Beach,

Florida, Dr. James H. McLean, Los Angeles County

Museum of Natural History (LACM), Los Angeles,

California, Mrs. Virginia 0. Maes, ANSP, and Mr. and

Mrs. Daniel Steger, Tampa, Florida.

Type depositories: Holotypes of all species de-

scribed are in the collection of the National Museum

of Natural History. Paratypes, deposited in various

other museums when sufficient material was avail-

able, are Usted in material of each species examined.

DMNH refers to the Delaware Museum of Natural

History.

GENUS Cerodrillia Bartsch & Rehder, 1939

Cerodrillia girardi new species

FIGS. 1, 2

Description: Shell with about 7]6. whorls, small, to

about 12.8 mm total length, solid, turreted, sub-

globose; color uniform waxy amber. Protoconch of 2

smooth, globose whorls passing without interruption

into post-nuclear whorls. Post-nuclear whorls SVi,

each with 7 or 8 strong, retractively curved, some-

what sigmoid ribs, broadest at rounded periphery.

Intercostal spaces broad, smooth except for micro-

scopic incremental Unes and extremely faint spiral

striation. Base short, marked with 7-9 spiral Unes

which increase in strength anteriorly. Aperture sub-

oval, broadest near middle. Columella nearly straight;

inner lip moderately wide, distinct. Outer lip thin,

backed by a thickened varix, irregularly curved, with

a shallow but distinct stromboid notch. Sinus deep,

broad, bordered posteriorly by a thick callus. Canal

very short, broad, shallow.

Material examined: Holotype: USNM 707001.

Length 8.8 mm, width 3.8 mm. Off Egmont Key,

Florida, Hourglass station D, 27°37'N, 83°58'W, 55

m; August 11, 1966. -Single adult paratypes, all from

station D, deposited at ANSP, LACM, MCZ, USNM,

American Museum of Natural History, New York

(AMNH), Delaware Museum of Natural History,

Greenville, Delaware (DMNH) and Florida Depart-

' Contribution no. 205

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July 1972

Vol. 86(1)

ment of Natural Resources Marine Research Lab-

oratory, St. Petersburg, Florida (FSBC I).

Remarks: Cerodrillia girardi sp. nov. is more

globose than C. clappi Bartsch and Rehder, C. perryae

Bartsch and Rehder, or C thea (Dall), though the

nucleus resembles that of C. thea. In general outHne it

is nearest C bealiana Schwengel & McGinty, with

which it is often collected, but the latter is a smaller

species with a much smaller and more pointed

nucleus.

Cerodrillia girardi was never collected in large

numbers during Project Hourglass, but low numbers

were taken frequently at station D. It was collected

only occasionally at stations E, L, and M (55 and 73

m stations). Most specimens other than type material

were juveniles or dead shells in poor condition. There

are specimens in the Steger collection from off the

lower Florida Keys in 49-55 m.

The species is named for Capt. Earl Girard, master

of the R/V Heman Cortez during the Hourglass

cruises.

GEH\}^BrachycytharaV^ooMrvg, 1928

Brachycythara barbarae new species

FIGS. 3, 4

Description: Shell with about 6 whorls, small, to

about 3.5 mm total length, biconic, translucent, with

httle or no spiral sculpture and strong axial ribs.

Protoconch with about 3 whorls, the tip distinctly

defined, not immersed in the next whorl; first two

nuclear whorls smooth, rapidly enlarging, somewhat

compressed axially; third whorl initially with fine,

curved, closely-spaced axial riblets which increase in

strength as they progress downward. Post-nuclear

whorls generally smooth, but faint spiral striae

sometimes present; axial ribs strong, sinuose, sharply

angled at the periphery, giving the whorls an angular

appearance; 8-9 such ribs on the body whorl. A faint

brown spiral band between suture and periphery of

spiral whorls; as many as six such bands below

periphery on body whorl, final band on the base most

broad. Eight or nine unornamented spiral threads on

base. Aperture narrow, about 40 per cent total length

of shell; outer lip thickened, rounded at shoulder;

sinus adjoining suture wide, rounded, very shallow;

parietal callus smooth, fairly broad; columella short,

truncate.

Material examined: Holotype: USNM 707003.

Length 3.4 mm, width 1.6 mm. Off Egmont Key,

Florida, Hourglass station D, 27°37'N, 83°58'W, 55

m; February 28, 1967. -Single adult paratypes, all

from station D, deposited at AMNH, ANSP, DMNH,

FSBC, LACM, MCZ, USNM, and the Steger

collection.

Remarks: Brachycythara biconica (C. B. Adams),

the only other Recent western Atlantic species near

B. barbarae sp. nov., has distinctly beaded spiral cords

on the post-nuclear whorls, giving the surface a

frosted appearance which immediately separates it

from B. barbarae. In addition, B. biconica possesses

more numerous axial ribs (10 -12 on body whorl)

which are rounded, not angled, at the periphery, and

a relatively longer aperture (about 50 per cent total

length of shell). The columella of B. biconica is

straight. The tip of the apex is immersed in the next

whorl, giving the nucleus a blunter appearance than

that oi B. barbarae. The lectotype of 5. biconica alba

(C. B. Adams) (Clench and Turner, 1950; pi. 32, fig.

1) has the frosted sculpture of B. biconica and is

apparently an unusual form of that species.

Bathymetric preferences for B. biconica and B.

barbarae are evident. In Hourglass collections, B.

biconica was common at 18 and 37 m stations but

seldom occurred at greater depths; it occurs in depths

as shallow as 1 m in the Florida Keys. Brachycythara

barbarae was common at 55 and 73 m Hourglass

stations, but was taken only occasionally at the

northern 37 m station (C), and was never taken at the

comparable southern station (K). There is one large

lot in the Steger collection taken southwest of

Sombrero Light, Monroe County, Florida, in 55 m. I

have not seen B. barbarae from shallower than 37 m.

The species is named for Mrs. Barbara Steger in

appreciation of her gracious hospitality during my

examination of the Steger collection.

GENUS Granoturris Fargo, 1953

Granoturris presleyi new species

FIGS. 5, 6

Description: Shell with about 7 whorls, small, to

about 4.7 mm total length, slender, turreted.

Protoconch flesh-colored, prominent, resembhng

Miraclathurella, of about IVi smooth whorls, tip

immersed in following whorl. First H post-nuclear

whorl strongly keeled at periphery as in Cryoturris,

with about 9 axial riblets. Subsequent whorls with

11-12 strong axial ribs, sharply angled at submedian

periphery. A spiral cord connecting peripheral angles,

another somewhat weaker cord midway between

periphery and anterior suture. Pre- and

post-peripheral sculpture of closely-spaced, frosted

Vol. 86(1)

THE NAUTILUS

FIGS. 1, 2 Cerodrillia girardi Lyons, Holotype, 8.8 mm, USNM 707001;

FIGS. 3, 4 Brachycythara barbarae Lyons, Holotype, 3.4 mm, USNM 707003;

FIGS. 5, 6 Granoturris presleyi Lyons, Holotype, 4. 7 mm, USNM 707005;

FIGS. 7, 9 Daphnella (Paradaphne) bartschi Da//, 10.4 mm, LACM collection;

FIG. 8 Daphnella (Paradaphne) margaretae Lyons, Holotype, 10.7mm, USNM 707006.

THE NAUTILUS

July 1972

Vol. 86(1)

spiral threads. Aperture elongate, subovate. Sinus

subsutural, a broad, shallow depression near shoulder.

Outer lip thin, unvariced, angled at shoulder. Anterior

canal short, moderately broad and deep. Post-nuclear

whorls white, with about 5 fine, brown spiral Unes on

post-peripheral portion, a more prominent, darker

brown Une immediately posterior to suture; 3 dark

brown Unes near middle of body whorl, 5 weaker

brown lines toward anterior canal.

Material examined: Holotype: USNM 707005,

Length 4.7 mm, width 1.8 mm. Off Egmont Key,

Florida, Hourglass station B, 27°37'N, 83°07'W, 18

m; November 20, 1967. -Three paratypes, all from

station B: 1 at ANSP, 2 at FSBC.

Remarks: Granoturris presleyi sp. nov. is the first

known Recent species from the western Atlantic; G.

padolina Fargo, from the North St. Petersburg

PUocene, differs from the Recent species by

possession of fewer axial ribs on whorls, and by its

larger size. The shoulders of G. padolina are normally

convex, according to Fargo; on G. presleyi, the

shoulders are always quite flat.

The species is named for Mr. Robert F. Presley,

ship's biologist of the R/V Neman Cortez during the

Hourglass cruises.

GENUS Dop/iwd/fl Hinds, 1844

SUBGENUS Paradaphne LusQron, 1954

Description: Shell small, to about 11.3 mm total

length, moderately thin, fusiform. Nucleus of 4-6

rounded, diagonally cancellate whorls, protractive

threads of cancellation somewhat stronger than

retractives on last whorl. Intersection of nuclear and

post-nuclear sculpture not sharply defined, pro-

tractive nuclear axials overlying for a small distance

post-nuclear spirals. Spire and body whorl with many

spiral lines crossed by axial threads, beaded at

intersections, with or without fine spiral threads in

interspaces. Aperture elongate-ovate, widest near

middle. Sutural sinus moderately deep, reversed

I^shaped, sloping anteriorly. Outer lip simple, curved,

tapered anteriorly, sUghtly thickened within; a very

weak notch at intersection with short, shallow,

moderately wide anterior canal.

Remarks: Powell (1966) dismissed Laseron's

Paradaphne for lack of characters of differentiation

from Daphnella, It is evident, however, that there

exists a circumtropical group of species, differing

markedly from Daphnella in nuclear and post-nuclear

sculpture. Daphnella botanica Hedley, 1918, the type

species o{ Paradaphne, belongs to this group, though

it is one of the species nearest in form to Daphnella

s. s.

Daphnella lymneiformis, the type species of

Daphnella, and closely related species have a smaller,

more slender protoconch of 2!^-3 whorls, with Uttle

variation in the strength of diagonal cancellation of

the last whorl. Intersection of nuclear and post-nu-

clear sculpture is abrupt, without the overlap zone of

Paradaphne. Size, sculpture, and number of whorls of

the protoconch of Paradaphne resemble those of

Metuonella Sorgenfrei, 1958, Cryptodaphne Powell,

1942, and Maoridaphne Powell, 1942, but these three

differ from Paradaphne by variously possessing strong

axial ribs, markedly different apertures and sinuses,

or both. In addition, the outer hp of adult Daphnella

s. s. is flared anteriorly, not tapered and constricted

as in Paradaphne. The aperture and outer lip of

Paradaphne closely resemble those of immature

Daphnella, but the inner thickening of some

specimens indicates that they are mature.

Eudaphne Bartsch, 1931 {non Reuss, 1922) was

proposed to contain Daphnella allemani (Bartsch,

1931). The name was later changed by Bartsch

(1933) to Eudaphnella. Keen (1958) suggested that

perhaps Eudaphne should be used in a subgeneric

sense for D. allemani and D. bartschi Dall, 1919.

However, D. allemani has the wide aperture, flared

outer lip rounded at the sutural sinus, and extremely

short canal of typical Daphnella and is so similar in all

other respects that subgeneric separation seerns

unwarranted, as noted by Powell (1966). Daphnella

bartschi is definitely assignable to Paradaphne, as are

D. retifera Dall, 1889, and a new species herein

described.

Daphnella (Paradaphne) margaretae new species

FIG. 8

Description: Shell with about 8 whorls, small, to

10.7 mm total length. Nucleus of about 5 rounded,

diagonally cancellate whorls, brown except fifth,

which has two spiral rows of large, round, white spots

best seen on wet specimens. Spire and body whorl

sculptured of fairly strong spiral lines crossed by

weaker axial threads, strongly beaded at intersections;

spiral threads within interspaces nearly obsolete.

Aperture slender, elongate; outer Up simple,

thickened within; sinus moderately deep, reversed L-

shaped. Siphonal canal broad, shaUow, brief, but

longer than that of most daphnelUds. Color cream,

with rich brown maculations. Three rows of large,

spot-Uke brown maculations on body whorl at

Vol. 86(1)

THE NAUTILUS

periphery, middle and posterior portion of base.

Material examined: Holotype: USNM 707006.

Length 10.7 mm, width 3.8 mm. South of Dry

Tortugas,Horida, 24°24'N, 82°58'W, 76-85 m; April

27, 1967; R/V Heman Cortez. --Para types: ANSP

324021. Length 5.4 mm, width 2.2 mm. On

Spondylus collected in 37-55 m off Palm Beach

County, Florida, 1970. -FSBC I 7896. Length 4.4

mm, width 2.0 mm. South of Sand Key, Monroe

County, Florida, 35-36 m, August 2, 1971; R/V

Heman Cortez.

Remarks: Radwin (1969: p.233) proposed the

term "cognates" to replace "analogues", as the

meaning of the former more clearly denotes "related

through the same origin" in the immediate biological

sense. Daphnella bartschi from the eastern Pacific and

D. margaretae sp. nov. are obvious cognates and have

probably evolved since emergence of the Central

American isthmus. D. margaretae differs from D.

bartschi (Figs. 7, 9) by possessing stronger spiral lines

crossed by weaker axial threads on the spire and body

whorl. The overall appearance of D. bartschi is of

even cancellation, whereas on D. margaretae spiral

sculpture is the dominant feature. Many fine spiral

threads in interspaces between post-nuclear spiral and

axial sculpture are clearly defined in D. bartschi, but

are nearly obsolete in D. margaretae. Overall, the shell

of D. margaretae is more slender than that of D.

bartschi The double row of large white spots on the

last nuclear whorl and maculations of spire and body

whorl are identical on both.

Daphnella retifera Dall, 1889, has one more

nuclear whorl and a more globose, thirmer shell than

either of the preceding species. Nuclear size, sculpture

of nuclear and post-nuclear whorls, and shape of the

aperture and outer lip indicate the relationship of this

species to the previous two. Interspaces between axial

and spiral sculpture are smooth. Color of the nucleus

is brown; post-nuclear whorls are white, with

scattered yellow maculations.

Daphnella (Paradaphne) margaretae is named for

Mrs. Margaret Kennedy, Delray Beach, Florida, who

provided one of the paratypes and whose articles on

western Atlantic Turridae in Seafari, the bulletin of

the Palm Beach County Shell Club, have stimulated

much interest in this group.

ACKNOWLEDGEMENTS

I thank Dr. James H. McLean and Mrs. Virginia

Maes for critically reading the manuscript and

providing additional information. They, Mrs. Mar-

garet Kennedy and Mr. and Mrs. Daniel Steger kindly

loaned specimens. The many persons who gave

assistance during inspection of collections at the

before-mentioned museums are gratefully acknow-

ledged. Special thanks are extended to Mrs. Sally

Kaicher who produced the photographs.

REFERENCES CITED

Bartsch, P. 1931. Descriptions of new marine

mollusks from Panama, with a figure of the

genotype of Engina, Proc. U. S. Nat. Mus., 79

(2881): 1-10, 1 pi.

Bartsch, P. 1933. Eudaphnella, a new generic name.

Nautilus, 47(2): 76-77.

Clench, W. J. and R. D. Turner. 1950. The western

Atlantic marine mollusks described by C. B.

Adams. Occ. Papers on MoUusks, 1(15): 233-403.

Dall, W. H. 1919. Descriptions of new species of

mollusks of the family Turritidae from the west

coast of America and adjacent regions. Proc. U. S.

Nat. Mus., 56(2288): 1-86, 24 pis.

Fargo, W. G. 1953. The Pliocene Turridae of Saint

Petersburg, Florida. In: Olsson, A, A. and A.

Harbison, 1953. Pliocene Mollusca of southern

Florida. Acad. Nat. Sci. Phila., Monograph 8, Pt.

2: 363-409, pis. 16-24.

Keen, A. M. 1958. Sea shells of tropical West

America; marine mollusks from Lower California

to Colombia, i-xi + 624 pp. Stanford Univ. Press,

Stanford, Calif.

Laseron, C. F. 1954. The New South Wales Turridae.

Roy. Zool. Soc. N. S. W. Handb., pp. 1-56.

Lyons, W. G. 1968. Mollusks of Project Hourglass.

Am. Malacol. Union Ann. Rpts. for 1968: 34-35.

Powell, A. W. B. 1966. The molluscan families

Speightiidae and Turridae. Bull. Auckland Inst.

and Mus., No. 5: 184 pp., 34 pis.

Radwin, G. E. 1969. A recent molluscan fauna from

the Caribbean coast of southeastern Panama.

Trans. San Diego Soc. Nat. Hist., 15(14): 229-236.

8

THE NAUTILUS

July 1972

Vol. 86(1)

SEVERAL ASPECTS OF THE REPRODUCTIVE BIOLOGY OF

LITTORINA IRRORATA (GASTROPODA)

Frasier O. Bingham

University of Miami

Rosenstiel School of Marine and Atmospheric Science ^

1 0 Rickenbacker Causeway

Miami, Florida 33149

ABSTRACT

The salt marsh periwinkle, Littorina irrorata, of eastern United States is dioecious. The

penis has a gray-brown, wrinkled basal region and a smooth, slender, white distal region

with a sperm groove located along the dorsal edge. Copulation, lasting 15 minutes or

longer, occurs in the warmer months and while the snails are out of water on marsh grass.

Females shed between 43,000 and 85,000 clear, planktonic, disc-shaped capsules, each of

which contains one gray egg. Spawning takes place at the air-water interface or just below

the surface of the water at a rate of four or five capsules per second for two to four

hours.

INTRODUCTION

The salt marsh periwinkle, Littorina irrorata (Say)

is common along the northern coast of the Gulf of

Mexico and the eastern coast of the United States

from central Florida to New York.

The observations reported here were made during a

detailed study of the behavior of the species in a salt

marsh near Panama City, Florida (30°14'N, 85°

4rW), during the period January through October,

1969 (Bingham, in press).

THE PENIS

The male copulatory organ was noted to develop

when male specimens reached a shell length of

approximately six mm. After this size is reached, the

separate sexes are easily distinguished by gently

lifting the outer shell lip of specimens that are

extended and firmly adhering to a substrate, and

noting the presence or absence of a penis slightly

below and posterior to the right tentacle (Fig. 1).

The penis has been seen to become larger and more

pigmented during the breeding season in some species

of Littorina (e.g. L. littorea by Tattersall, 1920; L.

augulifera by Linderking, 1954) and to show no such

seasonal variation in others (e.g. L. picta and L.

pintado by Struhsaker, 1966). No change in size or

color was noted in the penis of L. irrorata during the

period of this study which included the breeding

season.

* Contribution no. 1515.

The penis of L. irrorata can be divided externally

into two regions. The basal region (Fig. 2) is

gray-brown and has a wrinkled surface. The distal

region is white and has a fairly smooth surface. During

repose this region is slightly smaller in diameter and

approximately the same length as the basal region.

Located near the base of the basal region, so as to be

hidden from view when the organ is in its reposed

position (Fig. 1), is a light colored, slightly raised

area. Similar structures in L. littorea were found to be

glands and believed by Fretter and Graham (1962) to

secrete mucus used for holding the penis in position

during copulation. A sperm groove is located along

the dorsal edge of the entire length of the organ.

During copulation, the basal, region of the penis

becomes highly engorged with blood (Fig. 3) while

the distal region lengthens and decreases in diameter

so as to become thread-hke (Fig. 3).

COPULATION

Weekly observations of the snails in the salt marsh

were begun on January 4, 1 969. Copulation was first

noted during the afternoon of May 5 when about

25% of the animals were copulating, either at the

air-water interface or on grass shoots and leaves above

the water. The female of a copulating pair is always

firmly attached to the marsh grass with the foot

extended and, in most instances, in a spire-down

position. The male secures a firm hold on the female's

Vol. 86(1)

THE NAUTILUS

shell with its extended foot and comes to rest with

the right side of its shell hp slightly overlapping the

same portion of the female's shell. After the male is

in this position, the penis is directed into the mantle

cavity of the female and quickly inflates (Figure 4).

The length of time spent in copulation in 17 pairs

ranged from 24 to 36 minutes. During copulation and

for 15 minutes or longer thereafter, no locomotory

movements were seen in either sex.

High temperatures may be the stimulus for

copulation as suggested by the fact that during the

months of May, June and July, copulation was

observed on each field trip made in hot, clear weather

and no copulation was seen during two field trips

taken in cool, overcast weather conditions. In

studying this possibility, 150 specimens were

collected in the marsh on July 12 and brought to the

laboratory. Fifty specimens were placed in each of

three shallow pans which contained small amounts of

seawater and were covered with wire screens to

prevent the snails' escape. One pan was put outside in

the sunlight, one pan was left on a table in the

laboratory at 21°C, and one was placed in an oven set

at 40° C. No copulation occurred in the pan kept in

the laboratory. Copulation began in the sunlighted

pan and in the oven heated pan when a temperature

of 35°C was reached in each, and continued in both

pans for about two hours even though the pan in the

oven was taken out and allowed to cool to room

temperature (21*'C).

Tattersall (1920) noted that L. littorea kept in an

aquarium began to copulate soon after a water

change. He believed copulation to be induced either

by higher levels of dissolved oxygen or a decrease in

temperature. Alterations in temperature could also

have been involved when Lenderking(1954) observed

large numbers of L. augulifera copulating soon after

heavy rains.

Copulation in L. inorata was not seen in the marsh

after September 28, and specimens collected there-

after could not be induced to mate regardless of the

temperature at which they were maintained.

SPAWNING

Females which mated in the laboratory during the

afternoon of May 6 began spawning the next after-

noon at 1:10 PM, just 16 minutes after high tide

occurred in the area from which they were collected.

Specimens taken during the next several months

began spawning without regard to dayUght or dark-

ness at approximately the same time as the next high

tide after copulation.

A few females spawned while completely sub-

merged, but most were positioned at the air-water

interface (Fig. 5). Egg cases were continuously

expelled at a rate of four or five per second for a

period of two to four hours, and formed cloudy paths

to the bottom of the animals' containers. Two

females were isolated after copulation and their eggs

counted using a tally counter, grid slide and a known

volume of seawater in which the eggs had been

PENIS

FIG. 1. Repose position of the penis in L. irrorata.

FIG. 2. View of side of penis lying next to body. Penis in repose state. A - basal region; B - attachment gland; C ■

sperm groove; D - distal region.

FIG. 3. Penis engorged and the distal region in an extended state. A - basal region; B - attachment gland; C ■

sperm groove; D - distal regiori

10

THE NAUTILUS

July 1972

Vol. 86(1)

suspended by agitation. One snail laid approximately

43 thousand, and the other 85 thousand eggs.

THE EGG AND EGG CAPSULE

• L. irrorata lays clear, planktonic, disc-shaped egg

capsules which contain one gray egg each (Fig. 6).

The capsules measure from 250 to 280 n in diameter,

and are somewhat negatively buoyant in still seawater

of 26%o salinity.

LITERATURE CITED

Bingham, F. 0. In Press. The influence of en-

vironmental stimuli on the direction of movement

.i:t\>.<:-*.

•' :■'■•'**'•■

of the supraUttoral gastropod Littorina irrorata.

Bull. Mar. Sci.

Fretter, V. and A. Graham. 1962. British Prosobranch

Molluscs. Ray Society, London. 755 p.

Lenderking, R. E. 1954. Some recent observations on

the biology of Littorina angulifera Lam. of Bis-

cayne and Virginia Keys, Florida. Bull. Mar. Sci.

Gulf and Caribbean 3: 272-296.

Struhsaker, J. W. 1966. Breeding, spawning, spawning

periodicity and early development in the Hawaiian

Littorina: L. pintado (Wood), L. picta (Philippi)

and L. scabra (Linne). Proc. malac. Soc. Lond. 37:

137-166.

Tattersall, W. M. 1920. Notes on the breeding habits

and Ufe history of the periwinkle. Sci. Invest. Fish.

Br. Ire. 1: 1-11.

-250A

FIG. 4. Simplified drawing of copulating pair of L.

irrorata as seen through a glass substrate. A -male; B -

female.

FIG. 5. Simplified drawing of female L. irrorata

spawning

FIG. 6. Egg and egg capsule of L. irrorata. A - egg; B

egg capsule.

BOOK REVIEW

SEA SHELLS COMMON TO NORTH CAROLINA.

By Hugli J. Porter and Jim Tyler. 36 pp., 131 figs.,

colored cover on paperback. 1971. Free copy

obtained by writing: Division of Commercial and

Sports Fisheries, Dept. Natural Resources, More-

head City, N. C. 28557.

This is a well-illustrated and accurate guide to 1 29

marine mollusks of the coast of North CaroUna. It is a

paperback booklet designed for seashore visitors to

this part of the world. It contains both popular and

scientific names, brief descriptions, relative

occurrences, and habitats. The species are con-

veniently arranged in a series of simple keys and

illustrated by either drawings or clear photographs.

An index and glossary are included.

R. Tucker Abbott

Delaware Museum of Natural History

Vol. 86 (1)

THE NAUTILUS

11

GEOLOGIC HISTORY OF DEPOSIT-FEEDING PELECYPODS

David Nicol

Box 14376, University Station,

Gainesville, Florida 32601

ABSTRACT

All Paleozoic faunas average 10 per cent deposit-feeding pelecypods, with a range from

0 to 15 per cent. The Mesozoic pelecypod faunas average only 8 per cent deposit feeders,

with a range from 0 to 18 per cent. Of the Cenozoic faunas exluding the Recent, the

average percentage of deposit feeders is 16 per cent, with a range from 11 to 38 per cent.

There was a rapid increase in diversity of deposit-feeding pelecypods beginning in the

Paleocene. In the shallow-water Recent faunas the average percentage of deposit-feeding

pelecypods is 19 per cent, with a range of from 7 to 29 per cent, the Arctic having the

highest percentage. Deposit-feeding pelecypods are now at least as diverse as they have

ever been in the past. There is a higher percentage of deposit-feeding pelecypods (27 per

cent) in silty formations, than in sandy areas (12 per cent) or marl (10 per cent). There

appear to be at least 5 variables in the occurrence of deposit-feeding pelecypods - the

type of bottom sediment; geographic distribution; water temperature; bathymetric depth

and the evolutionary stage of development. The protobranchs originated in the

Ordovician, whereas the deposit-feeding tellinaceans are a relatively modem group,

appearing in the Early Cretaceous.

Protobranchs, which are deposit feeders, are a

minor element in most marine pelecypod faunas from

Ordovician to Recent. In a sample of 13 Paleozoic

faunas, protobranchs average 10 per cent of all

pelecypod species. This figure is quite consistent in

most of the Paleozoic faunas, although in 7 Permian

faunas the average is 11 per cent. The highest

percentage of all Paleozoic faunas, 1 5 per cent, is also

found in 2 Permian faunas.

The Triassic, Jurassic, and Early Cretaceous

(Albian and older) have lower percentages of proto-

branchs than is the average in the Paleozoic. The

average percentage for the Triassic and also the

Jurassic is 5 per cent and that for the Early

Cretaceous is only 4 per cent. The Late Cretaceous

faunas have an average of 7 per cent protobranchs,

but this average is high because of the Kemp Clay of

Texas, in which protobranchs comprise 16 per cent of

the pelecypod species (Stephenson, 1941). It is

interesting to note that the Kemp Clay is the

youngest Cretaceous formation in Texas and is Late

Maastrichtian in age. It is possible that the sudden

increase in protobranch species in a Late Cretaceous

fauna is an indication of colder seas at the end of the

Mesozoic.

The trend toward higher percentages of proto-

branchs continues in the Paleocene (3 faunas that

average 12 per cent protobranchs) and through the

Eocene (4 faunas average 13 per cent protobranchs).

Beginning with the Oligocene the percentage of

protobranchs decHnes considerably. Two OHgocene

pelecypod faunas average only 7 per cent proto-

branchs. Seven Miocene faunas average 9 per cent

protobranch species, and this includes a remarkable

fauna, the Miocene of Astoria, Oregon (Moore,

1963), in which 25 per cent of the pelecypod species

are protobranchs. The 2 Pliocene faunas average only

4 per cent protobranchs, and the 2 Pleistocene faunas

7 per cent protobranchs. The marked reduction of

protobranch species in shallow-water marine faunas

beginning in the Oligocene may be due to more

deposit- fee ding tellinaceans supplanting protobranchs

in this envirormient.

In the Recent marine faunas, protobranchs range

from 1 2 to 23 per cent only in the polar and most of

the cold-temperate faunas where the average tem-

12

THE NAUTILUS

July 1972

Vol. 86(1)

perature of the water is 10°C or less. The highest

percentage, 23 per cent protobranchs, is found in the

Arctic, but only 13 per cent of all Antarctic

pelecypod species are protobranchs. In the tropical

and warm-temperate marine faunas, the protobranchs

range from 1 to 7 per cent of each pelecypod fauna.

There are 3 fossil pelecypod faunas that are devoid

of protobranchs. The Permian Whitehorse fauna

(NeweU, 1940) has only 14 species of pelecypods,

none of which are protobranchs. The Early Triassic

fauna described by Ciriacks (1963) contains 23

species, but none are protobranchs. A Pleistocene

fauna from southwestern Japan described by

Hayasaka (1961) contains 87 species; none are

protobranchs, but there are 10 species of

deposit-feeding teUinaceans. The relatively coarse

sediment comprising the Whitehorse Sandstone was

not conducive to deposit feeding, and 14 pelecypod

species are suspension feeders. No explanation can be

given for the absence of protobranchs in the Early

Triassic fauna described by Ciriacks. The lack of

protobranchs in the Pleistocene fauna from south-

western Japan may be attributed to the fact that it

was a shallow warm-water environment suitable for

deposit-feeding tellinaceans but unsuitable for proto-

branchs.

No Jurassic or older pelecypod faunas analyzed

had undoubted deposit-feeding tellinaceans, (See

Pohlo, 1968, for deposit-feeding and

non-deposit-feeding teUinaceans) although this super-

family has been reported in rocks as old as Late

Triassic or Early Jurassic. Throughout the Cretaceous,

deposit-feeding teUinaceans (the TeUinidae) comprise

no more than 6 per cent of the total pelecypod

species in any of the 1 1 faunas analyzed, but none of

these faunas was devoid of the TeUinidae. Only the

Wangaloan (Paleocene) beds of New Zealand lack

deposit-feeding teUinaceans (Finlay and Marwick,

1937). The remainder of the Paleocene, the Eocene,

the Oligocene, and a few of the early Miocene faunas

have deposit-feeding teUinacean species comprising 5

to 9 per cent of the pelecypod species. Most Middle

Miocene faunas and aU Late Miocene faunas have 10

per cent or more deposit- feeding teUinaceans. This is

true also of the 2 PUocene and 2 Pleistocene faunas

analyzed. The oldest Miocene fauna that has at least

10 per cent deposit- feeding teUinaceans is the Chipola

Formation of Florida, and it is considered to be Late

Early Miocene in age. By Middle Miocene time the

diversity of deposit- feeding teUinaceans had reached

approximately its present level.

In Recent faunas there is a higher percentage of

deposit-feeding teUinaceans in warm water than in

cold water. In water temperature of 20°C or higher,

deposit-feeding teUinaceans range from 9 to 19 per

cent of the total pelecypod fauna. In colder waters

the deposit- feeding teUinaceans range from 0 to 10

per cent. There is an interesting geographic trend

superimposed on that of water temperature. In the

southern hemisphere the number of species of de-

posit-feeding teUinaceans is considerably lower than

in the northern hemisphere. The South AustraUan

and New Zealand faunas have only 4 per cent

deposit-feeding teUinacean species, the Tasmanian

and MageUanic faunas have only 3 per cent, and the

Antartic pelecypod fauna has none. On the other

hand, the fauna at Monterey Bay, California, has 10

per cent deposit- feeding teUinacean species, that of

Canada 9 per cent, and that of the Arctic 6 per cent.

Beginning with the Cretaceous, one can compare

the diversity of the protobranchs to the de-

posit-feeding teUinaceans in the various fossU

pelecypod faunas. The Early Cretaceous pelecypod

faunas have few species of deposit-feeding teUinids,

but there is an increase in number of species in

Cenomanian faunas, which are commonly considered

the oldest Late Cretaceous. The oldest pelecypod

fauna where teUinid species outnumber protobranch

species is in the Aptian, Early Cretaceous of Lebanon

(Yokes, 1940). However, there are only 5 de-

posit-feeding species in this fauna, comprising 9 per

cent of the total pelecypod species. Of the 9 Late

Cretaceous pelecypod faunas examined, 3 have more

teUinid species than protobranch species. The 3

Paleocene and 4 Eocene faunas have a greater number

of protobranch species than teUinacean de-

posit-feeding species, despite the fact that the

semeUds, which are also deposit- feeding telUnaceans,

first appeared in the Eocene. Of the 2 Oligocene

faunas, one has a greater number of deposit-feeding

teUinacean species than protobranch species. How-

ever, beginning with the Miocene the teUinacean

deposit-feeding species more commonly outnumber

the protobranch species. Of 7 Miocene pelecypod

faunas analyzed, only 2 have more species of proto-

branchs than of deposit-feeding teUinaceans. These 2

are the Shoal River Miocene of Florida (Gardner,

1926-1928) and the Miocene of Astoria, Oregon

Vol. 86(1)

THE NAUTILUS

13

(Moore, 1963). In all 4 Pliocene and Pleistocene

faunas, the deposit-feeding tellinacean species out-

number the protobranch species. Thus, from Miocene

onward, most of the fossU pelecypod faunas are

similar to Recent warm-water faunas (average tem-

perature 20°C or higher, Nicol, 1969) in that the

deposit-feeding tellinaceans outnumber the proto-

branch species. It is obvious from these data that the

ratio of percentages of teUinacean deposit-feeding

species to protobranch species in pelecypod faunas

older than Miocene is an unreUable indicator of water

temperature, and in even some of the Early and

Middle Miocene pelecypod faunas, water temperature

may not be too accurately indicated by the proto-

branch-tellinacean species ratio.

In summation on the diversity of deposit-feeding

pelecypods through time, all Paleozoic faunas average

10 per cent deposit feeders, with a range from 0 to 15

per cent. The Mesozoic pelecypod faunas average

only 8 per cent deposit feeders, with a range from 0

to 1 8 per cent. Of the Cenozoic faunas excluding the

Recent, the average percentage of deposit feeders is

16 per cent, with a range from 11 to 38 per cent. The

fauna with 38 per cent deposit-feeding pelecypods is

from the Middle Miocene of Astoria, Oregon. It is

interesting to note the rapid increase in diversity of

deposit-feeding pelecypods beginning in the

Paleocene. In the shallow-water Recent faunas the

average percentage of deposit-feeding pelecypods is

1 9 per cent, with a range of from 7 to 29 per cent,

the Arctic having the highest percentage but closely

followed by Canada. Deposit-feeding pelecypods are

now at least as diverse as they have ever been in the

past.

Purdy (1964) pointed out that there is a higher

percentage of deposit- feeding marine animals in silt

and clay substrates than in the coarse sediments such

as sands, where deposit feeders are few or absent. In

the fossil faunas analyzed, where there are reliable

data on the Uthology of the beds, the following

results were obtained. In 6 sandy formations the

average percentage of deposit-feeding pelecypods is

12 per cent. In 5 marl and Umestone formations the

average percentage of deposit- feeding pelecypods is

only 10 per cent. This is surprising in that the

percentage is lower than in the sandy formations. In 3

silty formations the average percentage of de-

posit-feeding pelecypods is 27 per cent

There appear to be at least 5 variables in the

distribution of, or diversity of, deposit-feeding

pelecypods. The type of bottom sediment is defin-

itely a factor. There is a greater percentage of

deposit- fee ding pelecypods in clays and sUts than in

sands and limestones. Geographic distribution can be

a factor in the diversity of deposit-feeding

pelecypods at a given locality. At present there are

considerably fewer deposit-feeding tellinaceans in the

southern hemisphere than in the northern hem-

isphere, and this mainly explains why the southern

hemisphere pelecypod faunas have a lower percentage

of deposit feeders. The paucity of deposit-feeding

tellinaceans in the Recent faunas of the southern

hemisphere may explain why there are none of these

pelecypods in the Wangaloan (Paleocene) of New

Zealand. In other words, the deposit-feeding

tellinaceans originated in the northern hemisphere

and are still not so diverse in the southern hem-

isphere. Water temperature is a factor in de-

posit-feeding pelecypod distribution today, with

protobranchs dominating the colder waters. Water

temperature is also closely linked with water depth.

In deeper water there is generally a higher percentage

of deposit feeders, and at great depths (2000 meters

or more) the protobranchs comprise at least 35 per

cent of the total pelecypod species. The stage of

diversity or evolution of the protobranchs and de-

posit-feeding tellinaceans is another important var-

iable. The protobranchs are an ancient stock, going all

the way back to the Ordovician, whereas the de-

posit-feeding teUinaceans are a relatively modern

group, appearing in the Early Cretaceous but not

attaining their present great diversity until about the

Middle Miocene.

In order to conserve space, I have not cited in the

main body of this paper each reference that I

analyzed, but all references are included in the

bibhography. Fourteen Recent and 50 fossil

pelecypod faunas are analyzed. Of the fossil faunas,

19 are Cenozoic, 18 are Mesozoic, and 13 are

Paleozoic in age. I have excluded the Recent deep-sea

pelecypod fauna because all of the fossil faunas

analyzed are from shallow water. Deep-water sed-

iments of the past are rarely available for study, and

modern deep-water faunas are quite different from

fossil shallow-water faunas.

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CoraUian Lamellibranchiata. Palaeontographical

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Bird, S. 0. 1968. A pelecypod fauna from the

14

THE NAUTILUS

July 1972

Vol. 86 (1)

Gaptank Formation (Pennsylvanian) West Texas.

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CarceUes, A. R., and S. I. Williamson 1951. Catalogo

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Ciriacks, K. W. 1963. Permian and Eotriassic bivalves

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Finlay, H. J. and J. Marwick 1937. The Wangaloan

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Gardner, J. A. 1935. The Midway Group of Texas.

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Gilbert, H. and J. de H. de Braucourt 1954.

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THE NAUTILUS

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Ferdinand Rau, Ohringen, Germany. 1-461.

Smith, A. G. and M. Gordon, Jr. 1948. The Marine

mollusks and brachiopods of Monterey Bay, Cal-

ifornia, and vicinity. Proc. Calif. Acad. Sci., 4th.

Ser. 26 (8): 147-245.

Stenzel, H. B., E. K. Krause and J. T. Twining 1957.

Pelecypoda from the type locality of the Stone

City Beds (Middle Eocene) of Texas. Univ. Texas

pub. 5704: 1-237.

Stephenson, L. W. 1923. The Cretaceous formations

of North Carohna. Part 1. Invertebrate fossils of

the Upper Cretaceous formations. North Carolina

Geol. Survey. 5 : 1-604.

Stephenson, L. W. 1941. The larger invertebrate

fossils of the Navarro Group of Texas. Univ. Texas

pub. 4101: 1-641.

Stephenson, L. W. 1953. Larger invertebrate fossils of

the Woodbine Formation (Cenomanian) of Texas.

U. S. Geol. Survey Prof Paper 242: 1-226.

Yokes, H. E. 1939. Molluscan faunas of the

Domengine and Arroyo Hondo Formations of the

California Eocene. Ann. New York Acad. Sci. 38:

1-246.

Yokes, H. E. 1 946. Contributions to the paleontology

of the Lebanon Mountains, RepubUc of Lebanon,

part 3. The pelecypod fauna of the "Olive Lo-

cality" (Aptian) at Abeih. BuU. Am. Mus. Nat.

Hist. 87(3): 143-215.

Yokes, H. E. 1957. Miocene fossils of Maryland.

Maryland Geol. Survey Bull. 20: 1-85.

Wade, B. 1926. The fauna of the Ripley Formation

on Coon Creek, Tennessee. U. S. Geol. Survey

Prof Paper 137: 1-272.

Warmke, G. L. and R. T. Abbott 1961, Caribbean

seashells, Livingston PubUshing Co., Narberth,

Pennsylvania. 1-348.

Wilson, A. E. 1956. Pelecypoda of the Ottawa

Formation of the Ottawa-St. Lawrence lowland.

Geol. Survey Canada BuU. 28: 1-102.

Winters, S. S. 1963. Supai Formation (Permian) of

eastern Arizona. Geol. Soc. Am. Mem. 89: 1-99.

Woodring, W. P. 1925. Miocene mollusks from

Bowden, Jamaica. Pelecypods and scaphopods.

Carnegie Inst. Washington pub. 366: 1-222.

16

THE NAUTILUS

July 1972

Vol. 86(1)

TWO NEW NORTH AMERICAN PULMONATA: PARA VITREA SERADENS

AND PHILOMYCUS SELLATUS

Leslie Hubricht

4026 35th Street

Meridian, Mississippi 39301

ABSTRACT

Paravitrea seradens Hubricht (Zonitidae) from Gauley Bridge, Fayette Co., West

Virginia, and the slug Philomycus sellatus Hubricht from Princeton, Jackson Co.,

Alabama, are described as new species. Additional geographical records are given for

Helicodiscus triodus Hubricht. Anatomical notes are made on Triodopsis occidentalis

(Pilsbry and Ferriss) and Polygyriscus virginianus (P. R. BurchJ.

Triodopsis occidentalis (Pilsbry & Ferriss)

Triodopsis obstricta occidentalis (Pilsbry and Ferriss),

Pilsbry, H. A., 1940, Land Mollusca of North

America (north of Mexico). Aca. Nat. Sci. Phil-

adelphia, Mono. 3, 1 : 829.

Mesodon sargentianus (Johnson & Pilsbry),

Vagvolgyi, J., 1968, BuU. Mus. Comp. Zool. 136:

210.

1 recently collected this species alive near Locust

Grove, Independence Co., Arkansas. The genitaha was

found to be similar to that of T. denotata (Ferussac)

(Pilsbry, fig. 473-6.). The shell appears to be a

connecting Unk between T. obstricta (Say) and T.

fosteri (F. C. Baker). The aperture is somewhat

intermediate between these two species. The

periphery varies from strongly angulate to carinate.

Paravitrea seradens new species

FIGS. la-c.

Description: Shell small, pale amber, subhyaline,

glossy; spire low, dome-shaped, sutures shallow,

sculpture of numerous irregularly spaced radial

grooves; periphery somewhat flattened giving the

shell a quadrate appearance in immature shells,

becoming more rounded at maturity; umbilicus deep

and well-like exhibiting all the whorls, contained over

5 times in the diameter of the shell, base flattened

and excavated around the umbihcus; whoris slowly

expanding, last whorl expanding more rapidly and

deflected downward to the aperture; aperture lu-

nate, obhque; lip thin, simple; internal armature of

one or more pairs of rather large teeth on the outer

and basal walls at all stages of growth, except in an

occasional very old shell.

Height 2.8 mm., diameter 5.5 mm., aperture

height 2.0 mm,, diameter 2.5 mm., umbilicus di-

ameter 1.0 mm,, 6.9 whorls. Holotype.

Distribution: West K/rgzw/a; Kanawha Co.: wooded

hillside, Hernshaw. Fayette Co.: below Cane Branch

Falls, 1.5 miles east of Gauley Bridge, holotype

173020 and paratypes 173021 Field Museum of

Natural History, other paratypes A8774 collection of

the author. Wyoming Co.: mountainside, 2.8 miles

northeast of Wolf Pen. Logan Co.: ravine, 4.8 miles

southwest of Holden. Mingo Co.: Gyandot River

bluff, opposite Justice; mountainside, 0.7 mile north

of Cinderella. McDowell Co.: ravine, 4 miles north-

east of Welch.

Remarks - Paravitrea seradens is most closely

related to P. capsella (Gould) differing in having at

least one pair of teeth at all stages of growth; in

having a more expanded aperture; in having the base

more impressed around the umbilicus; and in having a

larger umbilicus in immature shells, the umbilicus

being more well-like. The type locality for this species

is also the type locaHty for Helicodiscus triodus

Hubricht and Carychium clappi Hubricht.

Polygyriscus virginianus (P. R. Burch)

Polygyra virginianus P. R. Burch, 1 947, The Nautilus

61:40.

Polygyriscus virginianus (P. R. Burch), Burch J. B.,

1962, How to Know the Eastern Land Snails. Wm.

C, Brown Co., Dubuque, Iowa. p. 148.

Vol. 86(1)

THE NAUTILUS

17

On a recent visit to the type locality I succeeded

in collecting a series of living specimens of this

species by "mining" at the right spot. Immature

shells have epidermal fringes on the spiral ridges, and

look very much like immature Helicodiscus

fimbriatus Wetherby, but are without teeth. After

maturity these fringes wear off and they are com-

pletely gone in old shells. In adults there is a

transverse tooth on the base a short distance within

the aperture. This was not mentioned in the original

description. P. virginianus seems closely related to

Helicodiscus but whether it should be placed in that

genus is left open pending anatomical studies.

Helicodiscus triodus Hubricht

This species can be added to the growing list of

cave snails, as it has been found living in the two

caves listed below.

West Virginia: Monroe Co.: McClungs Cave,

Zenith. Greenbrier Co.: McClungs Cave, 1.5 miles

northeast of Maxwelton. These two caves are not

connected, although they have the same name. They

are about thirty miles apart.

Philomycus sellatus new species

FIGS. Id-e

Description: The mantle pattern is black on a

white or cream background in very young slugs; as

they become older the pattern becomes lighter, dark

gray in half-grown slugs and brownish gray in adults

and the background becomes darker so that the

pattern is somewhat obscured. The forward end of

the mantle is irregularly mottled; behind this is an

area in which the dark pigment is greatly reduced.

About one-third of the way back is a transverse dark

band with irregular margins. On the latter half of the

mantle there is a broad dorsal band which is bordered

by a series of elongated black spots. On each side

there is a narrow longitudinal band m\h irregularly

scattered mottling above and below. This mottUng

may become so thick as to obscure the lateral bands

and merge them with the dorsal band. The tentacles

are dark gray, and the foot is white. The maximum

length of adults when extended in crawling is about

100 mm.

Distribution: Alabama: Jackson Co.: wooded hill-

side, 1.7 miles northeast of Princeton, holotype

157322 F. M. N. H., paratypes 30066 collection of

the author; base of Putnam Mtn., 4 miles northwest

of Princeton; Woodville (H. E. Sargent); Russell Cave

FIG. lA-C. Paravitrea seradens Hubricht, holotype.

FIG. ID-E. Philomycus sellatus Hubricht, holotype.

National Monument. Madison Co.: wooded hillside,

1.3 miles northeast of New Hope: base of Sharp

Mtn., near Sneeds Spring, Sharps Cove, northeast of

Maysville. Tennessee: Franklin Co.: hilltop, 2.5 miles

south-southeast of Huntland. Marion Co.: Raven Den

Point Sink, 1.8 miles west of Martin Spring.

Remarks: Philomycus sellatus is most closely re-

lated to P. carolinianus (Bosc). The latter half of the

mantle is not distinguishable, but the transverse Ught

band followed by a dark band on the forward half

will readily separate it. Philomycus sellatus is the

common upland slug of northeastern Alabama and

adjacent Tennessee. It sometimes occurs with P.

carolinianus.

Illustrations by Miss Carole Wrigley provided

through the courtesy of Field Museum of Natural

History.

18

THE NAUTILUS

July 1972

Vol. 86(1)

A NOTE ON LAND SNAILS ASSOCIATED WITH KUDZU VINE

Dorothy E. Beetle

Peninsula Nature Museum

524 Clyde Morris Boulevard

Newport News, Virginia 23601

ABSTRACT

A heavy growth of Kudzu vine taking over a sandy slope and nearby woodland in

Newport News, Virginia, now supports a large population of land snails, consisting of

ten species. Among the genera represented are Triodopsis, Retinella, Hawaiia,

Ventridens, Zonitoides, Helicodiscus, Strobilops, Gastrocopta, and Vertigo.

On March 28, 1971 a fire raced along the north

boundary of the grounds of the Peninsula Nature

Museum in Newport News, Va. This area includes the

steep slope of a railroad overpass and adjacent forest

floor. This report lists the snails found exposed by

the complete removal of leaf litter.

Originally the land where the museum and the road

are situated was covered with a mixed pine and

hardwood forest. White and black oaks and hickories

are abundant and sweetgum, red maple, beech, tulip

tree and the shrubby dogwood and sourwood occur

in the area. The elevation is 45 feet above sea level.

The soil is coarse sand, with crossbedded sands,

gravels and clays extending downwards to

considerable depths. Newport News lies on a

peninsula which juts into the Chesapeake Bay. Its

climate is mild with abundant rain.

When the road and overpass were constructed in

1958, the steep sandy slopes were planted v/ith

Kudzu vine to prevent erosion. This rapidly formed a

dense mass which smothered volunteer plants. The

Kudzu has since invaded the adjacent woods. It has

smothered shrubs under festoons of vine and climbed

into the crowns of the tallest trees, killing them.

Under the trees scattered patches of moss, grass and a

few early spring flowers are the only plants which

survive.

In fall the Kudzu becomes dormant, leaving a

brittle mat of dead leaves resting slightly above the

soil on the vines which thickly crisscross along the

ground. These leaf mats can be lifted intact to view

the bare sand beneath. Even in the woods the sand

permits the rapid leaching of material and only a thin

humus layer is formed. Under the Kudzu vine

dessicated plant material consists almost entirely of

the previous year's growth. On the steep slopes of the

overpass the soil has almost no traces of plant debris.

In view of the exposed and Umited habitat of the

overpass, it was surprising to discover a large

population of land snails consisting of 10 species, plus

3 additional ones on the floor of the woods invaded

by Kudzu. The fire completely cleared the area of

plant material and the snails were picked up from the

surface of the sand.

STATION 1

One square yard, 35° slope, almost under the

overpass, 3 feet above ground level. The Triodopsis

and Gastrocopta were most numerous at this station.

One specimen of V. oscariana occurred here.

Triodopsis juxtidens

Retinella indentata paucilirata

Hawaiia minuscula

Ventridens cemoides

Zonitoides arboreus

Helicodiscus parallelus

Strobilops labyrinthica

Gastrocopta contracta

Vertigo oscariana

Vol. 86(1)

THE NAUTILUS

19

STATION 2

One square yard on low slope of embankment, one

foot above ground level.

Triodopsis juxtidens

Retinella indentata paucilirata

Helicodiscus parallelus

Succinea sp., juvenile

Gastrocopta contracta

STATION 3

Under standing dead trees, under and around log 4

inches in diameter.

Triodopsis juxtidens

Triodopsis fallax obsoleta

Triodopsis albolabris

Helicodiscus parallelus

Retinella indentata paucilirata

Hawaiia minuscula

Random collection from the forest floor also

turned up specimens of Mesodon thyroidus and

Ventridens cemoides. Triodopsis juxtidens was most

abundant all through the forest area; in some places

as many as a dozen individuals lying in one square

yard.

The woods in the vicinity of the nature museum

are second growth and receive fairly heavy use for

recreational purposes. While much of the understory

has been trampled down, the Kudzu vine has not yet

invaded this section. The same mollusks are present as

in the area of the Kudzu vine except for Succinea sp.,

Gastrocopta contracta and Vertigo oscariana. T.

juxtidens and//, parallelus are most abundant.

Gratitude is expressed for the assistance of F.

Wayne Grimm for identification of some of the

material.

LITERATURE CITED

Pilsbry, Henry A. 1940-1948 Land MoUusca of North

America (North of Mexico). Vols. 1-3. Acad. Nat.

Sci. Phila., Monogr. 3.

BOOK REVIEW

THE MARINE MOLLUSCS OF ARCTIC CANADA.

By Elizabeth Macpherson. PubUcations in Biolog-

ical Oceanography, no. 3. 149 pp., 54 maps, 7 pis.

Paperback. 1971. $3.50. Available by mail from

the Marketing Division, National Museums of

Canada, Ottawa 4, Canada.

This is a welcome handbook account of the shelled

marine mollusks of Arctic Canada exclusive of the

bivalves and cephalopods. Included are 1 08 species of

prosobranch gastropods, chitons and scaphopods.

Each species has its main synonymy, adequate de-

scription, Ust of Canadian localities, literature

records, overall range, distributional map, and an

accompanying photograph. There is a brief but

interesting discussion of the ecology and zoo-

geography of the mollusks of the Canadian Arctic. An

up-to-date bibhography and index are included.

The nomenclature is excellent, although a few

discrepancies have crept in. Lophyrochiton

Yakovleva, 1952, may be preceded by Stenosemus

Middendorff, 1847, and Lepidopleuroides Thiele,

1893. Buccinum tehue Gray, 1839 (non Schroter,

1805) should be scalariforme Moller, 1842. The

modern typography of the printed page, consisting of

alternating double and single columns and

not-too-clear photographs do not make this otherwise

excellent work easy to use.

R. Tucker Abbott

Delaware Museum of Natural History

20

THE NAUTILUS

July 1972

Vol. 86(1)

SEASONAL VARIABILITY IN CALYCULISM IN SPHAERIUM TRANSVERSUM (SAY)^

Waiiam F. Gale^

Department of Zoology and Entomology

Iowa State University, Ames, Iowa 50010

ABSTRACT

The nepionic shell of the freshwater bivalve, Sphaerium transversum, may be separated

from the older part of the shell by a sulcus, thus forming a bulging "cap " or caly cuius at

the umbo. Populations sampled from a stream in Iowa showed varying percentages of

specimens being calyculate or noncalyculate. Growth experiments in a field laboratory

indicated that arrested growth caused the formation of a calyculus, thus suggesting that

this character is of questionable taxonomic importance.

The calyculus or "cap" is the bulging upper

portion of the beak or umbo of some sphaeriid

shells. It consists of the shell of the nepionic young

(Thomas, 1965) and is separated from the rest of the

shell by a sulcus. While investigating the bottom

fauna of the Mississippi River near Fort Madison,

Iowa (Gale, 1969), I observed that some Sphaerium

transversum were distinctly calyculate and that others

were distinctly noncalyculate. Frequently, inter-

mediates were found in which the nepionic shell was

distinct, but not elevated above the surface of the

adjoining shell. Such shells were not considered

calyculate.

That S. transversum occurs in two forms has long

been recognized (Sterki, 1909). The significance of

the calyculus and why it is sometimes absent have not

been explained. Herrington (1968, personal commu-

nication) suggested that "capped beaks" might result

from a rest period. The presence of calyculi was a

characteristic used to justify the validity of

Musculium as a genus. Herrington (1962) suggested

that calycuh have little taxonomic value because they

occasionally occur in groups other than Musculium

and are variable within a species.

* Joumal Paper No. J-7035 of the Iowa Agriculture

and Home Economics Experiment Station, Ames,

Iowa. Project 1374. A contribution from the Iowa

Cooperative Fishery Unit sponsored by the Iowa

State Conservation Commission, Iowa State Univer-

sity of Science and Technology, and the Bureau of

Sport Fisheries and Wildlife (U. S. Department of

Interior). (Contract 14-16-0003-12204).

^ Present address: Ichthyological Associates,

Berwick, Pennsylvania 18603.

In bottom samples, the proportion of calyculate

clams diminished progressively between June and

September (Table 1). Similar trends occurred at

several sampling stations. Gams 2-3 mm long, with

the exception of those in May and December, usually

were not calyculate. Probably most of the clams 2-3

mm long had not grown since birth. Since the

calyculus represents the nepionic shell, it cannot be

observed until visible post-embryonic growth has

taken place. A few calyculate clams 2-3 mm long

were observed in the June-November collections, but

were missed in random selection of the subsamples.

In June, all clams examined were calyculate except

those 2-3 mm long. Most clams over 3.0 mm long in

June probably had over-wintered when 2-3 mm long.

In May only clams of the first two sizes were present

at station 14, the station where the June to December

samples were collected. Unless immigration occurred

at station 14 between May and June or unless the

samples were not representative, the small clams of

May grew into large calyculate clams by June. Thus,

clams that overwinter when 2-3 mm long, presumably

before growing, become calyculate as growth com-

mences the following spring.

In July, all clams 3-4 mm long and 30% of those

4-5 mm long were noncalyculate. These were prob-

ably born in June or July 1967. By August, non-

calyculate clams 6-7 mm long were beginning to

appear. A major die-off of calyculate clams in large

size classes occurred in August, and by September,

only noncalyculate clams remained. The absence of

calyculate clams at station 14 in September means

that the entire population consisted of clams born in

1967, probably since May.

Vol. 86(1) THE NAUTILUS 21

TABLE 1. Number of S. transversum examined (No.) in 1967 and percentage (%) with calyculate

shells. Specimens for April and May were pooled from 7 open-water stations about three

miles downstream from Ft. Madison, Iowa (Stations 4-10 on transect 3 of Gale, 1969).

Specimens for June through December were collected at a station about 0.5 miles further

downstream (transect 4 Station 14 of Gale, 1969). Clams 2-4 mm long were present in

April but were discarded after counting. When more than 10 clams were available for one

size group the specimens were randomly selected.

22

THE NAUTILUS

July 1972

Vol. 86(1)

In October, calyculate clams reappeared, with

smaller clams becoming calyculate. During November

and December, the population included both

calyculate and noncalyculate clams. Of clams 2-3 mm

long, 90% were calyculate in December. Thus, over-

wintering is not essential for calyculus formation; but

all small clams that over-wintered became calyculate,

while only part of those bom between September and

December became calyculate.

The possibility that an initial period of no growth

(resting state) influences subsequent shell devel-

opment is plausible and is supported by growth

experiments conducted in a field laboratory during

1967 (Gale, 1969). In the laboratory most clams

which he d an initial lag in growth became calyculate

and those which grew immediately after birth became

noncalyculate. Therefore, the presence or absence of

calycuU in S. transversum is not hereditarily deter-

mined within the species. Thomas (1959) observed

that newborn S. partumeium reared in the laboratory

frequently exhibited a lag in growth lasting up to 6

weeks, but clams that grew immediately gave birth to

young that also began to grow immediately. Pre-

sumably, clams that did not grow immediately

produced young that had a growth lag.

The stimulus that eUcited a resting state in

laboratory-reared clams in my study, is not known,

but certain clues may be discovered by examining the

methods in which the clams were treated. Clams, in

chambers where growth lags occurred, were born in a

pan of river water, picked up with a camel-hair brush,

measured with an ocular micrometer, and placed into

a vial of river water until four were collected. The vial

of clams and water was then poured into the proper

chamber. Small clams in chambers where growth lags

did not occur were born there and were not measured

or otherwise handled until the chambers were cleaned

after 33 days.

Physical and chemical changes may have occurred

in the pan of river water holding the parental stock.

None of the newborn was in the pan longer than an

hour, but the process of gathering them from the pan

and making measurements consumed almost an entire

afternoon; ample time for changes in water temper-

ature, dissolved oxygen, pH, etc.

Tactile stimulation might have elicited a resting

state. Besides being handled with a brush, young were

frequently pushed about by larger clams in the pan.

That some young were born in the well-lighted pan

and others in subdued Hght in the laboratory may

have been important. That some chambers contained

the decomposing remains of the parental stock may

also have been a factor. Factors causing a resting state

in the laboratory and in the field may not have been

the same.

The presence of calyculi in part of the population

was helpful in assessing growth rates of the clams in

their natural environment. For example, if we assume

that the largest noncalyculate clams (4-5 mm long) in

July (Table 1) became the largest noncalyculate clams

(6-7 mm) in August, the mean length increase was

about 2 mm. Also, noncalyculate clams 3-4 mm long

in July grew to 5-6 mm in August. Since clams were

noncalyculate in September, growth of calyculate

clams can be followed in fall.

The results of this study substantiate Herrington's

(1962) contention that calyculism is of questionable

taxonomic importance.

I wish to thank Dr. Kenneth D. Carlander and Dr.

R. Jess Muncy, Department of Zoology and En-

tomology, Iowa State University, who directed the

research and made helpful suggestions concerning the

manuscript,

LITERATURE CITED

Gale, W. F. 1969. Bottom fauna of Pool 19,

Mississippi River with emphasis on the Ufe history

of the fmgemail clam, Sphaerium transversum.

Ph.D. thesis, Iowa State Univ., Ames, Univ. Micro-

film No. 69-20642, Ann Arbor, Mich.

Herrington, H. B. 1962. A revision of the sphaeriidae

of North America (Mollusca: Pelecypoda). Univ.

Mich. Mus. Zool. Misc. Publ. 118. 74 p.

Sterki, V. 1909. Some observations and notes on

Musculium, The Natilus 23: 17-19.

Thomas, G. J. 1959. Self fertilization and production

of young in a Sphaeriid clam. The Nautilus 72:

131-140.

Thomas, G. J. 1965. Growth in one species of

sphaeriid clam. The Nautilus 79: 47-54.

Vol. 86(1)

THE NAUTILUS

23

SHORT PAPERS

SUBFOSSIL NON-MARINE GASTROPODS FROM

MOLOKAI, HAWAIIAN ISLANDS

Patrick Vinton Kirch

Bernice P. Bishop Museum

P. 0. Box 6037, Honolulu, Hawaii 96818

Rapid extinction of endemic land snails in the

Hawaiian Islands has occurred due to massive removal

of native forests, particularly in lowland areas

(Carlquist 1970). Hence it is extremely difficult to

find associations of endemic gastropods in their

original habitats. Perkins (1913, pp. xxvi-xxvii)

long-ago recognized the importance of fossiliferous,

pulmonate-bearing deposits in the reconstruction of

lowland snail associations. In 1969 and 1970 I

studied a Holocene, non-marine gastropod bearing,

subfossil deposit at the base of Halawa Valley,

Molokai Island. The deposit (50-Mo-Al-4) is an

eroded taluvial fan (Wentworth 1943) comprising

unconsolidated sand, alluvial outwash, and slumped

material.

Charcoal from the base of the fan was submitted

to Gakushuin University, Japan, for radiocarbon age

determination. The result (GaK-2744) was 750 ± 90

B. P. (1950). The geologic matrix indicates that the

fossils were deposited with debris eroded from the

adjacent subvalley following extensive forest clearing

in prehistoric times.

Table 1 lists the gastropods (pulmonates and

prosobranchs) from this deposit. Apparently

originally they comprised a lowland snail association,

and as such allow a partial reconstruction of the

lowland flora as well. The probable habitat of these

snails is Ukely to have been dominated by Acacia koa,

Chenopodium sp., Eugenia sandwicensis, and small

stands of Metrosiderous sp. (an important pulmonate

host). Today the region is dominated by Leucaena

glauca, an introduced species.

Acknowledgements. I should Hke to thank Dr.

Yosiho Kondo for aid in the identification of

specimens.

Table 1 . Gastropods from A 1-4

Halawa Valley, Molokai Island

ENDODONTIDAE

Endodonta sp.

ACHATINELLIDAE

Tomatellinops baldwini ( Ancey)

Lamellidea gracilis (Pease)

Elasmias sp.

Tomatellides procerula (Ancey)

Tomatellaria cincta (Ancey)

AMASTRIDAE

Leptachatina emerita Sykes

Leptachatina varia Cooke

Amastra (Amastrella) petricola (Newcomb)

Amastra (s.s.) humilis (Newcomb)

Amastra (Heteramastra) perversa Hyatt and Pilsbry

SUCCINEIDAE

Succinea sp.

HELICINIDAE

Pleuropoma laciniosa var. molokaiensis Neal

LITERATURE CITED

Carlquist, S. 1970. Hawaii: A Natural History. The

Natural History Press, New York.

Perkins, R. C. 1913. Fauna Hawaiiensis 1(6), Cam-

bridge University Press.

Wentworth, C. K. 1943. Soil avalanches on Oahu,

Hawaii. Geol. Soc. Amer. Bull. 54: 53-64.

ODOSTOMIA TURRITA FOUND ON

HOMARUS GAMMARUS^

Jon-Ame Sneli

Biological Station, N-7001 Trondheim, Norway

Several species of molluscs belonging to the family

Pyramidellidae have been reported as ectoparasites of

other marine invertebrates (Robertson & Orr 1961,

Cheng 1967), but only one observation of a

pyramidellid in association with a crustacean has been

reported (Fretter & Graham 1949).

In October 1969 a lobster (Homarus gammarus

(L.) = H. vulgaris H. Milne Edwards) of 5.1 kg weight

was brou^t to the Zoological Museum in Oslo. The

^ Contribution No.

Trondheim, Norway.

1 56, Biological Station,

24

THE NAUTILUS

July 1972

Vol. 86(1)

lobster was collected at Herad near Farsund on the

Norwegian coast. On the pinching legs and carapace

were observed a few Pomatoceros triqueter (L.) and

by close examination of the gills one specimen of

Odostomia turrita Hanley was found. The

identification of the Odostomia species was kindly

done by Mr. Tore Hoisaeter. The O. turrita had to sit

weU-attached to the gills as the lobster was boiled,

transported and handled by fishmongers before

delivered to the museum.

O. turrita is distributed from the north of Norway

south to the Mediterranean, but no information

about its host-specifity has been reported. According

to Robertson & Orr (1961) the discovery of a

pyramidellid "on" or "with" another invertebrate

does not mean necessarily that the invertebrate in

question is host to the pyramidellid. The only reliable

records of parasitic relationships are those in which a

pyramidellid has been seen to feed.

Observation of feeding was in this case impossible,

but the record is still interesting as no earlier

observation of Odostomia species associated with

decapod crustaceans has been reported. Fretter &

Graham (1949) have seen O. plicata (Montagu) thrust

its proboscis onto the antennae of amphipods at

Plymouth but this behaviour may well have been

accidental (Robertson & Orr 1961).

Although the case remains to be proved there is no

reason to believe that the O. turrita found on the

lobster gill could possibly be associated with the few

P. triqueter located on the exoskelton of the lobster.

LITERATURE CITED

Cheng, T. C. 1967. Marine molluscs as hosts for

symbioses with a review of known parasites of

commercially important species. Adv. Mar. Biol. 5:

1-424.

Fretter, V. & A. Graham. 1949. The structure and

mode of hfe of the Pyramidellidae, parasitic

Opisthobranchs. J. Mar. Biol. Ass. U. K. 28:

493-532.

Robertson, R. & V. Orr. 1961. Review of

pyramidellid hosts, with notes on an Odostomia

parasitic on a Chiton. Nautilus 74: 85-91.

PYRAMIDELLID GENERA OF PILSBRY

James X. Corgan

Austin Peay State University

Clarksville, Tennessee 37040

H, A. Pilsbry proposed four genus-group names for

taxa he considered referable to the gastropod family

Pyramidellidae Gray, 1 840. Each name was proposed

for a new subgenus and each subgenus was charac-

terized in a few brief words or sentences appended to

a specific description. There was Uttle in PUsbry's

writings to indicate that the subgenera were new. As a

result, standard nomenclators Hke the Nomenclator

Zoologicus overlooked some of the names. Most were

also omitted from an excellent catalogue of Pilsbry's

taxa by Clench and Turner (1962).

Today, the genus-group names Pilsbry introduced

in the PyramideUidae are httle used because the

pyramidellid faunas he studied have received little

subsequent attention. His taxa have been overlooked

and have not been evaluated by modern workers.

Pilsbry's names are available names. The groups he

discriminated seem to be meaningful taxa. Several

could be widely distributed in the world fauna. This

report lists Pilsbry's pyramidellid genera and provides

a brief appraisal of each.

Evaletta PUsbry, 1918

Type Species - By original designation, rwrfton/7/fl

(Evaletta) elizabethae Pilsbry, 1918, p. 315.

Though it was described as a subgenus of Tur-

bonilla Risso, 1826, Evaletta and Turbonilla have

little in common. As interpreted by Dall and Bartsch

(1909) and by Laws (1937), Turbonilla has prom-

inent axial sculpture and an emergent, heterostrophic

protoconch. In Evaletta the heterostrophic proto-

conch is deeply immersed and there is no axial

sculpture. Species of Evaletta have more whorls than

most Turbonilla. The growth form is more acicular

and there is generally a weakly developed tooth on

the inner lip. Evaletta differs from Turbonilla in so

many characters that it should be ranked as a distinct

genus.

Only two species have been referred to Evaletta:

Turbonilla (Evaletta) elizabethae Pilsbry (1918, p.

314, fig. 9) and T (E.) laysaensis PUsbry (1918, p.

315, fig. 10). Both are Recent species from the

Hawaiian Islands.

Vol. 86(1)

THE NAUTILUS

25

Goniodostomia Pilsbry and Johnson, 1917

Type Species - By original designation, Odostomia

(Goniodostomia) superans Pilsbry and Johnson,

1917, p. 181.

Corgan (1971) catalogued the content of this

nominal genus and interprets Goniodostomia Pilsbry

and Johnson, 1917, as a junior synonym of

Eulimastoma Bartsch, 1916.

Nesiodostomia Pilsbry, 1918

Type Species - By original designation, Odostomia

(Nesiodostomia) prima Pilsbry, 1918, p. 325; non

Odontostomia acutiuscula var. prima Boettger, 1907

{= Nesiodostomia montforti Corgan, 1972).

The characters, content, and distribution of

Nesiodostomia are discussed at length by Corgan

(1972). Nesiodostomia has been recorded from the

Tertiary of Japan but this reflects confusion with the

morphologically similar genus Puposymola Coss-

mann, 1921. All species that are correctly referred to

Nesiodostomia are part of Recent faunas of the

Hawaiian Islands.

Odostomidea Pilsbry And Johnson, 1917

Type Species - By monotypy, Odostomia (Odos-

tomidea) bartschiana Pilsbry and Johnson, 1917, p.

182.

Odostomidea Pilsbry and Johnson was proposed as

a subgenus of Odostomia Fleming, 1813, but there

are few similarities between the two taxa. Species of

Odostomia lack sculpture while species of

Odostomidea are highly sculptured. In Odostomia,

the nuclear whorls are generally deeply immersed in

the first teleoconch whorl while they are about half

immersed in Odostomidea. The tabulate shoulder of

Odostomidea contrasts markedly with the smoothly

convex whorl profile of Odostomia. The two taxa

seem generically distinct.

Two species have been referred to Odostomidea

Pilsbry and Johnson: Odostomia (Odostomidea)

bartschiana Pilsbry and Johnson (1917, p. 152,

Ulustrated by Pilsbry, 1922, pi. 36, fig. 15) and O.

(O.) mogindo PUsbry (1922, p. 393, pi. 36, fig. 6).

Both were described from the Oligocene of Hispanola

and both bear a strong resemblance to Recent species

that have been referred to Salassia Folin, 1872,

Eupyrgulina Melvill, 1910, and Odostomella

Bocquoy, Dautzenberg, and Dollfus, 1883. A survey

of the hterature suggests that all of these nominal

genera need both taxonomic and nomenclatural

review. Until a comprehensive study is completed, it

does not seem possible to evaluate Odostomidea

Pilsbry and Johnson, 1917.

REFERENCES CITED

Clench, W. J., and Turner, R. D. 1962, New names

introduced by H. A. Pilsbry in the Mollusca and

Crustacea: Acad. Nat. Sci. Philadelphia, Spec. Pap.

4. 218 pp.

Corgan, J. X. 1971, Review oi Parodostomia, Telloda,

Goniodostomia, and Eulimastoma (Gastropoda:

PyramideUacea): The Nautilus 85, p. 51-60.

Corgan, J. X. 1972, Pacific species oi Nesiodostomia

Pilsbry, 1918, and Puposymola Cossmann, 1921

(Gastropoda: PyramideUacea): The VeUgerH p.,

355-360.

Dall, W. H., and Bartsch, P., 1909, A monograph of

the West American pyramidellid moUusks: U. S.

Nat. Mus. Bull. 68. 358 pp. 30 pis.

Laws, C. R., 1937, Review of the Tertiary and Recent

Neozelanic pyramidellid molluscs. No. 1. The

genus Turbonilla: Trans. Royal Soc. New Zealand

66, p. 402- 422, pis. 32-33.

Pilsbry, H. A., 1918, Marine Mollusca of Hawaii,

IV- VII: Proc. Acad. Nat. Sci. Philadelphia 69 p.

309-333, pis. 20-22.

PUsbry, H. A., 1922, Revision of W.M. Gabb's

Tertiary MoUusca of Santo Domingo: Proc. Acad.

Nat. Sci. PhUadelphia 73 p. 305-435, pis. 16-47.

PUsbry, H. A. and Johnson, C. W., 1917, New

MoUusca from the Santo Domingo OUgocene:Proc.

Acad. Nat. Sci. PhUadelphia 69 p. 1 50-205.

OBSERVATIONS ON THE SIPHONAL

BEHAVIOR OF YOUNG SURF CLAMS,

SPISULA SOLIDISSIMA

Morris K. Jacobson

American Museum of Natural History

Central Park West at 79th Street, New York 10024

Not many observations on the siphonal behavior of

the surf clam, Spisula solidissima (DUlwyn), in nature

appear in the literature. The foUowing account of

such an observation made in Rockaway Beach, New

York may therefore prove to be of interest.

During the September 1971 ebb rides, the sandy

area inhabited by large numbers of immature surf

clams was uncovered for a few hours each day. The

clams measured from about 40 to 80 mm (1^ to

26

THE NAUTILUS

July 1972

Vol. 86(1)

about 3 inches) in length. Dense patches of clams

were scattered throughout the sandy substrate. The

small openings of the siphons at the surface of the

sand were the only part of the clam that was visible.

Morse (1919:188) described these siphon openings.

He found 28 small papillae on the Up of the anal

(dorsal or excurrent) siphon, but they were not

readily visible under field conditions, and the opening

appeared as a small, rather narrow oval with

apparently smooth edges. The opening of the

branchial (ventral or incurrent) siphon was twice the

width of the anal siphon and was much distorted by

"two excessively stout papillae springing from the

wall in the conjunction with the anal syphon. They

are curved outward half-way across the opening. One

of these is bifurcated." (Morse, I.e.). The 33

irregularly alternating long and short papUlae fringed

the outer margin and were clearly seen. Frequently

both siphons were visible, but when only one

appeared, it was almost always the opening of the

dorsal siphon.

During the short inter-wave periods, when the sand

was not being roiled by the wave action, the siphon

openings were exposed. As a wave flowed in carrying

a thick suspension of sand grains, the siphon openings

closed and the presence of the animal was not

detectable for a moment. In the short interval when

the upsweeping wave came to a brief halt, the sand in

suspension sank rapidly to the bottom and

simultaneously the siphon openings appeared. As the

water receded, sand grains were again placed in

suspension and the openings again disappeared. As

the sand swiftly settled, the siphons opened again and

remained open until the next onrush of water.

Apparently the film of water remaining on the sand

surface before the next flow of water was sufficient

to enable the mollusk to continue its feeding and

elimination activities.

At more or less regular intervals a small jet of

water, thickly laden with sand grains, was ejected

from the ventral siphon. This behavior was typical of

clams inhabiting the active wash zone. As the tide

came in, observations had to come to an end, but it

can be assumed that in areas covered by the tide and

not within the wash zone, the surf clams kept both

siphons open constantly during its feeding periods.

The type of siphonal activity described for Spisula

is apparently a defensive adaptation to minimize the

ingestion of sand particles. Any sand taken in is

probably stored briefly in the branchial siphon and

expelled at certain intervals. It was surprising to note

that the anal siphon was open more often than the

branchial or feeding siphon.

This manuscript was kindly reviewed by Dr.

William K. Emerson of the American Museum and Dr.

John Ropes of the National Fisheries Service, Oxford,

Maryland.

LITERATURE CITED

Morse, E. S. 1919. Observations on Uving

lamellibranchs of New England. Proc. Boston Soc.

Nat. Hist. 35 (5): 139-196.

Dates of Publication of The Nautilus

Volume 85, number 1: July 20, 1971

Volume 85, number 2: October 4, 1971

Volume 85, number 3: January 27, 1972

Volume 85, number 4: April 28, 1972

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NOVEMBER, 1972

THE

NAUTILUS

Vol. 86

Nos. 2 - 4

A quarterly

devoted to

malacology and

the interests of

conchologists

%» ^ '

0EC4 Wt

Woods Hoitt M|l»»

C-4<'

Founded 1889 by Henry A. Pilsbr/, Continued by H. Burrington Baker.

Editors: R. Tucker Abbott and Charles B. Wurtz

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EDITORIAL COMMITTEE

CONSULTING EDITORS

Dr. Arthur H. Clarke, Jr.

Department of Mollusks

National Museum of Canada

Ottawa, Ontario, Canada K1A-0M8

Dr. William J. Clench

Curator Emeritus

Museum of Comparative Zoology

Cambridge, Mass. 02138

Dr. William K. Emerson

Department of Living Invertebrates

The American Museum of Natural History

New York, New York 10024

Mr. Morris K. Jacobson

Department of Living Invertebrates

The American Museum of Natural History

New York, New York 10024

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Department of Geology

The Ohio State University

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Los Angeles County Museum of Natural History

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Biological Laboratory

National Marine Fisheries Service

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Division of Marine Geology

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Division of Mollusks

U. S. National Museum

Washington, D.C. 20560

Dr. G. Alan Solem

Department of Invertebrates

Field Museum of Natural History

Chicago, Illinois 60605

Dr. David H. Stansbery

Museum of Zoology

The Ohio State University

Columbus, Ohio 43210

Dr. Ruth D. Turner

Department of Mollusks

Museum of Comparative Zoology

Cambridge, Mass. 02138

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Division of Biology

School of Marine and Atmospheric Science

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EDITORS

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Delaware Museum of Natural History

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i«nne Bmlogicai LuboratoryT"^

L/e/?>J|JPvNAUTl[.US

DEC 4 1972

CONTENTS

Hal Lewis

Notes on the Genus Distorsio (Cymatiidae) with Descriptions of New Species 27

Tadashige Habe

Notes on the Genus Parancistrolepis Azuma (Buccinidae) 51

Gale G. Sphon

Berthella kaiiiae, a New Opisthobranch from the Eastern Pacific 53

E. P. Cheatuni

A New Species of Humboldtiana (Helminthoglyptidae) from the Sierra Vieja Mountains of Texas 56

Winston F. Ponder and Gilbert P. Whitley

Tom Iredale (1880-1972) [an obituary] 60

Ronald F. Thomas and Frazier 0. Bingham

Shell Structure in Spirula spirula (Cephalopoda) 63

Joseph Rosewater and Geerat J. Vermeij

The Amphi- Atlantic Distribution oi Littorina meleagris 67

Joseph Rosewater

Teratological Littorina scabra angiilifera 70

Samuel L. H. Fuller

Type Locality of Unio pumilus Lea, 1 838 (Unionidae) 72

Samuel L. H. Fuller

Unio caroliniana Bosc, 1 801 (Unionidae) 74

Marc J. Imlay

Greater Adaptability of Freshwater Mussels to Natural Rather than to Artificial Displacement 76

(continued over)

i

Barry Roth and Richard E. Petit

A New Species o^ Persicula from the Arabian Sea (MarginelUdae) 80

Horace G. Richards

Quaternary MoUusks from Fiji 81

Hank K. Mienis

On the Alleged Occurrence of Lambis violacea and Clithon coronata in Indonesia 82

Tadashige Habe and Kiyoshi Ito

Ancistrolepis kawamurai, a New Japanese Buccinidae 83

Samuel L. H. Fuller

An Underscribed Structural Feature in the Marsupium oi Elliptic) lanceolata (Lea, 1828)(Unionidae) ... .85

Derek S. Davis

An Unusual Form of Littorina littorea (Linne) Found in Nova Scotia 87

News

Book Reviews

(of) R. T. Abbott. 50; B. R. Wilson and K. Gillett. 59; M. Furlong and V. Pill. 66; L. G. Hertlein and U. S.

Grant, 4th. 71.

NOTICE TO SUBSCRIBERS

In order to have a new volume of The Nautilus begin in January, rather than July, we have issued the

remainder of volume 86 in a combined issue of nos. 2, 3, and 4. For further information on billing, see p. 82.

11

Vol. 86 (2 - 4)

THE NAUTILUS

27

NOTES ON THE GENUS DISTORSIO (CYMATIIDAE) WITH

DESCRIPTIONS OF NEW SPECIES

Hal Lewis

Research Associate, Dept. Malacology

Academy of Natural Sciences of Philadelphia

Philadelphia, Pa. 19103

ABSTRACT

Distorsio perdistorta Fulton, 1938, whose known distribution has previously been

limited to Japanese waters, has been discovered in the Gulf of Mexico, the Lesser Antilles,

and the Indian Ocean. Distorsio horrida Kuroda and Habe, 1964, also originally limited to

Japanese waters, is a synonym A different species from Japan, misidentified by previous

workers as "perdistorta," is described as a new subspecies, Distorsio constricta habei

Lewis. The American tertiary progenitors of this species are believed to be D. crassidens

(Conrad, 1848) of the Oligocene and D. simillimus (Sowerby, 1850) of the Miocene. The

living species D. constricta is recognized as having three geographical subspecies - con-

stricta (Broderip, 1833) from the tropical Eastern Pacific; macgjntyi Emerson and Puffer,

1953, from the Western Atlantic; and habei, new subspecies from Japan. Similar subspe-

cific distributions are known among the Cymatiidae, Cassidae and Volutidae.

An unusual periostracal structure is described and illustrated along with various ana-

tomical features o/ Distorsio.

Distorsio burgessi is described as a new species from Hawaii which is sympatric with

Distorsio anus (Linne).

The lectotype of Distorsio ridens Reeve is figured and D. ridens is clearly separated

from D. reticulata /?oc?mg, D. perdistorta Fw/fo«, and D. decussatus Valenciennes.

INTRODUCTION

While examining shells in the collection of Mr. and

Mrs. Riley Black of Fort Myers, Florida, three spec-

imens of a Distorsio were brought to my attention.

These matched a specimen first sent to me by Mrs.

Elsie Malone of Sanibel, Florida, which had been mis-

identified as Distorsio macgintyi Emerson and Puffer,

1953. Surprisingly, these specimens proved to be in-

distinguishable from Distorsio perdistorta Fulton,

1938, a species living in Japanese waters which had

been renamed Distorsio horrida by Kuroda and Habe

in 1964. Unfortunately they also misidentified an-

other taxon known as "Distorsio perdistorta" Kuroda

and Habe, 1964, (not Fulton, 1938), which now re-

quires a new name.

This paper will confirm Distorsio perdistorta

Fulton as a vaUd species; substantiate its occurrence

in the Western Atlantic and the Indian Ocean; clarify

its history in the literature; differentiate it from Dis-

torsio clathrata (Lamarck, 1816) 2in^ Distorsio mac-

gintyi Emerson and Puffer, 1953, in the Western At-

lantic; and describe the taxon previously known as

"Distorsio perdistorta" Kuroda and Habe, 1964.

HISTORICAL DISCUSSION OF

DISTORSIO PERDISTORTA

Emerson and Puffer, 1953, in their catalogue of

the Distorsio suggested the possibility that perdistorta

is a geographical subspecies of Distorsio reticulata

Roding, 1798, and said, "Fulton states that this spe-

cies possesses affinities withZ). ridens {Rqqvq, 1844)"

which they considered to be a "variant of D. re/^/cw-

/a/a 'Bol ten' Roding, 1798."

An examination of reticulata and perdistorta has

shown that there are consistent taxonomic differ-

ences. Table 3 was prepared to facilitate a comparison

using characters which are helpful in separating Dis-

torsio. The list of differences cited in Table 3 and the

28

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

specific description of perdistorta contained in this

paper demonstrates that these are separate species.

Actually, Fulton (1938, p. 56) hsted various char-

acters which "separates this from ridem Reeve."

They are "the shorter anterior canal and the much

closer granular or reticulate spiral sculpture . . .; the

line of numerous small plaits on the anterior part of

the columella is perpendicular not oblique as in

ridens. " Examinarion of the lectotype, syntypes and

Reeve's figure shows that ridens is clearly separable

from perdistorta. D. ridens is regulariy fusiform,

while perdistorta is grossly distorted. There are 25 to

30 axial ribs on the body whori of perdistorta and

only 1 2 on ridens, giving it a much more open sculp-

ture.

The spiral cords of perdistorta are separated by a

fine interstitial cord which is lacking in ridens. The

parietal shield of ridens is medium-oval in shape with

TABLE 1. Specimens o/Distortio perdistorta examined from the Gulf of Mexico.

Vol. 86(2-4)

THE NAUTILUS

29

Length

(mm)

Width

(mm)

Locality

Depth

(meters)

Madagascar:

60 34

43.5

20.5

P2. Chalutage 10

12°43'S-48°15'E

300-340

very coarse plicae along the columellar edge which

curve posteriorly away from the aperture past the

axis of the shell. The shield oi perdistorta is more ob-

long with finer teeth on the columellar edge which

are more or less in a straight line which ends at the

axis of the shell. (See Table 3 for a comparison be-

tween perdistorta, ridens, and reticulata).

Until now, Western Atlantic specimens of per-

distorta have been so rarely collected that they have

failed to come to the attention of those who have

worked with this genus. In 1951, Axel Olsson and

Thomas L. McGinty described Distorsio comtricta

floridana as a subspecies of the Panamic Distorsio

constricta Broderip 1833. Unfortunately, the name

had previously been used by Gardner (1947, p. 535,

pi. 53, fig. 8) for Personella floridana and it was nec-

essary for Emerson and Puffer (1953, p. 101) to re-

name it Distorsio mcgintyi. The spelling is here

emended to macgintyi to conform with the recom-

mendations on the formafion of names in Appendix

D of the 1961 Code. Among the material with which

Olsson and McGinty dealt were the Distorsio collect-

ed by the Blake expedition in the Gulf of Mexico,

1877-78 and in the Caribbean, 1879-80. While dis-

cussing the Distorsio in his well-known "Blake Re-

port", Dall (1889, p. 221) referred to Link's (1807,

p. 1 23) hst which limited the genus to two living spe-

cies. Dall stated that these were "the two species and

only two species of which it is composed, even at the

present day." While not mentioned by Dall, one of

these species was Distorsio anus (Linne, 1758), the

type for the genus, and the other was listed and dis-

cussed by him as Distortrix reticulata Link, 1807,

which is Distortio reticulata Roding, 1 798. Dall divid-

ed reticulata into "variety reticulata" and "variety

clathrata" and considered it to be the only species in

the Western Atlantic. Curiously, the taxon that Dall

considered to be "variety clathrata" is the subspecies

now known as D. constricta macgintyi, while what he

regarded as "variety reticulata" is actually Distorsio

clathrata {Lamarck, 1816).

30

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

Fig. 1, Protoconch o/Distorsio perdistorta,y4ca(iem'

of Natural Sciences of Philadelphia, 241660, 100 fms.

Tosa, Japan, height 2 mm.

The history and synonomy of A reticulata Roding

was discussed and clarified by Puffer (1953, p. 1 13).

The distribution of reticulata and anus is limited to

the Indo-Pacific.

Emerson and Puffer, 1953, and Clench and

Turner, 1957, limited the living species in the Western

Atlantic to Distorsio clathrata and Distorsio mac-

gintyi. Neither Dall nor later authors realized that the

"Blake" expedition had actually collected a third and

different species. I found seven specimens of this spe-

cies (four from the dry collection and three from the

alcohol collection) in the USNM. Some of these

proved to be the actual specimens listed by Dall as

"Distortrix reticulata" Link and were still labeled as

such. They match the holotype, and other specimens

of Distorsio perdistorta recently collected in the

Western Atlantic in every detail, including penis,

periostracum and operculum, althougli the shells are

smaller in size.

Examination of the literature of fossil mollusca

has failed to reveal any record of an ancestor of Dis-

torsio perdistorta in the Western Atlantic althougli

there is ample evidence of the ancestors of Distorsio

clathrata and Distorsio macgintyi. A special effort

was made to investigate the possibility that a Panamic

species, such as Distorsio decussata (Valenciennes,

1832), could prove to be a Panamic analog of Dis-

torsio perdistorta in either Recent or fossil form.

Woodring, (1928, p. 495, pi. 18, figs. 7, 8, 9) illust-

rates a mixture of species which he misidentifies as

"Distorsio decussatus simillimus (Sowerby, 1850)".

Figures 7 and 8 are actually Distorsio gatunensis

Toula. 1909, the fossil subspecies of Distorsio

decussata. Figure 9 is Distorsio simillima, the fossil

subspecies of constricta. The differences between per-

distorta and the living and fossil forms of decussata

suggest that a close relationship between the two spe-

cies is unlikely, in spite of their relatively close distri-

bution. This seems especially borne out when one ob-

serves the great similarity of Western Atlantic,

Japanese, and hidian Ocean specimens, in spite of

their extreme geographical separation.

Fig. 3, Left jaw of Distorsio perdistorta Fulton, Gulf

of Mexico, length: .75 mm.

Fig. 2, Radula of Distorsio perdistorta Fulton, 105

fms, WSW Tampa, Fla., width of rachidian: 120^.

Vol. 86 (2 - 4) THE NAUTILUS

TABLE 2. Specimens of Distorsio constricta habei Lewis, new subspecies, examined.

31

32

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November 1972

Vol. 86(2-4)

Fig. 4, Protoconch of Distorsio constricta habei

Lewis, height of protoconch: 1.8 mm.

Genus Distorsio Rbding, 1798

Distorsio Roding, 1798, Museum Boltenianum, pt. 2,

p. 133. Type species, Murex anus Linne, 1758,

subsequent designation J. E. Gray, 1847. (The

Latin noun distortio is feminine).

Description - Shells range in length from 25 to 90

mm. They are strongly sculptured with spiral cords

and axial ribs which can make them knobby or can-

cellated. The whorls are swollen and distorted be-

cause they bulge in order to accommodate the grow-

ing animal as it covers earlier whorls. There is a large

parietal shield which joins with an expanded

peristome to encircle the aperture. When partially

covered by new growth, the old parietal shields serve

as varices. The distorted whorls, parietal plicae and

grossly-formed denticles on the inner edge of the out-

er lip, create a very irregular aperture. The shield has

a groove and numerous irregular plicae on the lower

left. A short anterior siphonal canal usually turns up-

wards.

The radula is taenioglossate. It is situated in the tip

of an extremely long proboscis which lies folded in

the proboscis sheath (figs. 14 to 22). The proboscis is

capable of tremendous extension. This is typical of

the genus and quite different from the typical

cymatiid pleurembolic proboscis. In species which

have been examined, the jaws are small, transparent,

and not as strongly formed as in most genera of

Cymatiidae.

OPERCULA

Previous workers have suggested that the pattern

of the opercula of Distorsio varies intraspecifically be-

tween terminal (without a nucleus) and submarginal

(with a nucleus). Personal observation of numerous

broken and repaired opercula has indicated that there

is a high incidence of breakage and repair. This is

probably due to the difficulty of passing the

operculum through the extreme constriction of the

aperture. It is interesting to note that all of the

opercula of Distorsio which show clear evidence of re-

pair have a nucleus. Missing opercula are replaced by

newly formed ones, which also always have a nucleus.

1 have observed this characteristic change in pattern

from terminal (a-nuclear) to nuclear in the repaired

opercula of other genera of Cymatiidae such as

Cymatium s.s., Gyrineiim and Monoplex. Often the

growth rings depart from the semicircular arc which is

typical of the original terminal pattern. Furthermore,

the specimens which have remained terminal are

usually consistent in size and shape, while the nucle-

ated opercula vary considerably. When dealing with

perfect specimens of the opercula of Distorsio, the

primary muscle scar pattern and the shape delineated

by the varnish can prove to be a valuable taxonomic

character.

Distribution - Species of Distorsio are found in

most tropical portions of the world to a depth of 300

Fig. 5, Operculum of Distorsio perdistorta Fulton,

dredged WSW Tampa, Fla., 105 fms., height 9 mm.

hlg. 6, Operculum of Distorsio perdistorta Fulton,

Academy of Natural Sciences of Philadelphia,

241652, Tosa, Japan, height 7 mm.

I'ig. 7, Operculum of Distorsio perdistorta Fulton,

Academy of Natural Sciences of Philadelphia,

241652, Japan, height 6.9 mm.

Fig. 8, Operculum of Distorsio clathrata Lamarck,

140 ft. S.W. Cape San Bias, Florida, 1969, height 5

mm.

Fig. 9, Operculum of Distorsio constricta macgintyi

Fmerson & Puffer, South of Loggerhead Lt., Tor-

tugas, Fla., 25 fms., height 7.8 mm.

Fig. 10, Operculum (^>/ Distorsio habei Lewis, Acade-

my of Natural Sciences of Philadelphia, 189639, Tosa

Bay, Japan, 70 fms.. height 6 mm

Vol. 86 (2 - 4)

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33

8

10

Fig. 5-10, Explanation on opposite page.

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November 1972

Vol. 86 (2 - 4)

fathoms. The earliest American fossils of the sub-

genus Persunella appear to be from the Middle

Eocene, while the earhest Distorsio s.s. appears to be

Distorsio crassidens (Conrad, 1848), from the Oli-

gocene of Vicksburg, Mississippi

REMARKS ON THE SUBGENERA

Subgenus Rhysema Clench. Wm. J. and Turner.

Ruth D. 1957. Johnsonia, vol. 3, no. 36, p. 236. Type

species clathratus Lamarck, 1816, by original designa-

tion.

When Clench and Turner, 1957, described the sub-

genus Rhysema, they differentiated it from Distorsio

s.s. on the basis of two characters. They pointed out

that Distorsio anus, the type for Distorsio s.s., had a

parietal shield which "extends well above the upper

limits of the outer lip and covers the preceding two

whorls" and a siphonal canal which is "nearly verti-

cal." This limited Distorsio s.s. to one species and all

other living species were then considered to be in the

subgenus Rhysema. Up to the present day, there has

been no additional evidence to justify two subgenera.

It is true that the various species which comprise

Rhysema, do not have a parietal shield which covers

two preceding whorls, but the shield often covers one

preceding whorl and quite often exceeds the upper

limits of the outer lip. Furthermore, there are many

specimens of various species whose siphonal canals

are very recurved, although not vertical. These charac-

ters appear to be based upon differences of relative

degree. Wliile they miglit be considered valid when

separating species, they do not appear to define sep-

arate subgenera. A new species is described later in

tins paper which is very similar {o Distorsio anus. Be-

cause of this similarity most workers would agree that

it is properly placed in Distorsio s.s. The definition of

Rhysema would require placing it in Rhysema since

its parietal shield does not exceed the two preceding

whorls and its siphonal canal is not vertical. This

example alone demonstrates the difficulty one would

have in attempting to place the various species of Dis-

torsio into separate subgenera based on these charac-

ters.

As an added example, Distorsio clathrata differs as

much from Distorsio maegintyi as does Distorsio anus

from cither one. Distorsio anus is often considered a

shallow water species but. living specimens have been

found in 45 meters. While the various species of

Rhysema are considered to be from deep water, some

species have been found living intertidally. Until more

evidence becomes available, it seems unnecessary to

retain two subgenera which are so weakly differenti-

ated. Consequently, I consider Rhysema to be a syno-

nym oi Distorsio.

Distorsio perdistorta Fulton, 1938

Figs. 1,3, 5-7, 11-34

Synonomy -

1938 Distorsio perdistorta Fulton, Proceedings of the

Malacological Society of London, (March 16), vol.

23, pt. 1, pp 55-56, pi. 13, figs, 3 and 3a. Type

locahty; Kii, Japan. Type in the British Museum of

Natural History

1964 Distorsio (Rhysema) horrida Kuroda and Habe.

Shells of the Western Pacific in Color, vol. 2, p. 74,

pi 23, figs. 3 and 3 (both figures numbered 3).

Type locality, Tosa Bay, Shikoku, Japan. Type in

the National Science Museum, Tokyo, Japan.

Description - The largest shell examined reached

82 mm. in length. The color of the shell is white, but

pale orange-brown color may be present on the spiral

cords. There are 8 to 10 convex whorls producing a

spire of approximately 50°, The outer lip is thickened

with the outer margin curving forward. There are 8 or

9 denticles on the inner edge of the outer lip The

third denticle below the posterior anal canal is much

larger than the others and is opposite a deep indenta-

tion in the parietal wall, creating a constricted

apertural shape. The parietal shield is thin, rather ob-

long, being wider posteriorly and narrowing anterior-

ly. The shield can be smooth or sculptured with very

low spiral cords and axial ribs which form beads when

they cross. There are numerous irregularly formed

plicae on the left of the groove in the lower parietal

shield. These are variable in sculpture and number,

sometimes reaching the edge of the shield. The groove

itself is more open and straighter than in most species

of Distorsio. The siphonal canal is relatively straiglit

for the genus, short and turned slightly upwards. It is

bordered by 9 to 15 well-defined plicae, the greater

number usually occurring on the larger specimens.

The upper three plicae arc the largest and the re-

mainder diminish in size. The sculpture consists of 8

major spiral cords on the body whorl, the upper 5 or

6 being separated from one another by a single fine

interstitial cord. There are 8 or 9 varices with 20 to

Vol. 86 (2 - 4)

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35

these papillae were

■i^'y'-^ky^ip^^-^':^^ y presumed missing

t

Fig. 11, Dorsal aspect of the mantle edge o/Distorsio

perdistorta Fulton (female) flattened to show papil-

lae.

Fig. 12, Dorsal aspect o/Distorsio perdisioita Fulton

(male). The mantle edge of this specimen lacks papil-

lae.

\

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November 1972

Vol. 86 (2 - 4)

25 axial ribs between them which form beads when The operculum is small and distinct from that of

they cross and give the shell a reticulated appear- Distorsio clathrata although it is similar to Distorsio

ance. macgintyi (figs. 5 to 10).

exit of proboscis

sheath

proboscis

fully

retracted

proboscis sheath

opened dorsally

Fig. 13, Right lateral view o/ Distorsio perdistorta Fig. 14, £)orsa/ v/ew o/ Distorsio perdistorta Fw/ron.

Fulton.

Vol. 86 (2 - 4)

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The soft body of the animal is tan, lacking the

spots typical of many Cymatiidae. The mantle edge

of the male animal examined was lacking in papillae

(fig. 12); however the female specimen has large ob-

vious papillae which are bisected by an open groove

(fig. 11). Since only two specimens were examined,

the differences in the mantle edge should not be con-

strued as evidence of sexual dimorphism. The penis is

very large with an open penial groove (figs. 12- 13),

The rachidian and laterals of the radula are very

powerfully formed (fig. 2). There is a long central

cusp on the racliidian with 4 to 6 sharply-pointed

shorter cusps on either side of it. The lateral has 6 to

9 small, sharply pointed cusps on the anterior lateral

edge. There are no cusps on the marginals. Because of

the extraordinary proboscis structure and the ex-

tremely small opening througli which the proboscis is

extended, the radula, jaws and buccal mass are very

small in relationship to the body mass,

PERIOSTRACUM

The periostracum is yellowish and formed in nu-

merous very low axial blades which are fringed with

fine projections along their edges. There are long

hair-Uke processes at the beads of the shell sculpture

which gives fresh specimens a bristly appearance. Al-

though the Japanese specimens of this species are

more sparsely bladed than the Western Atlantic spec-

imens, both exhibit an extraordinary periostracal

structure. Below the visible upper surface, there is a

second layer of tunnel-like chambers which run spiral-

ly around the shell (figs. 23 to 25). There is a perio-

stracal floor which is attached to the shell and a roof

which is parallel to the floor and shell surface. The

roof is formed by flat connections between the small

periostracal processes. The axial walls of the tunnels

are formed by the axial blades and the .spiral walls by

flat, vertical connections between the processes. I

have observed similar structures in Distorsio reticu-

lata, D. clathmta, A decussata, A constricta con-

stricta, and D. constricta macgintyt Two species

which do not have this type of periostracum are A

anus and A constricta habei.

DISTRIBUTION

The distribution of Distorsio perdistorta has pre-

viously been known from Tosa Bay, Shikoku, Japan

to Kii, Honshu, Japan. Based on the hmited amount

of material which has been collected in the Westeri.

Atlanfic, it is difficult to judge the relative rarity of

this species, but it certainly can be said that it has not

proved to be as common as Distorsio clathrata or Dis-

torsio macgintyt Thus far the northern record in the

Western Atlantic is west of Tampa, Florida, Gulf of

Mexico, while the southern record is off Barbados, in

the Lesser Antilles, Recently, two specimens were re-

corded from north-west Madagascar in the

Mozambique Channel, Indian Ocean.

COMPARATIVE FEATURES

Fulton described Distorsio perdistorta as white,

but examination of numerous specimens indicates an

occasional pale-brown coloration limited to the spiral

cords. This characteristic position of color pigment is

present in both Western Atlantic and Japanese spec-

imens, and appears to be a useful character in Dis-

torsio.

Some workers might be tempted to designate the

Western Atlantic representatives as a subspecies of the

Japanese perdistorta because of geograpliical separa-

tion, heavier periostracum and an apparent tendency

to reach a larger size. However, careful examination

of all the material failed to show that these were sig-

nificant differences. On the contrary, these characters

varied within each geographical range. My work with

the Cymatiidae has clearly demonstrated to me their

great ability for intraspecific variation. Laxton,

(1971), has shown that two distinct populations of

Cymatium spengleri (Perry 1811), can vary in spire

angle, number of varices, and other characters, when

each lives in different ecological situations and feeds

upon different species of ascidians which are available

in different quantities. Bayer, (1971, pp. 114-115)

discusses the close resemblance of various Japanese

and Caribbean genera and species. It is widely known

that other species of Cymatiidae are worldv^de in dis-

tribution with various unexplained geograpliical rela-

tionsliips. The morphological consistency of the

Japanese and Western Atlantic specimens of Distorsio

perdistorta makes it unnecessary to estabhsh a new

subspecies in spite of their great geographical separa-

tion.

In order to clearly differentiate the threQ Hving

species of Distorsio in the Western Atlantic, I have at-

tempted to estabhsh a diagnostic chart of their more

obvious differences.

38

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November 1972

Vol. 86 (2 - 4)

Distorsio constricta habei

new subspecies, Lewis

Figs. 4, 10,38,39

Synonomy -

1964 Distorsio ( Rhyscma) perdistorta Fulton,

Kuroda and Habc, Shells of Ihc Western Pacific in

Color, vol 2, p. 74, pi. 23, fig. 1 (non Fulton,

1938).

Description - The largest shell examined reached

54 mm. in length. There are 9 convex whorls which

have a Hallened plane above the periphery formed be-

tween the first and second spiral cords. The spire is

produced at approximately 42°. The outer lip is

slightly thickened with the outer margin curving for-

ward. There are 8 denticles on the right edge of the

aperture which extend to the edge of the outer lip

and form low cords. The third denticle below the pos-

terior anal canal is much larger than the others and is

opposite a deep indentation in the parietal wall. The

I'ig. 15-19, Dissection of the tip of the proboscis of

Distorsio perdistorta Fulton (dorsal aspect):

hni buccal mass;

c. esophagus;

ev. esophagal valve;

ie. incision in esophagus;

j. jaws positioned considerably posterior to lip of

oral tube;

lo. crenulated lip of oral tube (probably used for

ingestion of food);

Ip. lip of proboscis;

me. nmscles of the esophagal valve;

parietal shield is thin, oval and variably sculptured by

spiral cords and axial ribs which form low beads when

they cross. Color is a diffused pale orange-brown,

with more intense color limited to the spiral cords.

There is a clearly formed groove in the lower parietal

shield, bordered on the right by 8 to 12 denticles on

the columcllar edge of the siphonal canal, The first

denticle is the largest and the remainder diminish in

size. There usually is dcnticulation on the left side of

the groove, but in occasional specimens, the groove

blends into the parietal shield without delineating

dcnticulation. The siphonal canal is straight, short

and curves slightly upwards^ The sculpture consists of

8 major spiral cords on the body whorl and 3 cords

on the dorsal surface of the anterior canaL Between

the first and second cords below the suture, there are

2 or 3 fine interstitial cords The second and third

cords are close together with one fine interstitial cord

separating them. The third and fourth cord are sep-

arated by 2 fine interstitial cords, while the remaining

Vol. 86(2-4)

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39

Fig. 15-19, Explanation on opposite page.

40

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November 1972

Vol. 86 (2 - 4)

(N

'■■^Ji

in

o

I/)

d)

(U

0)

(/)

•5*;;^

A.i;-^5

t

^

^ ^

^

to

to

in

bo

Vol. 86(2-4)

THE NAUTILUS

25 shell surface hollow chambers

Fig. 23, Single periostracal process o/Distorsio perdistorta Fulton.

Fig. 24-25, Details of periostracal structure o/Distorsio perdistorta Fulton, 125 fms. W. of Ft. Myers, Fla.

42

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November 1972

Vol. 86 (2 - 4)

major cords are usually separated by a single fine

cord. There are 8 to 10 varices with 12 to 17 axial

ribs between the varices. Where the cords cross the

axial ribs, they form beads.

The periostracum is thin, yellowdsh, formed in

spiral rows which follow the cords of the shell in con-

trast to the numerous, fringed, axial blades of the

other species. There are thin, low hairlike processes at

regular intervals on the spiral rows and they align

axially. Longer processes develop on the ribs of the

shell The periostracum is attached directly to the

shell and lacks the second layer such as that found in

perdistorta and the other two subspecies, constricta

and macgintyU The operculum is terminal (fig. 10). I

have not examined an animal of this species.

Description of the holotype - The sliell measures

53 mm. in length and 28 mm. in width. There are 8

denticles on the riglit edge of the aperture which ex-

tend across the expanded peristome to the edge of

the outer lip, forming low cords. The lower left edge

of the parietal shield is lacking in denticulation. There

are 9 denticles on the columellar edge of the shield.

There are 1 5 axial ribs on the body whorl.

Distribution and type locality - Most of the spec^

imens which I have examined were taken in Tosa Bay,

Shikoku, Japan, which is designated the type locality.

There is one specimen from Kii, Honshu, Japan.

Kuroda and Habe list the distribution as Boso Penin-

sula, Honshu, to Tosa Bay, Shikoku, Japan at depths

of 100 to 200 m They state that the species is un-

common. The holotype is in the ANSP no. 325380;

four paratypes in ANSP 325381 ; one paratype in Del.

Mus. Nat Hist. no. 50943.

Differentiating features - Distorsio habei is Very

similar in general appearance to Distorsio constricta

and Distorsio macgintyi The differences are not sig-

nificant enough to justify specific separation. How-

ever, unlike the Western Atlantic specimens of Dis-

torsio perdistorta, they can be separated when the

three subspecies are compared. The taxononiic char-

acters wliich are consistent enougli to use are: pig-

mentation, the pattern of cords and interstitials, den-

ticulation on the expanded peristome and the struc-

ture of the anterior siphonal canal.

The shells of all three subspecies are irregularly

stained with orange-brown, but the cords on the shell

of habei are always colored v^th a darker pigmenta-

tion. On the parietal sliield of macgintyi and con-

stricta there are strongly formed white beads outlined

by a rich brown color in between the beads which

gives the shields a markedly reticulated appearance.

Tliis character is very consistent in macgintyi but

varies in some specimens of constricta. 1 have exam-

ined large specimens of constricta from the Galapagos

Islands which have no beading on the shield at all. All

of the specimens of habei examined had beaded

sculpture on the sliield, but the strength of the bead-

ing and color are variable, lower, and finer than the

beading and color on macgintyi The shield of habei is

either white or very liglit orange-brown. In some spec-

imens of habei, the parietal shield was well-below the

suture, but all of the specimens of constricta and

macgintyi which I have examined had parietal shields

which were at the suture or above it.

The regular pattern of interstitial cords which is al-

ways present in habei is absent from constricta and

irregularly variable when present in macgintyi In

habei, the riglit posterior edge of the anterior canal

slants toward the posterior columellar edge, almost

closing the canal externally at the point where they

Fig. 26-27, Distorsio perdistorta Fulton, Holotype, British Museum (Natural History), Kii, Japan, 60 mm. x 35

mm

Fig. 28-29, Distorsio perdistorta Fulton. Gulf of Mexico, 57 mm. x 32 mm.

l-ig. 30-31, Distorsio perdistorta, Fw/fow. West of Tampa, Fla. llOfms. The heavy periostracum on this specimen

is variable within the species. 66 mm x 36 mm.

I'ig. 32-33, Distorsio perdistorta Fulton. Tosa Bay, Japan, 63 mm. x 33 mm.

I'ig. 34, Distorsio perdistorta, Fulton. Dredged West of Tampa, Fla. This specimen has a periostracum that closely

corresponds to the average Japanese specimen. 82 mm. x 44 mm.

l-ig. 35, Distorsio clathrata Lamarck. Dredged 65 fms. offTortugas, Florida, 67 mm. x 36 mm.

I'ig. 36-37, Distorsio decussata, Valenciennes. Trawled near Topolobamp, Sinaola, Mexico, 40 fms. 64.5 mm. x

33 mm

Vol. 86 (2 - 4)

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43

Fig. 26-37, Explanation on opposite page.

44

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November 1972

Vol. 86 (2 - 4)

come together. The edges of the anterior canals of

constricta and macgintyi remain parallel and leave the

canal open.

While discussing the holotype oihabei, I described

how the denticles on the right edge of the aperture

continue across the expanded peristome to the outer

edge of the shield, forming low cords. If the cords do

not continue from the apertural edge, there are no

denticles formed at the outer edge of the shield, al-

though there is a shallow groove which runs parallel

to the aperture down the length of the expanded

peristome. In both constricta and macgintyi, tliis

groove is stained with a darker orange-brown color.

Clearly defined denticles are formed on the right out-

side edge of the shield. Although separate, these

denticles align with those at the apertural edge.

In the specimens examined there were 8 to 13

axial ribs between the varices of constricta, (an aver-

age of 10.6); 11 to 20 on macgintyi (an average of

14.7); and 12 to 17 in habei (an average of 15.6).

Distorsio habei differs from Distorsio perdistorta

by being smaller, more distorted, having a more

angled whorl which is tabled above the periphery,

fewer axials (15.6) than perdistorta (20 to 25), a dif-

ferent pattern of interstitial cords, a different perio-

stracal structure and richer pigmentation than the

Japanese specimens of perdistorta. The spire angle of

habei is 42 while the spire angle of perdistorta is

50°.

HISTORICAL DISCUSSION OF HABEI

Kuroda and Habe in "Shells of the Western Pacific

in Color", (1964, vol. 2, p. 23, figs. 1 and 3) des-

cribed Distorsio horrida (their fig, 3) as a new species

and differentiated it from what they identified as

"Distorsio perdistorta Fulton" (their fig. 1). Exami-

nation of a photo of the type specimen of Distorsio

perdistorta kindly supplied by the British Museum of

Natural History and correspondence with Dr.

Tadashige Habe of the National Science Museum of

Tokyo proved that Distorsio horrida Kuroda and

Habe, 1964, is a synonym of Distorsio perdistorta

Fulton, 1938, and that "Distorsio perdistorta"

Kuroda and Habe (not Fulton) is an unnamed species.

This taxon is described as Distorsio constricta habei

in honor of Dr. Tadashige Habe whose many contri-

butions to malacology are known througliout the

world. Careful comparison of this species with spec-

imens of worldwide species of Distorsio has led me to

realize that Distorsio constricta habei from Japanese

waters along with Distorsio constricta constricta from

the Eastern Pacific and Distorsio constricta macgintyi

from the Western Atlantic are geographical sub-

species. These three subspecies give evidence once

again of the amazing distribution of some of the

Cymatiidae.

FOSSIL RELATIVES

There is some confusion in the literature that deals

with the ancestors of the recent Caribbean and

Panamic species of Distorsio, Various authors have

considered the Antillean Miocene fossil Distorsio

simillima (Sowerby, 1 850), to be a paleosubspecies of

three different recent Distorsio i.e.: decussata,

clathrata and constricta. The matter was further com-

plicated because they confused specimens of Dis-

Fig. 38-39, Distorsio constricta habei Lewis, Holotype, Tosa Bay, Shikoku, Japan, 53 mm. x 28 mm.

Fig. 40, Distorsio constricta macgintyi Emerson and Puffer, dredged 280 ft. S. W. of Key West, Fla., 43 mm. x 25

mm.

Fig. 41, Distorsio constricta constricta Broderip, dredged 200 m, off Southern Coast of Santa Cruz Is., Glalpagos,

49 mm x 27.5 mni

Fig. 42, Distorsio anus (Linne). Hawaii

Fig. 43-44, Distorsio burgessi Lewis, Hawaii, Holotype, The Academy of Natural Sciences of Philadelphia, No.

326470, 39 mm x 25 mm

Fig. 45-46, Distorsio ridens Reeve, Philippines, Syntype, The British Museum of Natural History, No. 1967630,

77.5 mm x 38 mm.

Fig. 47, Reeve's figure o/ Distorsio ridens.

Fig. 48, Distorsio ridens Reeve, Lectotype, American Museum of Natural History, 64.5 mm. x 35 mm

Vol. 86 (2 - 4)

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:p:m

47

F/^. 38-48. Explanation on opposite page.

46

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November 1972

Vol. 86 (2 - 4)

torsio simillima with Distorsio gatunensis Toula, 1909

(Miocene, Panama). Rutsch, 1930, (pp. 6ia61 1, 614,

pi. 17, fig. 6) examined and figured the holotype of

gatunensis and concluded that this species is actually

the fossil form of the recent decussata. This was con-

firmed by Emerson and Puffer, 1953, (p. 100) and

Weisbord, 1962, (p. 271). Woodring, 1928, (p. 300,

pL 18, figs. 7 to 9) illustrates two species under the

name "Distorsio decussatus simillimus". Actually, fig.

9 and fig. 1, pi. 19, are a specimen of Distorsio

simillima, the ancestor of constricta. Figures 7 and 8

are Distorsio gatunensis Toula, the ancestor of

decussata. A possible sequence of evolutionary events

indicates that the first true Distorsio s.s. was

crassidens (Conrad, 1848), Vicksburg, Mississippi,

Middle Oligocene, which spread throughout the

Caribbean developing into simillima during the Mio-

cene and eventually into the recent macgintyi,

Pilsbry, (1922, p. 360,) considered crassidens to be a

fossil subspecies of Distorsio constricta. However, he

was unaware of the existence of macgintyi, since that

species was not reported until 1951, and he was

under the impression that simillima had died out in

the Caribbean without a recent form. Fossil evidence

shows that simillima existed in the Tertiary Caribbean

faunal province from the Lower Miocene to the Plio-

cene and in the Eastern Pacific from the Middle Mio-

cene to the Pliocene. It must be remembered that the

Tertiary Caribbean Province extended into the East-

ern Pacific from southern Nicaragua to northern Peru.

Woodring, (1966, p. 427) suggests the possibility of

transportation of planictonic larvae by the Miocene

North Equatorial current along the south border of

the Canbbean Sea, through the Atrato Strait (and

others) into the Eastern Pacific. It has been reason-

ably established that this faunal migration was at its

heiglit during the Middle Miocene, a time which

corresponds with the fossil presence of simillima in

the Eastern Pacific. Since simillima was unknown in

the Eastern Pacific during the Lower Miocene, we can

assume that it existed in the Caribbean province for

approximately 5 million years before migration into

the Eastern Pacific. It is likely that Distorsio

macgintyi was the first recent species to develop from

the crassidens-simillima stock. Distorsio simillima,

having migrated to the Eastern Pacific during the Mid-

dle Miocene, became isolated at the end of the Terti-

ary and early Pleistocene after closure of the Central

American land bridge, and there was modified into

the present-day constricta. Eastern migration of an-

cient simillima accounts today for the Japanese sub-

species Distorsio constricta habei

Distorsio burgessi new species, Lewis

Figs. 43, 44

In August, 1963, an unidentified species of Dis-

torsio was illustrated by Dr. C. M. (Pat) Burgess on

the first page of vol. 1 1, no. 10, of the Hawaiian Shell

News. The specimen appeared to be closely related to

Distorsio anus Linne and until recently I have con-

sidered it to be a polymorphic form of A anus. Re-

cently sufficient material has come into my posses-

sion to show that consistent differences do exist be-

tween the two species which suggests that they are

sympatric.

Distorsio anus is widespread in its distribution and

shows morphological variation within a given popula-

tion as well as between widely separated populations.

However, there is such a great intergrading of form

that it would be impossible to tell where a given spec-

imen was taken. In the case of Distorsio burgessi, its

distribution is hmited to Hawaii and though found

with D. anus, it can readily be separated from it.

Tliough D. anus is very common and D. burgessi rela-

tively rare, large selections of anus examined which

were taken in the same general area as burgessi fail to

show specimens which intergrade with burgessi The

first specimen illustrated in Hawaiian Shell News was

taken, along with two others, in 14 fathoms, but the

holotype and paratypes were found by diving in shal-

lower depths, dispelling the idea that burgessi might

be a deep water form of anus. The differences be-

tween the two species are consistent but do require

careful observation. If the species were not sympatric

it would seem more reasonable to consider burgessi as

a subspecies of anus.

Distorsio burgessi is named in honor of Dr. C. M.

Burgess who first published it as an unidentified spe-

cies.

Description - The largest shell examined reached

60 mm. in length. There are 9 convex whorls which

have a flattened plane above the periphery formed be-

tween the first and second spiral cords„ The periphery

is defined by the second and third spiral cords joining

to form a double cord The spire is produced at an

angle of approximately 53°. Tlic outer lip is fonned

by the right edge of the parietal shield which projects

slightly past the' body whori. Behind the shield at the

point where it rneets the body whori there is a strong

axial cord which is separated from the preceding axial

Vol. 86(2-4)

THE NAUTILUS

cord by a deep groove. Tliis groove is crossed by the

spiral cords and gives the impression of a deep perfor-

ated line. The outer edge of the parietal shield is shal-

lowly ruffled by 9 low cords which are separated by 8

dark brown shallow grooves. Larger specimens have a

row of sharply formed teeth at the beginning of each

dark groove arranged in a line parallel to the outer Up.

These teeth are separated from the apertural denticles

by a secondary groove which is also parallel to the

outer lip and is liglitly stained in a dotted line be-

tween the teeth. The parietal shield is the most strik-

ing character of this species. It is ear-shaped, and the

lower left edge of the shield is flattened and conforms

to the body whorl behind it. The main mass of the

shield is sculptured by 9 strong, regular, spiral cords

and 5 or 6 axial ribs wliich create a checkerboard pat-

tern This effect is greatly enhanced by the rich

dark-brown pigmentation in the grooves between the

squares of the pattern. An unusual appearance is cre-

ated along the left and upper edges of the parietal

shield because the ribs stop before the left edge and

the spiral cords continue, while the cords stop before

the upper edge and the axial ribs continue. The aper-

ture is typically irregular and with 9 denticles. The

third, wliich is the largest, is opposite a deep indenta-

tion in the parietal wall. The shell is wliite, liglitly

stained with a very pale orange. The columellar phcae

and siphonal canal are straiglit and aligned on the axis

of the shell. The siphonal canal is very short and re-

curves dorsally at an angle of approximately 115°.

The sculpture consists of 8 spiral cords on the body

whorl and one cord on the dorsal surface of the

siphonal canal. There are usually 10 to 12 major axial

ribs which cross the cords and form low nodules^ The

periostracum is yellowish, very flat and thin, formed

in a regular pattern of low hairlike processes, with

slightly larger processes forming on the axial ribs. It is

attached directly to the sheU surface, lacking the

second layer found in perdistorta. I have not exam-

ined an animal nor the operculum of this species.

Description of the holotype - The shell measures

39 mm. in length and 25 mm in width. There are 12

denticles along the columellar edge to the end of the

siphonal canal. There are 12 major axial ribs on the

body whorl. The specimen has its periostracum in-

tact. Holotype ANSP no. 326470. 2 paratypes in the

Hal Lewis collection. 1 paratype Del. Mus. Nat. Hist.

1 paratype in the CUfton So Weaver collection.

Distribution - All of the specimens examined were

taken in Hawaii at approximately 21.1 8N Long.

158.07 W. Lat. off Oahu Island, Type locality:

Barber's Point, S. W, Oahu Island, Hawah. Collected

by E.

Differentiating features - This species differs from

Distorsio anus primarily on characters relating to the

parietal shield and anterior siphonal canal. The parie-

tal shield of Distorsio anus is oval to almost round

with deep ruffles usually present around the peri-

meter of the shield. It rises above the body whorl to

completely cover the preceding two whorls. In Dis-

torsio burgessi the ruffled edge is restricted to the

outer lip and the shield covers approximately Wi pre-

ceding whorls. The shield of anus is white to diffused

orange-tan, lacking the rich dark-brown pigmentation

typical of burgessi. The sculpture on the shield of

both species consists of 9 spiral cords but on anus

there are usually more numerous axial cords, giving

the shield surface a finer beaded and wrinkled look.

Even when the sculpture is coarser, it lacks the regu-

lar checkerboard pattern of burgessi. The anterior

canal of anus is always angled to the left, often curv-

ing slightly, in contrast to the straiglit axially aligned

canal of burgessi. The siphonal canal of anus is longer

and recurves dorsally at an angle of approximately

90°. The groove to the left of the siphonal canal on

the lower left parietal shield of anus opens into the

aperture via a secondary groove which is framed by

two large plicae. All of the specimens of burgessi

which were examined did not have this secondary

groove, and the primary groove in the lower left

shield was reduced, thinner and shallower than in

anus, forming a narrow dark-brown passage which fol-

lows the edge of the columellar-siphonal denticles. In

specimens examined, there were 10 to 12 axial ribs

on burgessi and 14 to 16 axial ribs on anus.

In spite of the rich pigmentation present on the

parietal shield of burgessi, it is lacking in the rich

body whorl pigmentation typical of Distorsio anus.

This appears once again to be an indication of the

specific importance of the position of pigmentation

on the shells of Distorsio.

Dis torsio ridens Reeve, 1844

Figs. 45-48

In the course of my work with the Cymatiidae, I

have observed that the species known as ridens Reeve

is often misidentified in collections. This very uncom-

48

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

mon species has had equal inistreatment in the litera-

ture. It has been listed as reticulata Roding (Maxwell

Smith, 1948, p. 23); properly figured but misidenti-

fied as reticulata (Wagner and Abbott, 1967, p. 85,

fig. 13-11 5); been confused with Distorsio smithi von

Maltzan, 1884 (Nickles 1950, p. 86, fig. \2>3),perdis-

torta Fulton, 1938 (Oyama, 1957, pi. 1, figs. 7-8),

and, in many other instances too numerous to men-

tion, improperly synonymized, misidentified and mis-

figured. The actual species described and figured by

Reeve (1844, Triton, pi. 12. sp. 46) has been subse-

quently figured properly by Try on (1881, pi. 17, fig.

177) and Webb (1935, pi. 51, fig. 10). Both of these

figures are copies of Reeve's original figure, and it is

probable that neither author had actually seen spec-

imens of the species. Oddly enough, I have been un-

able to find a published photograph of rid ens.

A recent visit to the British Museum of Natural

History to study types revealed that none of the

syntypes labeled ridens matched the actual specimen

figured by Reeve. It is important to realize that

Reeve's cymatiid drawings are accurate depictions of

the specimens which they represent, matching them

very well in size, color pattern, details of breakage,

etc. All of the syntypes were less colorful, and none

matched the figure in size, although they were clearly

the same species (fig. 45). However, there is a spec-

imen in the collecfion of the American Museum of

Natural History New York (cat. no. 6369) (fig. 48)

which so very closely matches Reeve's figure (our fig.

47) that I feel it is reasonable to designate it as the

lectotype. The specimen measures within a millimeter

of the figure, matches it exactly in the outline of the

shell, shape of the parietal shield, position of color

(absent from the syntypes), spire angle, broken

siphonal canal, number of denticles on the columellar

edge and minor details of breakage. The only differ-

ences appear to be artististic exaggerations of a

sece)nd row of plicae on the lower left parietal shield

and the gross manner in which the groove on the low-

er parietal shield enters into the aperture. Both of

these characters are unique to the figure, being absent

from the syntypes and all other specimens examined.

1 have never seen characters such as these on any

specimen of any species of Distorsio. This specimen is

from the Wm. A. Haines collection, the bulk of which

was given to the AMNH in 1879. Haines, an American

conchologist of the 19th Century, exchanged with

many famous conchologists of the day, and could

have obtained the specimen through his friend John

C. Jay, or even directly from Reeve. While at the

British Museum of Natural History, I carefully exam-

ined every cymatiid in the collection, including ma-

terial not separated and distributed. There were no

other specimens of ridens Reeve that could be con-

sidered to match the figured specimen.

The ridens situation is further complicated by the

similarity of ridens to Distorsio decussata

Valenciennes, 1832. There is a claimed "paratype" of

ridens in the collection of the Museum of Compara-

tive Zoology at Harvard University (MCZ 186600 C.

B. Adams coll.) with Philippine data but which actu-

ally is a specimen of decussata. This misidentification

is common. Distorsio decussata can quickly be sepa-

rated from ridens by the fact that decussata always

has a double spiral cord at the periphery, a character

which is consistent in fossil as well as Recent spec-

imens. Myra Keen (1971, p. 508, sp. 962) lists the

range of decussata from Cape Tepoca and Guaymas,

Sonora, Mexico, south to Manta, Ecuador, offshore

to depths of 82 m., and she properly considers Dis-

torsio ridens of authors, not Reeve, to be a synonym

of decussata.

It is hoped that the discussion of this matter, pub-

lication of photographs of the lectotype, syntype and

Reeve's figure will serve to clarify this species and es-

tablish it as a valid taxon, separable from reticulata,

per distort a and decussata.

ACKNOWLEDGEMENTS

I wish to thank Dr. George Davis, Associate Cura-

tor, Department of Malacology of the Academy of

Natural Sciences of Philadelphia for his valuable criti-

cal comments on the manuscript; Dr. R. Tucker

Abbott, du Pont Chair of Malacology. Delaware Mu-

seum of Natural History, who initially suggested that

this paper be written and greatly assisted in its organi-

zation and nomenclature; Dr. Robert Robertson,

Pilsbry Chair of Malacology, Academy of Natural

Sciences of Philadelphia, for his suggestions and dis-

cussion of the manuscript; The British Museum of

Natural History. MoUusca Section, for their kind co-

operation during my visit, and Mrs. Way for the

photo of the holotype of Distorsio perdistorta: Dr.

William Emerson, Chairman of the Department of

Living Invertebrates, American Museum of Natural

History of New York, for drawing my attention to

Vol. 86 (2 - 4)

THE NAUTILUS

4>

certain references in the literature of fossils; Dr.

Joseph Rosewater, Division of Mollusks, National Mu-

seum of Natural History, Washington, D. C, for his

kind cooperation and the loan of valuable material;

Dr. Robert Bullock, Museum of Comparative Zoology

at Harvard College, for the loan of valuable material;

and to Dr. Tadashige Habe, Curator of Invertebrate

Zoology, National Science Museum, Tokyo, for his

kind cooperation and correspondence.

LITERATURE CITED

Bayer, Frederick M. 1971. Biological Results of the

University of Miami Deep-Sea Expeditions. 79.

New and Unusual Mollusks Collected by R/V John

Elliott Pillsbury and R/V Gerda in the Tropical

Western Atlantic. Bulletin of Marine Science. 21

(1): 11-236, figs. 1-72. Coral Gables, Fla. (Univer-

sity of Miami Press) (March).

Broderip, W. J. 1833. Proceedings of the Zoological

Society of London, pt. 1, p. 5.

Clench, William and Turner, Ruth. 1957. Johnsonia,

Monograph of the Western Atlantic Mollusks, The

Family Cymatiidae in the Western Atlantic 3 (36):

1 89-244, 26 pis. (December 20).

Conrad, T. A. 1848. Observations on the Eocene

Formation, and Descriptions of One Hundred and

Five New Fossils of that Period, from the Vicinity

of Vicksburg, Mississippi; with an Appendix. Jour-

nal of the Academy of Natural Sciences of

Philadelphia 1, ser. 2, p. 188, pi. 11, fig. 40.

Dall, WUliam Healey 1889. Reports of the Results of

dredging ... by the U. S. Coast Survey Steamer

"Blake", pt. 2, Gastropoda and Scaphopoda. Bul-

letin of the Museum of Comparative Zoology, 18:

492 pp., pis. 10-40 (June).

Emerson, William K. and Puffer, Elton L. 1953. Cata-

logue of the Mulluscan genus Distorsio (Gastro-

poda, Cymatiidae). Proceedings of the Biological

Society of Washington 66: 93-108 (August 10).

Fulton, H. C. 1938. Descriptions and figures of new

Japanese Marine Shells. Proceedings of the Mala-

cologjcal Society of London 23 (1): 55-56, pL 3,

figs. 3 and 3a (March).

Gardner, JuHa 1947. The Molluscan Fauna of the

Alum Bluff Group of Florida. U. S. Geological

Survey Professional Paper, 142-H, pt. 8: 493-656,

pis. 52-62.

Habe, Tadashige Dr. 1964. Shells of the Western

Pacific in Color. 2, 233 pp., 65 pis.

Keen, A. Myra 1971. Sea Shells of Tropical West

America, p. 508, figs. 962 & 963.

Laxton, J. U. 1970. Shell growth in some recent New

Zealand Cymatiidae. Journal Exp. Marine Biology,

vol. 4, pp. 250-260.

Lamarck 1816 - Tableau Encyclopedique et

Methodique, Liste, p. 4, Atlas 3, pi. 413, figs. 4a

and 4b.

Link, H. F. 1807. Beschreibung der Natur-

alien-Sammlung der Universitat zu Rostock.

Nickles, Maurice 1950. Mollusques testaces marins de

la Cote occidentale d'Afrique. Manuels Ouest-Afri-

cains, 2: 86 & 87, fig. 133.

Olsson, A. and McGinty, T. L. 1951. A Distorsio new

to the Florida Fauna. The Nautilus, Philadelphia

65(1): 26-28, pi. 1, figs. 5, 6, 10, 11, 12 (July).

Oyama, Katura 1957. The Molluscan Shells, Science

and Photography Club, Distorsio 1, figs. 1-12.

Pilsbry, Henry A. 1922. A Revision of W. M. Gabb's

Tertiary Mollusca of Santo Domingo. Proceedings

of the Academy of Natural Sciences of Philadel-

phia 73, pis. 16-47.

Puffer, Elton L. 1953. Distorsio reticulata vs. Dis-

torsio clathrata in the West Indies. Proceedings of

the Biological Society of Washington 66: 109-124,

pis. 6 and 7 (August).

Reeve, Lovell A. 1 844.. Conchologja Iconica 2, Tri-

ton, pis. 1-20.

Rutsch, R. von 1930. Einige interessante Pag-

stropoden aus dem Tertiar der Staatem Falc6n und

Lara (Venezuela). Eclog. Geol. Helve tiae, 23 (2):

604-614, pi. 17.

Smith, Maxwell 1948. Triton Helmet and Harp Shells,

p. 23, pi. 8, fig. 10.

Sowerby, G. B. II 1850. Descriptions of new species

of Fossil Shells found by J. S. Heniker Esq. The

Quarterly Journal of the Geological Society of

London, pt. 1, Proceedings of the Geological Soci-

ety 6: 44-53, pis. 9-10.

Toula, Franz 1909. Fine jungtertiare Fauna von

Gatun am Panama-Kanal. Jahrbuch der Kaiser-

lick-Konigliehen Geol. Reichsanstalt, pp. 673-760,

4 pis.

Valenciennes, A. 1832 (in) Humbolt and Bonpland.

Recueil d'Observations de Zoologje et d'Anatomie

Comparee 2: 306-307.

von Maltzan, Hermann Freiherrn 1884. Diagnosen

neuer Senegambischer Gastropoden. Nachrichts-

blatt der deutschen Malakozoologichen Gesell-

schaft, no. 5, pp. 65 & 66.

50

THE NAUTILUS

November 1972

Vol. 86(2-4)

Wagner, Robert J. L. and Abbott, R. Tucker 1967,

Von Nostrand's Standard Catalog of Shells. Second

Ed., p. 85, fig. 13-115.

Webb, Walter Freeman 1935. Handbook for Shell

Collectors, Sixteenth Ed., p. 105, pi. 51, fig. 10.

Weisbord, Norman E. 1962. Late Cenozoic Gastro-

pods from N. Venezuela. Bulletin of American Pa-

leontology, vol. 42, no. 193 (March 5).

Woodring, Wendell P. 1928. Miocene MoUusks from

Bowden, Jamaica, Part 2. Gastropods and discus-

sion of results, no. 385, 564 pp., 40 pis. (Carnegie

Institute of Washington Pub.).

Woodring. Wendell P. 1966. The Panama Land Bridge

as a Sea Barrier. Proceedings of the American

Philosophical Society, vol. 110, no. 6, pp.

425-433, 4 tables.

BOOK REVIEW

THE KINGDOM OF THE SEASHELL by R. Tucker

Abbott. Crown Publishers, Inc., 419 Park Avenue

South, New York, New York. 256 pp., 92 black

and white plates, 1 78 colored plates and 13 draw-

ings. 1972, $14.95.

This delightful book presents a whole spectrum of

topics on marine mollusks from their division into six

classes to how and where to find them; their breeding

habits, structure, coloration and the ways in which

they have been used in the arts, religion and history.

Certain families, such as the volutes, cowries, murices,

cones and scallops, are selected to portray their im-

portance to man.

Many of the colored plates are among the finest

illustrations that have ever been published in this

field. These, as well as the black and white photo-

graphs, were gleaned from several sources; all are ac-

knowledged at the end of the book. This volume is

not a textbook on mollusks, even though it contains

much general and basic information, but rather a por-

trayal of the many facets of this remarkable group of

animals. Dr. Abbott has an unusual ability to grasp

those facts which are important and interesting about

seashells and to combine them all in a lively book

which reads like a novel. It is essentially a book for

the uniniated, but moUusk enthusiasts, whose efforts

have been devoted largely to collecting shells, will

find much to broaden their views. Anyone who ap-

preciates a well-written, artistically arranged book

will want to own one.

The book is fully indexed and contains a useful,

selected bibliography, the titles being grouped under

appropriate subject headings.

William J. Clench

Curator Emenitus

Museum Comp. Zoology

Vol. 86 (2 - 4)

THE NAUTILUS

NOTES ON THE GENUS PARANCISTROLEPIS AZUMA (BUCCINIDAE)

Tadashige Habe

National Science Museum, Tokyo, 160, Japan

The genus Parancistrolepis was established by

Azuma in 1965 for Japelionf?) kinoshitai Kuroda,

1931, because of the remarkable radular features,

which superficially resemble the radula of the family

Fasciolariidae rather than that of the family Buccini-

dae in having a three-cuspidate central tooth and a

eight-to-nine-cuspidate marginal tooth.

This genus was also assigned Ancistrolepis fujitai

Kuroda, 1931, by him, only because of the shell char-

acters and the extremely small operculum which

closely resembled those of the type species. This spe-

cies differs from the type species by its thick velvety

periostracum on the surface and in being surrounded

by prominent spiral cords. Ancistrolepis hiranoi

Shikama, 1962, is an absolute synonym o{ A. fujitai

Kuroda.

The writer has observed and figures here the radu-

lae of Parancistrolepis kinoshitai (Kuroda) and

Ancistrolepis fujitai Kuroda tentatively assigned to

the genus Parancistrolepis by Azuma. These two spe-

cies have the same radular formula, suggesting they

belong to the same genus as follows:

In Parancistrolepis kinoshitai, the central tooth is

subquadrate in shape and has three small cusps at its

hind margin and the marginal tooth is large and trans-

versely broad and has eight to ten cusps.

In Ancistrolepis fujitai, the central tooth has four

small cusps and the marginal tooth 14 cusps just as

does the type species.

The radula o{ fujitai does not agree well with that

of the genus Ancistrolepis Dall, 1895, in spite of the

close resemblance of its shell features. Therefore, we

assign it io Parancistrolepis Azuma, 1965.

An examination of the anatomy o{ Ancistrolepis

grammata (Dall, 1 907) shows the radula of the genus

Ancistrolepis has a four-to-six-cuspidate central tooth

and a four-cuspidate marginal tooth. Unfortunately,

the radula oi Ancistrolepis eucosmia (Dall, 1891), the

type species of the genus Ancistrolepis, has never

been examined.

The genus Parancistrolepis is a peculiar group in

having the thin shell ornamented by prominent spiral

cords and covered by a thick periostracum, in having

an extremely small operculum, and by its marginal

tooth having many cusps. These features are far apart

from the related subfamilies Neptuneinae and Ancis-

trolepisinae. I therefore estabhsh a new subfamily

Parancistrolepisinae.

Fig. 1. Parancistrolepis fujitai (Kuroda) from off

Kushiro, Hokkaido Island, Japan,

52

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

LITERATURE CITED

Azuma, M. 1965. On the Radulae oi Ancistrolepis

kinoshitai (Kuroda) and A. trochoideus ovoideus

Habe & Ito. Jap. Jour. Malac, Venus 24(2):

127-129, text figs. 1-2.

Habe, T. & Ito, K. 1965. Shells of the World in color,

vol. 1 (The North Pacific) 176 pp., 56 pis.

Hoikusha, Osaka.

Kuroda, T. 1931. On the Japanese Species of the

Chrysodomid Whelk, Ancistrolepis and Japelon.

Venus 2(5): 221-234, text figs. 1-18.

Shikama, T. 1962. On some Noteworthy Shells from

off Chosi, Chiba Prefecture. Sci. Rep. Yokohama

Natl. Univ., (2)8: 29-86, pis. 1-3.

Fig. 2. Half row of radula of Parancistrolepis fujitai

(Kuroda) from off Kushiro, Hokkaido Island, Japan.

Fig. 3. Radula of P. kinoshitai (Kuroda) from En-

shunada, Honshu Island, Japan,

Vol. 86 (2 - 4)

THE NAUTILUS

53

BERTHELLA KANIAE, A NEW OPISTHOBRANCH FROM

THE EASTERN PACIFIC

Gale G. Sphon

Los Angeles County Museum of Natural History

Los Angeles, Calif. 90007

ABSTRACT

Berthella kaniae Sphon, a member of the opisthobranch family Pleurobranchidae, is

described as a new species from Isla Siboga, Perlas Islands, Panama (type locality) and

Punta Mita, Nayarit, Mexico. It is close to B. sideralis (Loven) and B. californica (Dall).

A new species of Berthella was collected by the

author from Punta Mita, Nayarit, Mexico, in 1960,

and through the kindness of Mrs. K. B. Meyer of the

Smithsonian Tropical Research Institute in Panama, a

second specimen was obtained from the Perlas Is-

lands, Panama. I take great pleasure in naming this

species for her.

Berthella kaniae new species

Figs. 1-8

Description - Ground color translucent, ranging

from deep yellow to almost white. Color more in-

tense on dorsum; fading along edges of mantle. Tips

of frontal veil, rolled rhinophores, and area around

genital aperture colored reddish brown. Notum, gill,

sides and frontal veil spotted with same reddish

brown color. Notum (of holotype) covered with

white powdering seen only under 10 magnification,

SheU located mid-dorsal area, wholly internal, thin,

delicate, translucent, white; haliotiform in shape.

Ventral side of shell iridescent. Umbo small, spire of

two small whorls. Sculpture of very fine irregular

growth lines radiating from umbo region and showing

through shell. Gill rachis smooth; 18 leaves on the

dorsal half. Genital opening simple without lobule; lo-

cated in front of gjll. Radula 80 rows with 100-105

teeth per half row. No rachidian tooth. Dental formu-

la, 80(100-105.0.105-100). Teeth equal in size for en-

tire half-row except for the outermost 3 or 4 which

get progressively smaller. Curve of hook of individual

lateral teeth remains constant. Mandibular plates v^th

approximately 58 rows and about 37 platelets per

row. Scales of platelets approximately equal-size ex-

cept for the outermost 4 or 5 which become pro-

gressively smaller. No denticles or barbs on the plate-

lets.

Type material - Holotype: California Academy of

Sciences Invertebrate Zoology Type collection no.

560 consists of the entire animal and an egg mass that

was laid after the specimen had been collected from

15-30 feet, Isla Siboga (type locality), Perlas Islands,

Panama, by Joyce Young on February 15, 1972. It

measured 13 X 6.5 mm. when alive and fully relaxed.

The preserved animal measures 9 X 5,5 mm. Color

transparencies of the holotype (CASIZ slide col-

lection 2741) and of the paratype (CASIZ slide col-

lection 2742) have also been deposited with the

California Academy of Sciences. Both photos were

taken of the animals in life. Paratype: Los Angeles

County Museum of Natural History, Invertebrate

Zoology type collection no. 1453 consists of the

shell, a radula slide and a sUde of the jaw plates. The

paratype was collected by the author in 2 feet of

water on the underside of a dead coral head at Punta

Mita, Nayarit, Mexico, on January 21, 1970. Color

transparencies of both the holotype and paratype

have also been deposited at the Los Angeles County

Museum of Natural History,

Discussion - There are two other species of

Berthella known to occur in the eastern Pacific:

Berthella sideralis (Loven, 1846) and Berthella cali-

fornica (Dall, 1900). The simplest, and most arfificial,

way to separate B. kaniae from these two species is

on the basis of distribution and color. Berthella

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

t »

• •

*i

**' 4»

V

.>

m:

w

'^aSi.-w"*'

V%

.'.1 - 1

8

Vol. 86(2-4)

THE NAUTILUS

'^'^

sideralis was originally described from the Norwegian

coast and later collected by Dall (this was questioned

by MacFarland. 1966) from 25 fathoms off Unalaska,

Alaska. Bcrthella califoruica ranges from Crescent

City, Del Norte County, to La JoUa, San Diego Coun-

ty, California. Berthella kaniae is a tropical-sub-

tropical species found in Nayarit, Mexico, and the

Perlas Islands, Panama. Both B. sideralis and B. cali-

fornica are white in color. Berthella kaniae has a yel-

lowish ground color and is tipped and spotted with

red dish- brown.

Illustrations by MacFarland (1966. pi. 13) show

the median lateral radular tooth o{ Berthella sideralis

to be thin, narrow and have a slight bend at the tip.

This varies along the half row to rather short, stubby,

hooked innermost teeth. The outermost teeth have

only a shght bend and no hook at the end of a long

narrow tooth. MacFarland's illustrations (from Bergh,

1904) for the radular teeth o{ Berthella californica il-

lustrate almost sickle shaped teeth. Berthella kaniae

has little variation along the entire half row of teeth,

but the teeth are sharply hooked. None of the three

species appear to have serrations on the teeth mar-

gins.

The mandibular platelets of all three species are of

comparable shape, but those of Berthella sideraUs are

serrated while both B. californica and B. kaniae are

smooth. The shells of all three species are comparable

in form.

Acknowledgments - I am extremely grateful to

Mrs. Kaniaulono B. Meyer for supplying the holotype

specimen, photo and data which made it possible to

complete the description. I also wish to thank Mr.

David K. MuUiner for permission to use his photo-

graph of the paratype.

On May 19, 1972 (after the original submitting of

this paper) a third specimen of Berthella kaniae

Sphon was found in 15 feet of water at Isla Siboga,

Islas de las Perlas, Bay of Panama. The animal was

nestled in a crevice on the underside of a small clump

of the coral Pocillopora sp. and as it was being pried

out with a knife, it autotomized a large piece of the

notal border. By the time the animal was removed, a

second piece had been cast off so that the whole

notal border (about half the entire notum) was miss-

ing from the animal. Figure 1 of the holotype clearly

shows that that specimen had also autotomized its

notal border. Although the phenomena of autotomy

appears frequently in the nudibranchs and saco-

glossans as a defensive mechanism, to my knowledge

this is the first report of it occurring in the pleuro-

branchs.

LITERATURE CITED

Bergh, L. S. R. 1904. Malacol. Unters. Semper Set. 9,

6(3): 119-181.

Dall, W. H. 1900. A new species o{ Pleurobranchus

from California. The Nautilus 14 (8): 92-93.

Loven, S. L. 1846. Index Moll. Litora Scand. Occi-

dentalis habitantium. Ofvers. Kgl. Vet. - Akad.

Forhandl. 3: 136-140.

MacFarland, F, M. 1966. Studies of opistho-

branchiate mollusks of the Pacific coast of North

America. Mem. Calif. Acad. Sci. 4: 1-546.

Figs. 1-8. Berthella kaniae Sphon, new species. Fig 1, Holotype (CASIZ 560, X7). Fig 2, Paratype (LACM

1453). Fig. 3, Shell of paratype, dorsal view (X7.5). Fig. 4, Shell of paratype, ventral view(X7.5). Fig. 5, Group

of mandibular platelets (from paratype). Fig. 6, Single mandibular platelet (from paratype). Fig. 7 , Pair of lateral

teeth (from paratype). Fig. 8, Single lateral tooth (from paratype).

56

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

A NEW SPECIES OF HUMBOLDTIANA (HELMINTHOGLYPTIDAE)

FROM THE SIERRA VIEJA MOUNTAINS OF TEXAS

E. P. Cheatum

Department of Biology, Southern Methodist University,

Dallas, Texas 75222

ABSTRACT

Humboldtiana fullingtoni Cheatum, a new pulmonate species, and the tenth species of

this genus recorded from the Texas Trans-Pecos mountainous area, is described from

5,000 ft., southern end of the Sierra Vieja Mountains, Presidio Co., about 40 miles SSW

of Valentine, Texas. It is closest in shell morphology to H. hoegeana praesidii Pilsbr)'. Its

soft parts are grayish brown, while those of other Texas species in the genus are yellow,

orange or golden. Significant distinguishing characters in the genus are discussed.

Humboldtiana fullingtoni new species

Figs. 1,2,3

Description of holotype - Shell rather thin, subglo-

bose and slightly depressed; ornamented by three,

wide dark-brown bands on the basal whorl; two bands

at the beginning of the penult whorl, the lower of

these two fading out, thus leaving only the upper

band which gradually fades out about one-half turn

from the embryonic whorls. Width of the bands on

basal whorl (measured just back of the hp): upper 4.0

mm.; middle 4.3; and lower 4.6. With the exception

of rather close-set growth lines, some of which are

marked with whitish streaks and splotches, shell

smooth and devoid of distinct granulations, although

under high power very minute granulations are visi-

ble. Inner basal portion of peristome strongly reflect-

ed over the umbilicus in the form of a slightly

rolled-over triangular plate. Narrow umbilical chink

not visible from a direct apertural view. Embryonic

whorl convex and the initial one-half embryonic

whorl smooth; following whorl marked with minute

irregularly-arranged radial and sliglitly curved striae.

Inner lip sHghtly thickened; ground color of inner

apertural wall the same wood-brown of outer shell

and broken by dark bands on the basal whorl. Shell

height: 31.2 mm.; diameter: 38.6 mm.; apertural

height: 23.6 mm.; apertural wddth: 21.2 mm.; spiral

angle: 122° (using Parodiz measuring methods,

1951); sutural angle: 2°; columellar angle: 15°; 4

whorls.

3

FIGS. I, 2 and 3, Humboldtiana fullingtoni Cheatum,

new species. Holotype, 31.2 mm. in height, 38.6 mm.

in width. Apical views of \\. ferrissiana /^//^. 4) and H.

chisosensis (fig. 5).

Vol. 86 (2 - 4)

THE NAUTILUS

57

Living animal uniform fuscous. When withdrawing

into the shell the animal discharges a considerable

quantity of a clear, frothy viscous substance. The ani-

mal when full> extended measured approximately 65

mm. in length.

Holotype: No. 2186 - 5A Dallas Museum of Natu-

ral History; paratypes M.C.Z.; USNM 706882; Univer-

sity of Michigan Museum of Zoology; Carnegie Muse-

um; and the Academy of Natural Sciences of Philadel-

phia.

COMMENTS

The new species, Humboldtiana fullingtoni, was

collected April 20, 1971, at an altitude of approxi-

mately 5,000 ft. along the rimrocks overlooking the

Rio Grande River near the southern end of Sierra

Vieja Mountains in Presidio County, Texas. The type

locality is situated about 40 miles south-southwest of

Valentine, Texas.

^0^^' "'""^M

8

FIGS. 6-9, Apical views of Humboldtiana cheatumi

(fig. 6), edithae (fig. 7), H. agavophila (fig. 8), and H.

fullingtoni /"/ig. 9).

The snail-collecting expedition was conducted

under auspices of the Dallas Museum of Natural His-

tory, Mr. Hal Kirby, Director. The first Humboldtiana

were discovered by Mr. Richard Fullington, Curator

of Invertebrates at the Museum and I am naming this

species in his honor. Most of the ten living snails and

twenty-four "bones" collected were found at depths

of one and one-half to over three feet between and

beneath rocks. The living snails were attached to

rocks and the dead shells were dug out of dirt and

humus at the above depths. The soil and humus even

at these depths were powder-dry since the last rainfall

received in that area had occurred the preceding

September. Some of the living snails had secreted as

many as four apertural epiphragms in order to pre-

vent water loss.

In size this new species conforms with four other

species - H. edithae Parodiz which was collected on

Mt. Emory in the Chisos Mountains, H. agavophila

Pratt, collected at Laguna Meadow just west of Mt.

Emory, H. cheatumi Pilsbry collected in the Davis

Mountains; and, (in the collections at the Dallas Mu-

seum of Natural History two shells) H. chisosensis

Pilsbry which measure 39 mm. in diameter. The latter

were collected by Mr. E. H, Miner in 1951, but the

precise locality in the Chisos range was not given. As

more shells of Humboldtiana are collected, undoubt-

edly the size range of the various species will be ex-

tended.

Although the color bands of H. fullingtoni show

considerable variation in width, all are dark-brown in

color. In one large, partly broken, fresh shell the in-

terspace between the middle and lower bands is whit-

ish and this wide whitish area extends around the

basal whorl. This shell, compared with other speci-

mens collected at the same place, exhibits extreme

variation in the width of color bands. Measurements

of the bands (just back of the Hp): upper, 3.7 mm.;

middle, 1.7; and lower 6.3. Such variation precludes

the use of color band width as a distinct taxonomic

feature. In all the other shells the ground color is of a

uniform light-brown with the exception of the whit-

ish streaks and splotches which are irregular in distri-

bution.

According to Pilsbry 's (1939) description of H.

hoegeana praesidii, H. fullingtoni conformed closer to

this subspecies than any other species described for

the genus. Since I had not had the opportunity of

examining the type of//, hoegeana praesidii {U. S. N,

58

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

TABLE 1. Measurements ofParatypes in mm.

Diameter Height Aperture Heiglit Apertural Width

Whorls

4

31/2

3^i

3-M

y/i

31/2

31/2

yA

4

M. 134160), I asked Dr. Harald Rehder, Senior

Zoologist, Division of Mollusks at the U. S. N. M. to

compare a paratype of H. fuUingtoni with the type

shell of//, hoegeana praesidii. After a careful exam-

ination of the two shells, Dr. Rehder made the fol-

lowing comments (Correspondence, June 29, 1971):

"Your species is indeed closer to this subspecies than

to any other Texas form . . . The apical whorls of//.

fuUingtoni do show under high power fine riblets that

are faintly granulated in places. On the third whorl

these granules increase in strength and then on the

penultimate whorl they disappear. In H. hoegeana

praesidii this granulation appears to be much more

obscure, thougli our specimens, including the type,

are rather worn and this granulation may have been

rubbed off. Your new species is furthermore larger

and darker in color, with the bands wider than in H,

hoegeana praesidii ' ',

Apical views of H. ferrissiana, H. chisosensis, H.

cheatumi, H. edithae and //. agavophila (figs. 4, 5, 6,

7 and 8 respectively) show the distinct shell granula-

tions in contrast to Fig. 9 of H. fuUingtoni in which

granulations are absent.

In living snails of the Texas Humboldtiana which

we have had the opportunity to examine, all have had

various parts of the body either a yellow, orange or

golden color. The color of the living animal of //.

fuUingtoni is a uniform fuscous or grayish brown.

When considering shell characteristics of the vari-

ous species of Texas Humboldtiana it seems that the

most reliable distinguishing characters to be used are

the presence or absence of visible granulations, shape

of the granulations, smoothness or ornamentation of

the embryonic whorls, their shape (flat or rounded),

ground and band color of the shell and shell angles.

Shell color and markings must be based upon fresh

shells. In old shells the periostracum is so frequently

eroded so as to completely obliterate granulations

and other distinctive shell markings.

Ten species of Humboldtiana have now been

described from the mountainous areas of the Texas

Trans-Pecos, and these are the following Pilsbry

(1939):

Humboldtiana chisosensis Pilsbry

Humboldtiana ferrissiana Pilsbry

Humboldtiana hoegeana praesidii Pilsbry

Humboldtiana cheatumi Pilsbry

Humboldtiana palmeri Clench and Rehder

Humboldtiana texana Pilsbry

Humboldtiana ultima Pilsbry

Pratt (1971), described Humboldtiana agavophila

from the Chisos Mts.; Parodiz (1954) described //i/m-

boldtiana edithae from Mt. Emory in the Chisos Mts.;

and this species, Humboldtiana fuUingtoni is the

tenth species to be described from the Texas

Trans-Pecos. Pilsbry (1927), had listed//, chisosensis,

H. texana and H. ultima from Texas. In his key, based

upon shell characters, the major breaks involved

granulations and whether or not the shell was

smoothish or with wrinkles along the growth lines. //.

hoegeana praesidii and // fuUingtoni appear to be the

only Texas Humboldtiana without any traces of visi-

ble granulation.

It is my opinion that the taxonomic status of the

species under the genus Humboldtiana is debatable. It

is hoped that in the not too distant future careful

morphological studies of the soft parts will help to

clarify the taxonomic "picture".

Vol. 86 (2 - 4)

THE NAUTILUS

LITERATURE CITED

Pilsbry, H. A. 1927, "The Structure and Affinities of

Humboldtiana and related Helicid Genera of

Mexico and Texas", Proc. Acad. Nat. Sci, Phila.

79: 165-192.

Pilsbry, H. A. 1939, "Land Mollusca of North

America (North of Mexico)", 1, Part 1: 395-410.

Pratt, Lloyd 1971, "Humboldtiana agavophila, A

New Helminthoglyptid Land Snail from the Chisos

Mountains, Big Bend National Park", Southwest

Naturalist 15(4): 429-435.

Parodiz, Juan J. 1951, "Methods de conquiHo-

metria", Physis, Buenos Aires, No. 58, 20: 241-248.

Parodiz, Juan J. 1954, "A New Species of Hum-

boldtiana from Texas", The Nautilus 67: 107-108.

BOOK REVIEW

AUSTRALIAN SHELLS, illustrating and describing

600 species of marine gastropods found in Austra-

lian waters. By Barry R. Wilson and Keith Gillett,

with a preface by R. Tucker Abbott. 168 pp., 106

polychrome plates, 34 monochrome figures; page

size 9 by 11 'A inches. Charles E. Tut tie Co., Rut-

land, Vermont 05701, 1971. $21.50.

Biologist Barry Wilson and photographer Keith

Gillett have combined their talents to produce a pro-

fusely illustrated and informative handbook on the

shelled gastropods one would most likely encounter

by shore collecting and diving in Australian waters.

The introductory text includes brief discussions of

the classification, nomenclature and biology of mol-

lusks in general.

A plea is made for the conservation of Australian

mollusks, before irresponsible collecting reduces read-

ily accessible shores near populated areas "... to bar-

ren biological deserts . . ." In this regard, they cite

California as an example where "... some seaboard

schools and universities have to import specimens

from other areas or take their students to places

hundreds of miles away before they are able to look

at living animals instead of only seeing pictures of

them in books," a condition, unfortunately, not

hmited to our west coast.

For each family, a brief introduction including dis-

tributional, biological, and ecological data, is given,

together with a description of each species figured on

the full-page, color plates, which are conveniently

placed facing the text. Limitations of space have per-

mitted the inclusion of only 600 species; for example

of the 23 species of abalones living in Australian

waters, 12 are illustrated in the book. The descrip-

tions include data on size, range, and abundance, to-

gether with synonymous names that have been used

for Australian forms.

This book is outstanding for the numerous color

photographs and line drawings of living mollusks.

Hopefully, the inclusion of biological and ecological

data in this book will stimulate collectors to observe

and record such information, and to be more restric-

tive in the selection of specimens for their cabinets.

The authors suggest that only a few well-preserved

specimens of a species should be taken, and they rec-

ommend that females associated with eggs should be

left undisturbed.

There are few errors of commission for an under-

taking of this magnitude. Parts of the text are some-

what out of date. Some of the nomenclature is not

current. However, in most cases, the familial groups

requiring nomenclatural changes are undergoing in-

tense study by malacologists, and specialists have pro-

posed differing classifications, as in the case of the

family Muricidae.

This beautifully illustrated book is highly recom-

mended to anyone seeking information on the marine

life inhabiting AustraUan coastal waters.

William K. Emerson

Chairman and Curator

Departmen t of L iving Invertebrates

Tlie American Museum of Natural History

60

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

Fig. J. Photograph of Tom Iredale taken in September, 1938.

Vol. 86 (2 - 4)

THE NAUTILUS

TOM IREDALE (1880-1 972)

Winston F. Ponder and Gilbert P. Whitley

The Australian Museum

6-8 College St., Sydney, Australia

The name Tom Iredale will be familiar to most zo-

ologists and certainly to all malacologists and ornith-

ologists. Iredale was a remarkable man, for in his long

career he published over 360 papers and articles and

erected more than 2,600 new taxa. He worked not

only on molluscs and birds but also published papers

on mammals and fishes. Iredale has been called the

"Australian Linnaeus" - an apt title for a man of his

talents.

Iredale was born in Cumberland, England, on 24

March, 1880, and, when 21 years of age, sailed for

health reasons to Christchurch, New Zealand. He be-

came friends with W.R.B. Oliver, with whom he visit-

ed the Kermadec Islands in 1908. While on the

Kermadec Islands he made a life-long friend of Roy

Bell who collected large quantities of material for him

on Lord Howe and Norfolk Islands, in Victoria and

southern New South Wales. While in New Zealand he

published his first three papers in the Transactions of

the New Zealand Institute ( 1 907, 1 908).

Iredale left New Zealand in 1908 to return to

England, but on the way Charles Hedley persuaded

him to stay for the Australian Association for the Ad-

vancement of Science Congress in Brisbane. Thus in

February, 1909, Iredale had a first-hand experience

with Queensland molluscs by collecting at Caloundra

and also Port Curtis.

From 1909 Iredale spent much time in the British

Museum (Nat. Hist.) as a freelance worker, identify-

ing birds and molluscs. He worked with Mathews on

Birds of Australia and wrote several papers dealing

mainly with molluscan nomenclatural and bibli-

ographic problems. One of his finest efforts was his

critical commentary on Suter's Manual of the New

Zealand Mollusca, published in 1915. He also helped

Sherborn with his great Index Animalium and col-

laborated with C. D. O'Donoghue to produce a List

of British Nudibranchiate Mollusca, which was pub-

lished in 1923. He considered liis most spectacular

molluscan discovery to be Sherbornia mirabilis.

In April, 1920, H. E. Gregory, the Director of the

Bernice Paualii Bishop Museum at Honolulu, offered

Iredale a position on the Museum Staff, but this was

not accepted.

In 1923 Iredale came to Sydney for the Pan Paci-

fic Science Congress and went on a collecting trip to

Victoria with Pilsbry, who was also in Australia for

the Congress. Pilsbry, some years later, wrote to Ire-

dale (19th July, 1928):-

"Do you still ride on those market trains, such as

that we took from Port Pliillip? I have never really

Uked pigs since that ride in the society of so many

dead ones."

"I think it wonderful of you and Basset Hull to

dedicate your Chiton (excuse me! Loricate) book to

me. Of course I think you make the genera too small,

but this is a mere matter of opinion . . ."

Iredale was to cause a lot of controversy with his

philosophy on genera and species. After his arrival in

Australia in 1923, he pubUshed a paper based on Roy

Bell's collections from Twofold Bay, southern New

South Wales, in which he erected 1 1 1 new names.

After working v^th Hedley at the Australian

Museum, he was appointed to the position of Assist-

ant to Joyce Allan in the Conchology Department in

1924. In the same year he was appointed to the posi-

tion of Conchologist and was assisted by Joyce Allan,

with Phyllis Clarke engaged as artist. Hedley left the

Australian Museum in 1924 to become the Director

of the Great Barrier Reef Committee, a position

which he occupied for two years before his death. In

Iredale's early days at the museum two other

well-known conchologists, John Brazier and Thomas

Whitelegge, who had been retrenched during the de-

62

THE NAUTILUS

November 1972

Vol. 86(2-4)

pression of 1893, also visited the museum from time

to time.

Iredale's moUuscan work at the Australian Muse-

um was prolific and ranged from chitons to cuttle-

bones, marine borers to deepwater molluscs and land

shells.

Although Iredale was basically a museum worker,

he loved field work and considered this very im-

portant to gain a proper understanding of the animals

on which he was working. He made several collecting

trips to Queensland, including participating in the

Great Barrier Reef Expedition in 1928 and 1929, and

did much local collecting in New South Wales. He

visited Lord Howe Island in December and January,

1931-2.

A bibliography and an index to his new names was

provided in 1956 by McMichael and Whitley {Aus-

tralian Zoologist, 12(3): 211-250) and one of us

(G.P.W.) is currently revising this and preparing a

more detailed account of Iredale's life and work.

Many find Iredale's work difficult because of his

brief descriptions and seemingly casual introduction

of new names. These people have probably not seen

him at work using his unsurpassed knowledge of the

Hterature, tracing back to their beginnings any species

names under consideration. He never relied on pub-

lished synonymies, everything was checked and re-

checked with a book in one hand and a specimen in

the other. Hours of observation and research were

condensed, perhaps unfortunately, into a line or two.

He never used a typewriter, all of his work being

handwritten, including the finished manuscript. A

number of unfinished manuscripts and notes are pre-

sei^ved in the Australian Museum.

Iredale's philosophy on generic groupings changed

somewhat over the years. His classifications in eariier

papers, strongly abused by some conservative con-

temporaries, are now often adopted as "modern class-

ifications." In 1912 he wrote "I am convinced that all

future workers, to produce any lasting results, must

undertake monographic studies, and moreover, must

study series, note variation, also determine the sub-

species from such series, and discriminate between

species, subspecies and varieties. I foresee the time

when there will be more genera, fewer species and

more subspecies, with entire elimination of varieties."

What could be closer to the approach of the modern

taxonoirdst?

Unfortunately for Iredale, Australia has a vast

fauna with many perplexing and exciting unknowns.

Because of this, he was rarely able to live up to his

desired standards, although, with the help of Hull, he

produced excellent monographs of the Australian and

New Zealand "loricates" which were completed in

1927 and 1932, respectively. Several important mol-

luscan reference works were produced by Iredale, in-

cluding lists of the Australian, Lord Howe and Nor-

folk Islands and Papuan land molluscs, the freshwater

molluscs of Australia and the marine molluscs of New

South Wales (with D. F. McMichael). His nomencla-

tural and bibliographic contributions result mostly

from his period spent in the British Museum and after

his retirement in 1944, he produced a number of

bibliographic and historical accounts of Australian

conchologists.

Up until the time of his death, on 12 April, 1972,

at Curl Curl, a seaside suburb of Sydney, just after his

92nd birthday, Iredale was an Honorary Associate of

the Australian Museum and except for his last four

years, frequently worked in the museum. Many

genera and species of molluscs, birds, fishes, etc., have

been named after him, the last known to him was

Favartia (Miirexiella) iredalei Ponder, 1972.

Not only was Iredale's work extraordinary, but his

sharp wit, coupled with unusual dress and a broad

grin made him a personality few could forget. Perhaps

most of all he will be remembered by those who

knew him in Australia and elsewhere for his cheerful

help and assistance which was given generously and

willingly. Certainly many people will feel a deep per-

sonal loss at Tom Iredale's passing.

Vol. 86(2-4)

THE NAUTILUS

63

SHELL STRUCTURE IN SPIRULA SPIRULA (CEPHALOPODA)'

Ronald F. Thomas & Frasier 0. Bingham

University of Miami

Rosenstiel School of Marine and Atmospheric Science

Miami, Florida 33149

ABSTRACT

The external surface of the shell of Spirula exhibits laminar mesa-like projections

suggesting deposition over an extended period. No pores, through which transfer of ma-

terials could take place, were seen in the siphuncle at a magnification of 10,000X.

Tlie small cephalopod Spirula spirula (Linnaeus,

1758) is rarely seen alive althougli its internal shell is

commonly washed onto beaches in many parts of the

world. The animal inhabits depths of 200-1500 me-

ters (Voss, 1956) and is widely distributed (Bruun,

1943). The biology, function of the shell, vertical dis-

tribution and world-wide distribution, as known, have

been covered in detail by Bruun (1943). The shell of

S. spirula appears to be thin and fragile; however it

has been found to withstand pressures between 2000

and 3500 p.s.i. without being crushed (Denton &

Gilpin-Brown, 1971).

The ability of the animal to actively change the

contents of completed chambers has been dealt with

by several authors. Chun (1914) concluded that after

a chamber was completely formed, there was no

transfer of gases or liquids into or out of that cham-

ber. Bruun (1943) thought that gases could pass

through the siphuncle or shell. Denton &

Gilpin-Brown (1971) note that some chambers of the

shell contain hquids. It is their opinion that the ani-

mal is able to regulate the gas or liquid contents of

the chamber and thus employ the shell in buoyancy

control.

The surface structure of the shell and of the

siphuncle have not previously been described. A lack

of contrast and large sculpture on the shell surface

makes it difficult to see the surface detail with a liglit

microscope. The high resolution and depth of field of

* Contribution No. 1 564 from the University of

Miami, Rosenstiel School of Marine and Atmospheric

Science, Florida 33149

the scanning electron microscope made it possible to

observe the surface detail shown in this study. Pre-

sented here are scanning electron microscope photo-

graphs of the external sculpture of the shell of S.

spirula as well as the internal structure of the

siphuncle.

In large specimens of S. spirula the shell contains

about 35 chambers. The first chamber formed by the

animal (figures 1 & 2) is small, spherical and compara-

tively smooth. This chamber probably corresponds to

the unsculptured protoconch of other molluscs in

which shell growth after settling displays adult sculp-

ture characteristics. The first chamber is somewhat

separated from the remainder of the shell by a promi-

nent constriction (figure 2). The second chamber

(figure 2) exhibits traces of surface sculpture in the

form of minute ridges scattered over the exterior sur-

face. The sculpture on the second chamber is not so

well developed as that seen on following chambers.

Figures 3 & 4 show the ridges of the seventh cham-

ber. These ridges are unconnected, mesa-like pro-

jections of the shell surface and are laminar in struc-

ture (figure 4). The ridges are tall and distinctly

layered, indicating that their formation may involve

periodic deposition througli the lifetime of the ani-

mal. Appelloff (1893) found that the thickness of the

external surface of a newly formed chamber increases

during the addition of further chambers. The in-

creased thickness seen by AppellotT (1893) is indi-

cated by this study to be a consequence of periodic

deposition of thin layers of CaCo on the tops of the

mesa-like projections of the shell surface. In more re-

cently formed chambers such as the 32nd chamber

(figures 5 & 6), the ridges are not so well-formed as

64

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

those on the seventh chamber. They are set farther

apart and arranged in a linear pattern parallel and per-

pendicular to the axis of shell growth. The laminar

structure is still present although the ridges are not as

highly elevated as in the earher chambers. It appears

to us that ridges are formed near the region of new

shell growth and later additions of CaCo increase

their size and obscure their original Unear arrange-

ment.

The external surface of the siphuncle was scanned

at magnifications up to 10,000 X. The surface ap-

peared smooth and no pores which would permit ex-

change between materials in the chambers and in the

siphuncle were noted, but this does not exclude the

possibility that such pores exist and are too small to

show up at this magnification. The inner surface of

the siphuncle is quite rough (figures 7 & 8) and the

crystalline structure of the CaCo matrix is evident.

Vol. 86 (2 - 4)

THE NAUTILUS

65

Fig. 1. The first 14 chambers of the shell ofS. spirula.

20X. Fig. 2. The first and second chambers of the

shell. lOOX. Fig 3. The seventh chamber. Note the

numerous ridges that cover the entire surface. lOOX.

Fig. 4. A portion of figure 3, at higher magnification.

Note the laminar structure of the ridge. fOOOX. Fig.

5. The smaller and linearly arranged ridges of the

32nd chamber. SOX. Fig 6. Sculpture of the 32nd

chamber. 200X. Fig. 7. Calcium carbonate crystals in

the inner surface of the siphuncle. 2,000X. Fig. 8.

Laminated calcium carbonate crystals. 5,000X.

66

THE NAUTILUS

November 1972

Vol. 86(2-4)

ACKNOWLEDGMENTS

We wish to thank Dr. Donald Marszalek and Mr.

Wally Charm for their assistance and operation of the

scanning electron microscope. This work was sup-

ported in part by National Science Foundation grant

GB-24994X.

LITERATURE CITED

Appelloff, A. 1893. Die Schalen von Sepia, Spirula

und Nautilus . Kongl. Svenska Veten-

skaps-Akademiens Handl. 25: 1-106.

Bruun, A. 1943. The biology oi Spirula spirula (L.).

Dana Report No. 24: 1-46.

Chun, C. 1914. Die Cephalopoden. II. Teil. Myopsida,

Octopoda. Wissensch. Ergebn. Deutsch.

Tiefsee-Exp. "Valdivia" 1898-1899, 18: 405-552.

Denton, E. J. & J. B. Gilpin-Brown. 1971. Further

observations on the buoyancy of Spirula. Jour.

Marine Bio. Assoc. U. K. 51: 363-373.

Voss, G. L. 1956. A review of the cephalopods of the

Gulf of Mexico. Bull Marine Science 6: 85-178.

BOOK REVIEW

EDIBLE? INCREDIBLE! By Marjorie Furlong and

Virginia Fill. Faperback, 62 pp. (24 with full

color). Ellis Robinson Publ. Co. April 1972. $2.00.

Obtainable from Mr. Fom Rice, Box 33, Port

Gamble, Wa. 98364.

This is a charming and colorful guide to the com-

mon edible invertebrates and seaweeds of the Pacific

Northwest coast of North America. Clear colored

photographs, succinct descriptions, habitats, and

cooking hints are given not only for the plausible edi-

bles, such as clams, abalones and crabs, but also for

the incredible gourmet deliglits. such as sea urchins,

barnacles, sea cucumbers and chitons. Missing is a

warning that over 200 venturesome people in North

America have suddenly died of paralytic shellfish poi-

soning resulting from eating cooked mussels, clams

and periwinkles at certain seasons in certain areas, es-

pecially in northern California and both sides of

Canada.

R. Fucker Abbott

Delaware Museum of Natural History

Vol. 86 (2 - 4)

THE NAUTILUS

THE AMPHI-ATLANTIC DISTRIBUTION OF UlTORINA MELEAGRIS

Joseph Rosewater^ and Geerat J. Vermeij^

* Division of Mollusks, Smithsonian Inst., Wash., D. C. 20560

2 Dept. Zoology, Univ. of Maryland, College Park, Md. 20742

ABSTRACT

The gastropod Littorina meleagris (Potiez and Michaud, 1838), is now known to be

the second tropical species of Littorinidae found on both sides of the Atlantic. It has

recently appeared in Bermuda, and the new West African records may be the result of a

modern ship-borne introduction.

Littorina meleagris (Potiez and Michaud, 1838) is

a common western Atlantic littorinid gastropod living

in close association with algal mats at mid to high in-

tertidal levels (Lewis, 1960; Vermeij, in press a,b) (see

figs. 3, 4). Bequaert (1943) gave the geographic range

of the species as including southern Florida, the

Bahamas, the Greater and Lesser Antilles, and the

Caribbean coasts of Honduras and Colombia. Addi-

tional locahty records in the collections of the Na-

tional Museum of Natural History include Port

Aransas, Texas; Quintana Roo, Mexico; Portete,

Costa Rica (for locality see Houbrick, 1968); Devil's

Beach, Panama Canal Zone; Curacao; La Orchila Id.

(Rehder, 1962) and Catia la Mar, Venezuela; and

Barbados. It was recorded from Bermuda by Abbott

and Jensen (1967), who believed it to have been re-

cently introduced there (see map).

Wliile examining collections in 1968 at the Zoo-

logical Museum in Copenhagen, Rosewater noted the

presence of L. meleagris among tropical West African

material collected by the Atlantide Expedition in

1946 (Takoradi, Ghana) and by the Galathea Expe-

dition in 1950 (Teshi, Ghana). The specimens were

later borrowed for study through the courtesy of Dr.

J^^rgen Knudsen. The shells were all collected in dead

condition, many containing hermit crabs, and were

mixed with specimens of the morphologically similar

L. punctata (Gmelin, 1791) (figs. 1, 2, 5, 6.).

In July, 1971, Vermeij collected living specimens

of L. meleagris at Takoradi, in the western part of

Ghana. Most specimens were found in algal turf in the

low intertidal echinoid (Echinometra lucunter) zone

in association with Tricolia sp., Siphonaria pectinata

(Linnaeus, 1758), Fissurella nubecula (Linnaeus,

1758), and Patella safiana (Lamarck, 1819). Several

additional specimens were collected in permanent,

shallow, high intertidal pools, in which L. punctata,

L. cingulifera (Dunker, 1 845), and Merit a at rat a

GmeUn, 1791, were the dominant mollusks.

No other records are presently available for L.

meleagris from West Africa, neither Nickl^s (1950)

nor Buchanan (1954) mentioning it in their surveys

of the mollusks of Senegal and Ghana respectively.

The species could not be found in collections from

the 8 other localities visited by Vermeij in Ghana

during June and July, 1971, nor were specimens col-

lected by him in Senegal or Sierre Leone. The possi-

bility therefore arises that L. meleagris is a recently

introduced species in West Africa, especially since one

of the two localities from which it is known

(Takoradi) is a shipping port. The second locality

Map 1 . Distribution of Littorina meleagris (solid dots)

and L. scabra angulifera (open circles) in the east-

em and western Atlantic.

68

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

(Teshi) lies between Accra and the new shipping port

of Tema.

It is well-known that several boreal littorinids

(Littorina littorea, L. saxatilis and L. obtusata) occur

in both the eastern and western Atlantic. Up to now,

only one tropical species, the mangrove-associated L.

scabra angulifera (Lamarck, 1822), (see figs. 7, 8) was

known to be amphi-Atlantic in distribution (Rose-

water, 1970, pp. 455-458). The very restricted

known range of L. meleagris in West Africa and the

strictly tropical distribution of the species on both

sides of the Atlantic contrasts strongly with the signi-

ficantly greater latitudinal range of L. s. angulifera in

both the eastern and western Atlantic (see map, also

Bequaert, 1943; Nickles, 1950). Briggs (1967) and

Scheltema (1971) have noted that many amphi-Atlan-

tic fishes and mollusks, like the two amphi-Atlantic

tropical httorines, have a more restricted and often

more patchy distribution in the eastern Atlantic as

compared to the western Atlantic. This tendency may

be related partly to the much narrower latitudinal ex-

tent of tropical water (Ekman, 1953) and more vari-

able sea temperature regime in West Africa; as well as

to the pattern of east-flowing surface ocean currents

(Scheltema, 1971). In this connection, Lewis (1960)

has pointed out that L. meleagris produces floating

egg capsules which hatch into swimming veligers.

While L. meleagris was not included in a geo-

graphic distribution analysis of the rocky intertidal

mollusks at Takoradi (Vermeij, 1972), the habitat

characteristics of the species agree well with those of

other amphi-Atlantic species at that locality. In gener-

al, species not restricted to the eastern Atlantic were

found either to live cryptically beneath stones or in

other locally sheltered habitats, or to be eurytopic

with respect to shore level (Vermeij, 1972, in press,

a).

Differences between populations of L. meleagris in

the eastern and western Atlantic appear to be very lit-

tle greater than between adjacent western Atlantic

populations. Large numbers of specimens were not

available for analysis, especially from West Africa;

however, the following measurements are given (see

Table 1).

LITERATURE CITED

Abbott, R. T. and R. Jensen. 1967. Molluscan Faunal

Changes Around Bermuda. Science 155 (3763);

687-688.

Bequaert, J. C. 1943. The Genus Littorina in the

Western Atlantic. Johnsonia 1 (7): 1-27.

Briggs, J. C, 1967. Dispersal of Tropical Marine Shore

Animals: Coriolis Parameters or Competition? Na-

ture 216 (51 13): 350.

Buchanan, J. B. 1954. Marine Molluscs of the Gola

Coast, West Africa. Journal of the West African

Science Association 1 (1): 30-45.

Ekman, S. 1953. Zoogeography of the Sea. Sidgwick

and Jackson, Ltd., London, xiv + 417 pp.

Houbrick, J. R. 1968. A Survey of the Littoral Ma-

rine Mollusks of the Caribbean Coast of Costa

Rica. The Veliger 11 (1): 4-23.

Lewis, J. B. 1960. The Fauna of Rocky Shores of

Barbados, West Indies. Canadian Journal of Zoolo-

gy 38: 391-435.

Nickles, M. 1950. Mollusques Testaces Marins de la

C6te Occidentale d'Afrique. Manuels

Ouest-Africans 2 Paul Lechevalier, Editor, Paris, X

+ 269 pp.

Rehder, H. A. 1962. Contribucion al Conocimiento

de los Moluscos Marinos del Archipelago de los

Roques y La Orchila. Memoria de la Sociedad de

Ciencias Naturales La Salle 22 (62): 1 16-138.

TABLE L Comparative measurements of Caribbean and Takoradi, Ghana, colonies of Littorina meleagris.

Length of Aperture

.57

.61

.65

.64

Vol. 86 (2 - 4)

THE NAUTJLUS

Rosewater, J. 1970. The Family Littorinidae in the

Indo Pacific. Part I. The Subfamily Littorininae.

IndoPacific Mollusca 2(11): 41 7-506.

Scheltema, R. S. 1971. Larval Dispersal as a Means of

Genetic Exchange Between Geographically Sepa-

rated Populations of Shallow-water Benthic Marine

Gastropods. Biological Bulletin 140: 284-322.

Vermeij, G. J. 1972. Endemism and Environment:

Some Shore Molluscs of the Tropical Atlantic. The

American Naturalist 106(947): 89-101.

Vermeij, G. J. In Press, a. West Indian Molluscan

Communities in the Rocky Intertidal Zone: A

Morphological Approach. In Smith, R. A. (Ed.),

Thomas F. Goreau Memorial Volume. University

of Miami Press.

Vermeij, G. J. In Press, b. Morphological Patterns in

High Intertidal Gastropods: An Essay on Adaptive

Strategies and Their Limitations. Marine Biology.

Fig. 1-4, Littorina meleagris. 1, 2 from Takoradi,

Ghana, USNM 707154, 6.4 mm. length. 3, 4 from

English Harbor, Antigua, USNM 423627b, 6.8 mm.

Fig. 5, 6, Littorina punctata, from Cotonou, Daho-

mey, USNM 707160, 10.5 mm. Fig 7, 8, Littorina

scabra angulifera, from Monrovia, Liberia, USNM

673999, 22.6 mm

70

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

TERATOLOGICAL LITTORINA SCABRA ANGULIFERA

Joseph Rosewater

Division of MoUusks

Smithsonian Institution

Washington, D. C. 20560

The occurrence of monstrosities in Littorinidae is

well-known, especially in the boreal species, Littorina

littorea (Linne), L. obtusata (Linne), and L. saxatilis

(Olivi) (Jeffreys, 1865, pp. 354-376; 1869, pi. 65; S.

V. Wood, 1848, pp. 118, 119, pi. 10, fig. 14, a-k;

Reeve, 1857, pi. 4, fig. 18a; Dautzenberg and Fischer,

1914, pi. 3; Bequaert, 1943, pi. 3, fig. 7.) Anomalies

of the shell apparently result either from injuries or

genetic aberrations and show up as sculptural defects,

changes in direction of coiling, or other obvious mal-

formations. Readily visible anomalies of the animal

consist of branching or other abnormalities of the

tentacles and penis, and many others have been noted

(Pelseneer, 1920). It seems probable that the con-

siderable frequency with which teratological Littorina

littorea was reported in Great Britain reflects the fact

that hundreds of tons of this species were collected

for food and that "pennywinkle" mongers were re-

sponsive to conchologists who paid well for unusual

specimens. A number of such specimens from Bil-

lingsgate Market, England, are present in the Jeffreys

Collection in the National Museum of Natural Histo-

ry-

Variation in shell morphology often is extremely

prevalent in some species of tropical Littorinidae (see

Rosewater, 1970, pi. 352). Anomalous individuals sel-

dom have been reported, however, and it is not

known whether this is because they occur with less

frequency in the tropics or simply have not been dis-

covered. An extremely turreted individual of Lit-

torina scabra angulifera (Lamarck, 1822) (figs. 1, 2)

was collected from Island SQ 1, near Squirrel Key,

Monroe County, Florida, by D. Simberloff, on June

19, 1969. A large population of the species was pre-

sent on Red Mangrove. Although the malformation is

extreme, when the specimen was first seen it was easi-

ly recognized as L. scabra angulifera. An inquiry was

made of the collector, through Dr. W. Heard, Florida

State University, Tallahassee, Florida, to determine

whether or not any other specimens exhibiting this

phenotype had been observed. The figured specimen

was the only one seen. A subsequent examination of

the collection of this species in the National Museum

of Natural History yielded one additional, smaller.

similarly malformed specimen, also from Florida, but

with no definite locality (figs. 3, 4). A perusal of the

reported and figured teratological specimens of Lit-

torinidae shows the turreted anomaly to be rather

common. The Squirrel Key specimen may be the re-

sult of a genetic defect since the turreted condition is

present in the earliest post-nuclear whorls of the shell,

the protoconch being missing. The "Florida" spec-

imen may have resulted from an injury because the

first 3-4 whorls appear normal and the turreted condi-

tion develops following an apparent disruption of

growth during the fourth revolution.

Figs. 1-6. Turreted and Normal Linoxm^ ■i.cd^ixdL'dngxx-

liferaf Lamarck, 1822).

Figs. 1, 2. Squirrel Key, Monroe Co., Florida (USNM

700046; 34.6x20.4 mm.).

Figs. 3, 4. ''Florida" (USNM 594251; 17.4 x 11.7

mm, ).

Figs. 5, 6. Normal specimen from mouth of Shark

River, Ponce de Leon Bay, southwest Florida ( USNM

129174; 36.9 X 20.1 mm,).

Vol. 86(2-4)

THE NAUTILUS

LITERATURE CITED

Bequaert, J. C. 1943. The Genus Littorina in the

Western Atlantic Ocean. Johnsonia 1(7): 1-27, 7

pis.

Dautzenberg, Ph. and H. Fischer. 1914. Etude sur le

Littorina obtusata et ses variations. Journal de

Conchyliologie 62: 87-128, 3 pis.

Jeffreys, J. G. 1865. British Conchology 3: 1-393-

1869./Zj/d 5, pL65.

BOOK REVIEW

THE GEOLOGY AND PALEONTOLOGY OF THE

MARINE PLIOCENE OF SAN DIEGO, CALI-

FORNIA (PALEONTOLOGY: PELECYPODA).

By Leo George Hertlein and U. S. Grant, IV. San

Diego Society of Natural History Memoir 2 (Part

2B), pp. 135-411, text figs. 7-13, pis. 27-57, paper-

back. July 21, 1972. $15.00 postpaid. Available

by mail from the San Diego Society of Natural

History, P. O. Box 1390, Balboa Park, San Diego,

California 92 11 2.

This quarto volume is literally a mine of informa-

tion on many families of marine pelecypods. The

taxonomic discussion includes 4 subclasses, 8 orders,

22 superfamilies, 39 famihes, 82 genera, and 144 spe-

cies and subspecies. Ten species are not positively

identified but are compared to known species, and

two additional ones are cited only to genus. Eleven

species, not seen by Hertlein and Grant but reported

by other authors, are included. The species with

small-sized shells are not neglected. One genus, one

subgenus, nine species, and four subspecies are

described as new. The new genus is Irusella (Vener-

idae), and the new subgenus is Axinola (Glycymerid-

idae). The new species and subspecies are: Nucula

(Ennucula) balboana, Mytilus (Crenomytilus) coaling-

ensis stembergi, Chlamys (Chlamys) hastata ellisi,

Chlamys (Argopecten) abietis abbotti, Lima (Limaria)

orcutti, Aligena diegoana, Bomia frankiana, Dosinia

ponderosa diegoana, Chione allisoni, Chione kanakof-

fi, Psephidia stephensae, Semele ashleyi, Thracia kana-

koffi.

The authors have made many taxonomic keys to

genera, subgenera, species, and subspecies. Four of

the seven text figures are drawings of pectinid genera

that show the exterior ornamentation of the valves.

Most of the photographs of the species are of good

quahty, and there are 348 of them.

Pelseneer, Paul. 1920. Les variations et Leur Heredite

chez les Mollusques. Brussels, pp. 1-826.

Reeve, L. A. 1857-1858. Conchologia Iconica 10, Lit-

torina, pis. 1-18.

Rosewater, J. 1970. The Family Littorinidae in the

Indo-Pacific, Part I. The subfamily Littorininae.

Indo-Pacific Mollusca, 2(1 1): 417-509, pis.

325-386.

Wood, Searles V. 1848. A Monograph of the Crag

Mollusca, Part I, Univalves, pp. xii -i- 1-208, 21 pis.

Most of the categories above the genus are like

those found in the Treatise on Invertebrate Pale-

ontology, and Hertlein and Grant cite the references

and refer to the first usages and spellings of these

higher categories. The type species of each genus and

subgenus is given with mode of designation. There is

additional information given on the genera and sub-

genera, such as geologic range, a brief description, and

sometimes geographic range and other species that

have been allocated to the genus besides the ones

found in the San Diego Formation. The descriptions

of the species are detailed and include the synonymy,

repository of type specimens, type locaHty, geologic

range, the original description, morphologic variabil-

ity, and a discussion of related species. The taxono-

mic descriptions comprise 200 pages.

References and supplementary data, which are

numbered consecutively within the text in parenthe-

ses, are listed after the descriptive taxonomic part of

the monograph. This portion of the volume comprises

37 pages, and there are 1,384 entries. It is in this sec-

tion that one finds information on the names of the

categories above the genus, and the type species of

many genera and subgenera that are related to those

described in the taxonomic discussion. Some addi-

tional information is given on ecology, geologic

ranges, and biology and geographic distribution of

genera, subgenera, and species.

Four pages are devoted to a list of collecting locali-

ties. The 23-page index is quite complete.

The most impressive thing about this monograph is

the incredible amount of scientific hterature that

Hertlein and Grant examined and cited. This work

will be useful to anyone doing research on marine

Pelecypoda, and it contains much good raw data for

the paleoecologist to analyze.

David Nicol

Box 14376, University Station

Gainesville, Florida 32601

72

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

TYPE LOCALITY OF UNIO PUMILVS LEA, 1838 (UNIONIDAE)

Samuel L. H. Fuller

Department of Limnology

Academy of Natural Sciences of Philadelphia

Philadelphia, Pa. 19103

ABSTRACT

A method for verifying the synonymies of Fusconaia masoni (Conrad 1834} and Lex-

ingtonia subplana (Conrad 1837) is discussed. The type locality of Unio pumilus Lea

1838 is corrected from the Neuse River system to the Cape Fear River system (both in

North Carolina). U. pumilus is a synonym of F. masoni, newly recorded from the Cape

Fear River.

The occurrence of a member of the primitive

unionid genus Fusconaia Simpson 1900 in the

Savannah River system was recorded by Fuller

(1971), who identified the species as maso^/ Conrad

1834, originally described as a Unio from the same

river system. Insofar as U. masoni appears to be the

earliest available name for this species, there is no

problem in accepting the combination F. masoni

(Conrad, 1834). This species should not be confused

with the conchologjcally very similar U. subplanus

Conrad 1837, the type species of the genus Lexing-

tonia Ortmann 1914. The distinctiveness oi subplana

has been verified on anatomical ground. Furthermore,

it is apparently limited to the upper James River (Ort-

mann, 1914).

It will be impossible to construct the correct

synonymies of Fusconaia masoni and Lexingtonia

subplana until their ranges are fully understood: the

Fig. 1. Known distribution of Fusconaia masoni (Conrad 1834) in the Cape Fear River

system of North Carolina, with the type locality of Vnio pumilus Lea 1838. Dotted lines

represent routes between Smithfield and Fayetteville available to Isaac Lea in 1827.

Vol. 86 (2 - 4)

THE NAUTILUS

type specimens of the several relevant taxa are devoid

of beak sculpture and soft parts, which are the only

discriminants between these two genera (Ortmann,

1914). Those taxa which were described from areas

where only one of these species proves to occur can

justifiably be assigned to synonymy; some of those

from areas where the two species prove sympatric

may have to be considered nomina dubia. In order to

apply such criteria, one must know the type localities

in question.

One of these taxa is Unio pumilus Lea, 1838,

whose type locality was cited by Johnson (1970:

301) as in the Neuse River system of North Carolina,

whereas the adjacent Cape Fear River system is cor-

rect. The point is of some importance because I have

Fusconaia masoni from the Cape Fear River: thus

Unio pumilus falls within that species' range.

In describing Unio pumilus, Isaac Lea (1838: 23)

wrote: "A single individual of this species was found

by me in 1827, in crossing the Black River, on the

road to Fayetteville from Smithfield." Contemporary

maps of North Carolina show that in 1808 there was

a single road connecting these towns and that a sec-

ond had been constructed by 1833 (Cumming, 1966:

pis. 9-10). The earher road crossed the Black River at

a point in the modern Harnett County about five

miles northeast of Erwin and about four miles

east-southeast of Coats (Fig. 1). The crossing of the

later route was about three miles southeast of Erwin,

some 1 0 river miles downstream of the original cross-

ing.

These points cannot be fixed with precision be-

cause modern roads do not necessarily follow their

original courses and because of the difficulty in

reconcihng early and modern cartography. Nonethe-

less, the type locality of Unio pumilus definitely is

somewhere between these two points as approximate-

ly defined. It is doubtless at or near the upstream

point because the later, more circuitous route (Fig. 1)

was less likely to be used in public transportation; be-

cause Cumming's (1966) post- 1833 maps do not

show this road (which suggests its relative unimport-

ance), and because Lea mentioned only "the" road

from Smithfield to Fayetteville.

On both early and modern maps is shown a Black

Creek in the Neuse River system, which flows sub-

parallel to Black River immediately to the southwest

of Smithfield. Lea had to cross both streams on his

1827 journey (Fig. 1). Their nominal similarity mis-

led Johnson (1970: 301), who thus amended the type

locality of Unio pumilus: "Black River [a tributary of

the Neuse River] on the road to Fayetteville from

Smithfield [=about 10 mi. W of Benson, Johnston

Co.] , North Carolina." The type locality is (about

five niiles) west of Benson, but this point is now in

Harnett County, and Black River is in the Cape Fear

River system.

This somewhat confusing situation is complicated

by the existence in this general area of another Black

River, also a tributary of the Cape Fear. This stream

is created by the confluence of Six Run and Great

Coharie Creeks, whose headwaters rise a few miles

south of Smithfield, but can be traversed on only the

most devious route to Fayetteville (Fig. 1). Moreover,

even Cumming's (1966) earlier maps clearly distin-

guish between this Black River and the streams "Six

Runs" and "Great Cohera" or "Big Cohary." Thus no

stream in this drainage could have been nominally or

geographically confused by Lea with the Black River

he crossed in 1827.

In view of all these considerations, it seems best to

restrict the type locality of Unio pumilus to: Black

River [Cape Fear River system: South River drain-

age] , on the road to Fayetteville from Smithfield

[=about 5 mi. NE Erv^n and about 4 mi. ESE Coats,

Harnett Co.] , North Carolina.

Thanks are due J. B. Post (Free Library, Philadel-

phia) and W. F. Caddell, Jr. (North Carolina State

Highway Commission) for aid in locating early and

modern maps of portions of North Carolina and to R.

T. Abbott for a critical reading of the manuscript.

LITERATURE CITED

Cumming, W. P. 1966. North Carolina in Maps. State

Department of Archives and History, Raleigli. Pp.

viii+ 1-36, pis. 1-15.

Fuller, S. L. H. 1971. A Brief Field Guide to the

Fresh-Water Mussels (Mollusca: Bivalvia: Union-

acea) of the Savannah River System. ASB Bulletin

18(4): 137-146, text figs. 1-14, 1 pL

Johnson, R. I. 1970. The Systematics and Zoogeo-

graphy of the Unionidae (Mollusca: Bivalvia) of

the Southern Atlantic Slope Region. Bulletin of

the Museum of Comparative Zoology 140(6):

263-450, tables 1-4, text figs. 1-5, pis. 1-22.

Lea, I. 1838. Descriptions of New Freshwater and

Land Shells. Transactions of the American Philo

sophical Society (N. S.), 6(1): 1-154, pis. 1-24.

Ortmann, A. E. 1914. Studies in Naiades [Part 3.].

The Nautilus 28 (3): 28-34.

74

THE NAUTILUS

November 1972

Vol. 86(2-4)

UNIO CAROLINIANA BOSC, 1801 (UNIONIDAE)

Samuel L. H. Fuller

Department of Limnology

Academy of Natural Sciences of Philadelphia

Philadelphia, Pennsylvania 19103

ABSTRACT

The date of authority for Unio caroliniana Bosc is established as 1801. Conflicting

interpretations of this taxon 's identity are reviewed. U, caroliniana should be considered a

nomen dubium.

Bosc (1801: 142, pi. 23, fig. 2) described and

figured Unio caroliniana in Volume 3 of the first edi-

tion of his Histoire Naturelle des Coquilles. This

volume is dated "An X," an abbreviation for the

tenth year of the French revolutionary (or republi-

can )calendar - i.e., the period from 23 September

1801 to 23 September 1802. The unsatisfactory

designation "1801/02" would have to be used as the

date of authority for Unio caroliniana had Dodge

(1952: 3>?>) not shown that Volume 3 of the

"Histoire" appeared no later than 23 October 1801.

Bosc (ibid.) compared his description and figure of

Unio caroliniana to figure 5 on plate 239 in the

so-called Encyclopedic Methodique of Lamarck and

collaborators. The figures in question resemble the

shell of a North American fresh-water mussel. Unio-

merus tetralasmus (Say 1831) (concept of Johnson,

1970: 339). Noticing the unclearness of Bosc's figure

(whose posterior hinge is particularly obscure). Lea

( 1852: 44) felt that it might as easily have been in-

tended {() represent the holarctic Margaritifera mar-

garitifera (Linnaeus 1758) (Margaritifcridae). This is

certainly possible because the nature of the lateral

tcclh m this figure might be considered to represent

those which may occur in younger A/, margaritifera

from Europe (Woodward, 1854) and North America

(Ortmann, 191<)).

On (he other hand, the type locality ofUnio caro-

liniana is "Ics caux dormanfcs (stagnant | en Caroline"

(Bosc, ibid.), whereas the range oi Margari I ij era mar-

garitifera HI the Atlantic drainage of the United States

extends no farther south than the upper Little

Schuylkill River drainage in northeastern Pennsyl-

vania (Ortmann, 1919: 3-5), and this species' depen-

dence upon pristine and flowing waters is

well-known. Moreover, the figures of Bosc and in the

Encyclopedic Methodique might be thouglit to

demonstrate the rougli periostracum, conspicuous

growth rests, postdorsal angulation, and somewhat

arcuate ventral margin of Uniomerus from larger

Carolinian rivers. Finally, Bosc (ibid.) redescribed M.

margaritifera (as Unio margaritifera) on the same

page where Unio caroliniana was described; one as-

sumes that he distinguished clearly between them.

For these reasons, one is tempted to grant Unio

caroliniana Bosc 1801 priority over Unio tetralasmus

Say 1831. Thus a perplexed Simpson (1900: 741,

1914: 711) did with a query include Unio caroliniana.

in his synonymy of Unio (Uniomerus) obesus Lea

1831. However, the figure of Unio caroliniana miglit

also illustrate Elliptio complanata (Liglitfoot 1 786), a

widespread species in the Atlantic drainage (see John-

son, 1970: 321), whose ready confusion with Unio-

merus is notorious.

Unfortunately, there seems to be no extant vouch-

er material which might help settle the matter. For

instance, E. Fischer (in correspondence) stated that a

type specimen of Unio caroliniana has not been locat-

ed in the National Museum of Natural History at

Paris, where the specimens figured by Bosc or in the

Encyclopedic Methodique are as likely to have been

deposited as at another institution, if any.

An ambiguous figure, an inadequate description

("Ovale, alongee; les sommets ronges. "), and the

probable loss of the figured type - these factors com-

bine to disguise the correct identity of Unio carotin-

Vol. 86(2-4)

THE NAUTILUS

75

iaim, which had best be considered a nonieii dubiuni.

In recent discussions of Uniomenis from the Atlantic

and Gulf drainages of the United States, Clench and

Turner (1956- 177-179) and Johnson (1970:

339-343) understandably ignored the name.

Confusion over the status of Unio caroliniana is

reflected in the curious evolution of its spelling and

citation in the literature:

Unio caroliniana [error for carolinianus] . Bosc,

1801: 142, pi. 23, fig. 2; 1824: 139, pi. 23, fig.

2; 1830: 139, pi. 34, fig. 2.

Margaron (Margaritana) Carolinianus Bosc [1801] ,

Lea. 1852: 44.

Unio carolinensis Bosc 1824, Simpson, 1900: 741;

1914: 711.

Fortunately, no one appears to have confused this

taxon with Unio caroliensis Pacome 1889, a syno-

nym of the European Unio crassus batavus Maton and

Rackett 1807 (Haas, 1969: 51)

Thanks are due R. T. Abbott for criticism of the

manuscript and for guidance among unfamiliar ele-

ments of the late 18th and early 19th century mol-

luscan literature.

LITERATURE CITED

Bosc, L. A. G. 1801. Histoire Naturelle des Coquilles

[First Edition], Volume 3: 1-292, plates. Deter-

ville (Paris).

Bosc, L. A. G. 1824. Histoire Naturelle des Coquilles

[Second Edition], Volume 3: 1-287, plates.

Verdiere (Paris).

Bosc, L. A„ G. 1830. Histoire Naturelle des Coquilles

[Third Edition], Volume 3: 1-287, plates. Raynal

(Paris).

Clench, W. J., and R. D. Turner. 1956. Freshwater

Mollusks of Alabama, Georgia, and Florida from

the Escambia to the Suwanee River. Bulletin of

the Florida State Museum 1(3): 97-239, pis. 1-9.

Dodge, H. 1952. A Historical Review of the Mollusks

of Linnaeus. Part 1. The Classes Loricata and Pele-

cypoda. Bulletin of the American Museum of

Natural History 100( 1): 1-264.

Haas, F. 1969. Superfamilia Unionacea. Das Tier-

reich, Lief. 88: x + 1-663.

Johnson, R. I. 1970. The Systematics and Zoogeo-

graphy of the Unionidae (MoUusca: Bivalvia) of

the Southern Atlantic Slope Region. Bulletin of

the Museum of Comparative Zoology 140(6):

263-450, tables 1-4, text figs. 105, pis. 1-22.

Lea, I. 1852. A Synopsis of the Family of Naiades

[Third Edition]. Blanchard and Lea (Phila-

delphia). Pp. xx -h 17-88.

Ortmann, A. E. 1919. A Monograph of the Naiades of

Pennsylvania. Part 3. Systemafic Account of the

Genera and Species. Memoirs of the Carnegie

Museum 8(1): 1-384, text figs. 1-34, pis. 1-21.

Simpson. C. T. 1900. Synopsis of the Naiades, or

Pearly Fresh-Water Mussels. Proceedings of the

United States National Museum 22(1205): viii -h

501-1044.pl. 18.

Simpson, C. T. 1914. A Descriptive Catalogue of the

Naiades, or Pearly Fresh-Water Mussels. Bryant

Walker (Detroit). Pp. xii + 1-1 540.

Woodward, S. P. 1 854. A Manual of the Mollusca,

Part 2: 159-330, text figs. 90-223, pis. 14-24. John

Weale (London).

76

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

GREATER ADAPTABILITY OF FRESHWATER MUSSELS

TO NATURAL RATHER THAN TO ARTIFICIAL DISPLACEMENT

Marc J. Imlay

Office of Endangered Species

Bureau of Sport Fisheries and Wildlife

Fish and Wildlife Service

Washington, D. C. 20240

ABSTRACT

Artificial displacement of freshwater mussels in the stream bottom (substrate

smothering, commercial harvesting, or removal and replacement ) was detrimental to mus-

sels more often than was natural displacement (natural sand formation, storm disturb-

ance, etc.). A biological explanation is presented that contributes to understanding why

water projects (dredging, channelization, impoundments, quarry washing) are so devas-

tating to freshwater mussels.

Freshwater mussels are artificially displaced in

their natural stream bottom environment in many

ways, including simple removal from the substrate,

being smothered by a substrate, and being disturbed

in their substrate during harvesting of a mussel bed.

The mussel may have the physical capacity to recover

from these events by digging back in or out, as the

case may be, re-orienting to the direction of current

flow, and regaining a desirable water depth and type

of substrate by migratory movement. But there is

some evidence that the mussel is more prone to ac-

complish these feats when the displacement is of

natural, rather than artificial, origin. Animal behavior-

ists are learning that organisms utilize their abilities to

surmount difficulties when certain response-triggering

mechanisms, much more probably associated with

naturally occurring problems, are present. To site one

example (Cloudsley-Thompson, 1961: 118), in reset-

ting daily activity patterns to a changing photo-peri-

od, the animal responds in accordance with its own

daily rhytiim of resetting sensitivity. Even bright light

or total darkness has no effect at other times of the

day.

SMOTIIl RING

Freshwater mussels presumably can extricate

themselves from a thin covering of sand or silt by em-

ploying the foot and valves (Barnes, 1955, 1962). It is

conceivable, however, that an unnatural covering oc-

curring at an atypical season wt)uld not trigger a

natural response mechanism for digging out, even

thougli the mussel has the physical means to do so.

The results of the following laboratory experiment on

mussel response to smothering as a form of artificial

displacement indicate that the latter possibility oc-

curs more often in practice. Mussels were placed on

their sides in large battery jars, covered with sand,

silt, or other substrates and provided with about one

foot of continuously aerated Lake Superior water at

16 C.

Detritus. - On July 6, 1967, nine A nodonta grandis

collected from a very slow-moving creek in one foot

of water were covered with three inches of detritus

consisting of leaf, twig, and other loose material from

the creek. A few mussels emerged each day until by

July 1 1 all the mussels had completely emerged.

However, sixteen Fusconaia flava were covered on

July 1 8 and in seven days two Fusconaia flava were

almost emerged, one half way up, but thirteen were

on the bottom (ten dead).

On July 14, eigiit Ligumia recta were covered 'Adth

the loose detritus and in four days three were at least

partly emerged but at least three were on the bottom.

Sand. - On July 14, seventeen Fusconaia Jlava were

covered with 7.5 inches of sand from the St. Croix

River. On July 18, nine A nodonta grandis were also

covered v^th the river sand. None of the Fusconaia

Jlava had emerged in four days. By seven days all the

Anodonta grandis were still at the bottom with six

dead.

However, muddy sand was collected from Upper

St. Croix Lake and the leaves and twigs sieved out.

This lake sand was much muckier than the river sand.

On July 14, nine Anodonta grandis were covered with

Vol. 86 (2 - 4)

THE NAUTILUS

77

the sand. By four days, eight Anodonta grandis were

emerged. On July 18, nine Ligiunia recta were cov-

ered by the lake sand. By seven days one was

emerged, three were two-thirds the way up (two

dead), and five were on the bottom (four dead).

Very fine, mucky sand, mixed with thick clay, was

obtained from St. Louis Bay in Duluth and 4.5 inches

of it placed July 19 on top of ten Lasmigona costata

and ten Lampsilis siliquoidea. By six days one

Lampsilis emerged in normal feeding position. With

four others the siphons broke surface and were active.

Another Lampsilis was almost emerged. One

Lampsilis was on the bottom but alive. Two others

were dead. Of the Lasmigona, nine were on the bot-

tom, seven alive and two dead. On July 25, ten

Lampsilis siliquoidea from Grand Lake, Minnesota

and eight Anodonta grandis were covered, along with

five Lasmigona costata and four Lampsilis of river ori-

gin. By six days two Anodonta grandis and two lake

Lampsilis siliquoidea were emerged, along with two

river Lampsilis, but five Anodonta grandis were on

the bottom alive and eiglit lake Lampsilis siliquoidea

were on the bottom (seven dead). The remaining two

river Lampsilis were both dead on the bottom along

with the five Lasmigona costata (one dead).

Silt. - On August 3, sand was obtained 50-100 feet

from the dam at Fish Lake outlet, Minnesota, and silt

300-400 feet from the dam. On August 5, under 7

inches of the silt, were placed twelve Anodonta

grandis and twelve under a duplicate silt test,

fourteen under 7 inches of the sand and thirteen

under a duplicate sand test. On August 14, in sand,

one was emerged and another was dead one inch

down. On August 24, no more had emerged in all

four tests. But in one sand test all eleven clams were

dead on the bottom and in the other sand test four-

teen were on the bottom dead. In one silt test, all

twelve were on the bottom dead and in the other silt

test ten were found on the bottom dead.

Grit. - Grit (material too coarse to be considered

sand) was obtained from the Spht Rock area of Lake

Superior and, on July 19, placed on top of thirteen

Fusconaia flava, four Amblema costata and

twenty-one Lampsilis siliquoidea.

On July 25, one Fusconaia flava and Lampsilis

siliquoidea were fully emerged, two Fusconaia flava

and one Lampsilis siliquoidea half emerged, four

Lampsilis with siphons emerged, one Amblema costa-

ta wath siphons emerged. Two Lampsilis and one Fus-

conaia flava had climbed four-fifths up the grit.

But, on the bottom there were nine Fusconaia

flava, three Amblema costata and ten Lampsilis

siliquoidea of which three were dead.

On July 25, sixteen Fusconaia flava and ten

Lampsilis siliquoidea from Grand Lake, Minnesota,

were covered with grit. By August 7, four Fusconaia

flava had emerged. By August 24, no more clams had

emerged, but about one inch from the bottom were

two dead Grand Lake Lampsilis. Two Fusconaia flava

were ahve, but on the bottom. The other ten Fus-

conaia flava and eight Grand Lake Lampsilis were

dead on the bottom.

Application. - The demonstrated failure of mussels

more often than not to climb out of smothering con-

ditions explains in part the devastating effect of

dredging (Van der Schalie, 1941: 308), channeUza-

tion, silt behind dams, and quarry washing (Stans-

bery, 1970), or for that matter, of dust storms and

surface run-off (Ellis, 1936).

Even Anodonta grandis was not immune to

smothering, yet Dennis (1971) notes "As the com-

mon name 'floater' impUes, this species is Hglit weight

with a thin shell, enabling it to survive in loose sand,

mud, and silt."

As an example of the difference to mussels be-

tween artificial and natural sand formation, Grier

(1922) may be quoted: "An old bed of 'niggerheads'

existing at Wild's Landing was found to be absolutely

covered with sand deflected by the dams . . . the best

collecting in this area was from the sand bars."

Scruggs (1960: 28) observed in impounded areas of

the Tennessee River recently plagued by 0.5-1.25

inches of silt cover that "a relatively large number of

dead shells, of different species and sizes, were found

under layers of silt one inch or deeper at several sta-

fions. These shells were embedded in a normal posi-

tion with the valves closed; the shells showed litfle

sign of erosion indicating that mortality had occurred

within recent months."

METHODS OF HARVEST

Grier (1926) considered the practice of harvesting

mussels with the shoulder rake a principal reason for

the decline of the clamming industry. Van der Schalie

(1941) has commented that there has been "an ex-

cessive amount of mussel-shell gathering with the use

of apparatus injurious to the mussel beds." He urges

(1948) enforcement against the illegal practices of

commercial hand-picking and forking because only

the acceptable method (with the crow-foot bar skim-

78

THE NAUTILUS

November 1972

Vol. 86(2-4)

ming above the surface and removing just the larger,

liiglier mussels) does not remove the young ones so

necessary in turn for replacement of the older stock.

Furthermore, "some of the larger specimens" im-

portant for reproduction are "down at least five or

six inches." ". . . completely beneath the surface of

the river bottom It is seriously doubted if the

crow-foot dredges used in commercial operations

would bring up specimens from such depth in the

mud and gravel" (Van Cleave, 1940).

Dennis (1971) explains the damage to mussel beds

by hand-picking or raking as a matter of removing too

many shells for restoring to be possible. I suggest that

an additional explanation may be that brailing either

removes a mussel entirely from the water or does not

disturb it at all, leaving the substrate below relatively

undisturbed. Forking and hand-picking, on the other

hand, with the kicking of the substrate, overturning

of rocks, raking the substrate, and throwing back

mussels too young for harvest may amount to artifi-

cial displacement of the mussels not collected.

REMOVAL FROM SUBSTRATE

Isely (I9I4) examined 164 "Quadrulae" in Okla-

homa for migration a year after artificial displace-

ment into Shootly Creek from the Chikaskia River.

He recovered 84.8 percent of the original tagged mus-

sels and showed little, if any, niigration. Yet these

very test mussels had been actively moving naturally

before collection as shown by track marks. Similarly,

46 percent of Anodonta gmndis were found in their

original location, isely did not report on whether the

mussels had burrowed in or not, let alone their orient-

ation relative to the direction of stream flow. Isely

placed some Quadrulae in one foot of water and they

migrated to deeper water.

On May 27, 1971 1 placed twenty large male

/.ampsilis ventricosa on their sides at one location in

three feet of water near the bank in the Eau Claire

River, Wisconsin. On July 1 1, nineteen of these mus-

sels were found in the same four by four foot place-

ment area (thus no migration). All wlmc burrowed in

except one mussel almost (m its side. All were orient-

ed posteriorly upstream ami fourteen essentially

parallel to the current flow.

Hi)vvcver, earlier, in the late autumn, I left about

fifty mussels (including Lampsilis vcntricosa) on their

sides in a four by four foot sandy area near the bank

of the I:au Claire River and three months later the

mussels were still on their sides except for two or

three which had burrowed in. There was no migration

from the placement area. Thus, artificial displacement

at an atypical season was too extreme for the animals

to adapt to it.

Stansbery (personal communication) also observed

that mussels collected in summer and left on their

sides in a pile for many weeks remained th^t way.

It may be speculated that certain species of mus-

sels establish a partial domicile by growing into a

shape which conforms to the particular microhabitat

pressures and requirements at the location of the

particular mussel. This follows the familiar experience

of finding arcuate and even odd-shaped specimens in

areas v^th boulders, stones, and small patches of suit-

able sandy substrate. Such an older mussel would be

less adaptable to displacement than a younger mussel.

It would appear that regulations for commercial clam-

mers to toss back undersized mussels or mussels that

are protected as endangered species may accomplish

little because the mussels in question have been artifi-

cially displaced and are not expected to necessarily

regain a viable posture in the substrate Bates (1971:

32) reported that hand-picked specimens fared better

in regards to simple mortality after three weeks of re-

placement in the streams, then did brail-collected

specimens but even they had eleven percent mortality

(450 specimens in test) in just three weeks.

An experiment showing exceptions to the rule

may also be described. On August 21, ten Lampsilis

siliquoidea from Grand Lake, Minnesota, were placed

on top of a substrate in a bucket in the laboratory.

This was their natural substrate collected from where

the clams were collected, just inside the perimeter of

the emergent vegetation zone in the lake. Running

water (at about 16 C) was made to flow in copious

quantity past the animals and by the next day eiglit

o\' the clams were in a normal feeding position. The

substrate felt hard, yet the clams had pushed their

way in.

Similarly, Trueman (1968) placed Margaritifera

margaritifcra in an aquarium containing sand from

the river they were collected from. The mussels gener-

ally burrowed in normal position after a day or so of

migrating around the aquarium.

On the other hand. Roscoe and Redelings (1964)

transplanted twenty-five specimens of Margaritifera

margaritifcra and some lay almost parallel to the bot-

tom, covered to much extent by stones. "Most of the

clams in the bed had the siphons directed upstream,

but among the transplants the clams assumed every

Vol. 86 (2 - 4)

THE NAUTILUS

79

angle from siphons directed toward the current to

siphons directed away from the current " Yet, in the

same river, these investigators observed whole beds

had been washed downstream by natural storms into

two new areas.

Tliat mussels are displaced naturally and adapt

well is attested to by the following field experiences:

A fruitful collection over a one-hundred yard stretch

on the Eau Claire River, Wisconsin, yielded several

species in large numbers but these species had been

collected out from that location two years earlier. It

is not known whether the repopulation came from

downstream or upstream, but it evidently represents a

natural displacement from somewhere else in the

strean^ These displaced mussels were not laying on

their sides or misoriented to the current, but ap-

peared to be in normal position in the substrate.

Furthermore, a similar experience (collecting out, re-

population, adaption to displacement) was encoun-

tered at another location on the Eau Claire River.

The type of distinction shown here between artifi-

cial and natural imposition on the mussels may be in-

volved in the unhappy circumstance that artificial

river-lakes such as the large impoundments on the

Tennessee River and elsewhere are so inimical to mus-

sels but natural river-lakes such as Lake Pepin, Minne-

sota, or Mussel Shoals, Alabama, supported even

more species and numbers of mussels than have large

rivers.

LITERATURE CITED

Barnes, G. E. 1955. The behavior of Anodonta

cygiiea L. and its neurophysiological basis. Jour.

Exp. Biol. 32:158-174.

Barnes, G. E. 1962. The behavior of unrestrained

Anodonta. Animal Behavior 10:174-176.

Bates, J. M. 1971. Ohio Mussel Fisheries Invesfiga-

tion, Final Report, Part 1, Center for Aquaric Bi-

ology, Department of Biology, Eastern Michigan

University, Ypsilanti, Michigan 48197.

Cloudsley-Thompson, J. L. 1961. Rhythmic activity

in animal physiology and behaviour. Academic

Press, New York. 236 p.

Dennis, S. D. 1971. Pennsylvania Mussel Studies.

Final Report, Center for Aquaric Biology, Eastern

Michigan University, Department of Biology,

Ypsilanri, Michigan 48197. 138 p.

Ellis, M. M. 1936. Erosion silt as a factor in aquatic

environments. Ecology 17:29-42.

Grier, N. M. 1922, Final report on the study and ap-

praisal of mussel resources in selected areas of the

Upper Mississippi River. Amer. Midland Naturalist

8:1-33.

Grier, N. M. 1926. Report on the study and appraisal

of mussel resources in selected areas of the Upper

Mississippi River, 1920-25. Amer. Midland Natu-

ralist 10:89-110.

Isely, F. B. 1914. Experimental study of the growth

and migration of freshwater mussels. Bureau of

Fisheries, Document 792. 27 p.

Roscoe, E. J., and S. Redelings. 1964. The ecology of

the freshwater pearl mussel Margaritifera margariti-

fera(L.). Sterkiana 16:19-32.

Scruggs, G, D. 1960. Status of freshwater mussel

stocks in the Tennessee River special scientific re-

port - Fisheries No. 370. U. S. Fish and Wildlife

Service, Washington, D.C. 41 p.

Stansbery, D. H. 1970. Rare and endangered fresh-

water mollusks of the Mississippi and St. Lawrence

River systems of North America. Malacologia

10:9-22.

Stansbery, D. H. 1971. Personal communication. Mu-

seum of Zoology, Ohio State University, Colum-

bus, Ohio.

Trueman, E. R. 1968. The locomotion of the fresh-

water chm Margaritifera margaritifera (Unionacea:

margaritanidae). Malacologia 6:401-410.

Van Cleave, H. J. 1 940. Ten years of observation on a

freshwater mussel population. Ecology

21:363-370.

Van der Schalie, H. 1941. Zoogeography of naiades in

the Grand and Muskegon Rivers of Michigan as re-

lated to glacial history. Papers of the Michigan

Academy of Science, Arts & Letters 26:297-310.

Van der Schalie, H. 1948. The commercially valuable

mussels of the Grand River in Michigan. Misc.

publ. 4 of Institute for Fisheries Research. Michi-

gan Department of Conservarion. 42 p.

80

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

A NEW SPECIES OY PERSICULA FROM

THE ARABIAN SEA (MARGINELLIDAE)

Barry Roth

1217 Waller Street, San Francisco, California 94 11 7

and Richard E. Petit

P. O. Box 133, N. Myrtle Beach, South Carolina 29582

Persicula masirana Roth and Petit, new species

Description of holotype: Shell of medium size for

the genus, solid, ovate, narrower anteriorly; surface

polished, unsculptured except for faint growth lines.

Color mottled yellowish tan with a broad, reddish

brown peripheral band and eleven narrow, spiral rows

of alternating white and brown spots. The first row of

spots placed immediately below the suture, the rows

on either margin of the peripheral band larger than

the rest, and the four rows on the band itself being

very small and fine. A narrow spiral row of white

spots occurs just anterior to the brown band, and

another just posterior to it. Spire not visible, im-

mersed in a pad of brown callus. Outer lip white,

arching above spire, terminating posteriorly in a wash

of white enamel which partly covers apical callus pad;

lip margin rounded, slightly thickened externally, its

anterior half having about 12 faint, oblong denticles a

short distance inside aperture. Aperture narrow,

widening anteriorly, white inside, with a curved sul-

cus at posterior end and a deep, oblique anterior ca-

nal. Parietal wall convex, covered with translucent,

white callus which thickens to a node in front of the

posterior sulcus. Columella with eight folds, including

one at base of columella; the two most anterior folds

strong, extending onto body whorl; remaining folds

much smaller, confined to aperture, becoming in-

creasingly faint posteriorly. Length 10.0 mm.; great-

est diameter 7.2 mm.

Holotype, no. 51327, Delaware Museum of Natur-

al History; type locality: Ras Jidufah (local name:

Monument Beach) at northeast tip of Masirah Island

(21°N, 59°E), in the Arabian Sea, off the coast of

the Sultanate of Muscat and Oman; collected by Sgt.

Eric Thompson, Royal Air Force, early February,

1971. The holotype was found in beach drift. Al-

though we have been able to examine only the holo-

type, notes from the collector indicate that it was not

the only specimen collected.

Discussion: With its pattern of spiral rows of spots

and single peripheral band, Persicula masirana differs

from all other species of the genus. It is most nearly

like Persicula catenata (Montagu, 1803) from Florida

and the Caribbean. The latter species averages 6 mm.

long; it has a tan shell, spiral rows of white sagittate

spots which are margined with brown lines on the

side nearest the outer lip. and three brown spiral

bands, each crossed by a single row of V-shaped dash-

es. Less similar are the Caribbean P. sagittata (Hinds,

1 844) and eastern Pacific P. phrygia (Sowerby,

1846), which lack brown banding entirely.

Among Indian Ocean species, Persicula pulchella

(Kiener, 1834), reported from Australia to the Per-

sian Gulf, has waved axial brown Hues on a white

ground.

The genus Persicula is an old one, with forms

much like some Recent species first appearing in the

Eocene. Persicula masirana and the similarly-pat-

terned Caribbean species may be isolated survivors of

a Tethyan stock which has undergone little modifica-

tion since the early Tertiary.

LITERATURE CITED

Hinds, R. B. 1844. Description of Marginellae collect-

ed during the voyage of H. M.S. Sulphur, and from

the collection of Mr. Cuming. Proc, Zool. Soc.

London 14: 72-77.

Kiener, L, C. 1 834. Species general et iconographie

des coquilles vivantes . . .: genre marginelle. Paris

(J.-B. Bailliere et fils). 30 p.

Montagu, G. 1803. Testacea Britanica or natural his-

tory of British shells, marine, land and fresh-water,

including the most minute: systematically ar-

ranged and embellished with figures. Romsey (J. S.

Hollis). xxxvii + 606 p.

Sowerby, G. B. 1846. Monograph of the genus Mor-

ginella. In: Thesaurus conchyliorum, or figures and

descripfions of shells. 1: 373-406,

Figs. 1, 2. Persicula masirana Roth & Petit, new spe-

cies. Dorsal and ventral views of holotype, length 10

mm

Vol. 86 (2 - 4)

THE NAUTILUS

QUATERNARY MOLLUSKS FROM FIJP

Horace G. Richards

Academy of Natural Sciences of Philadelphia

Philadelphia, Pa. 19103

In August. 1969, while visiting Viti Levu in the

Fiji Islands, I had the opportunity of maidng a small

collection of fossil mollusks in the Lautoka area. The

exact locality is:

Along a road which branches off the Wairambetia

Road, which branches off to the SSW from the

Natambua Road in northwest Viti Levu. This

locality is close to, or identical with, localities C

1 133 and RB 32 of the Fiji Geological Survey De-

partment and is about 0.5 mile NW of Pila Trig.

The elevation is slightly over 200 feet above sea

level.

The mollusks studied include specimens collected

by myself and my student-assistant, Richard Nelson

in 1969, augmented by material collected by Mr. Nel-

son in 1970, material supplied by Peter Rodda and

other members of the Fiji Geological Survey Depart-

ment, and specimens borrowed from the British Mu-

seum through the kindness of Dr. Patrick Nuttall.

Apparently two geological ages are represented by

the material studied. Specimens from sliglitly higher

elevations than the road are apparently Late Tertiary,

and are assigned to the Mba Group (Rodda, 1967),

following mapping by W. R. Dickinson. These spec-

imens have been transferred to Dr. Harry Ladd, of the

U. S. National Museum in Washington, D. C, who is

studying the Tertiary mollusks of Fiji. (Most of the

specimens loaned by the British Museum are appar-

ently of Tertiary age and v^l be studied by Dr.

Ladd.)

The material obtained from close to the road is ap-

parently of Quaternary age, since all species are

known to be living in the Fiji waters today. The fauna

is of special interest because it apparently marks the

highest elevation for Quaternary fossils on the island.

However this is not surprising in view of the tectonic

history of the Fiji Islands. Schofield (1971) records

high sea levels (Pleistocene ?) on the Lau Islands, up

^ Part of this study was made possible by a grant from

the Penrose Fund of the American Philosophical So-

ciety.

to elevation 770 feet on Tuvutha, and to 500 and 600

feet on other islands of the group. For data on the

geology of Fiji, see Ladd (1934), Bartholomew

(1959), Rodda (1967) and Schofield (1970). Th.

fauna here recorded is apparently considerably oldei

than the material mentioned by Schofield.

In addition to thanking Mr. Rodda, Mr. Nelson

and Dr. Nuttall for help in gathering this fauna, 1

want to express grafitude to Dr. Harry Ladd, of the

U. S. National Museum and Dr. R. Tucker Abbott, of

the Delaware Natural History Museum, and to an-

other student-assistant. Earl Shapiro, who assisted in

the study of the fossils at the Academy. Dr. John

Wells, of Cornell University, identified corals listed

below.

The following fossils have been identified from the

Quaternary portion of the locality:

Gastropoda: Trochus (Trochus) maculatus Linne,

Tectus (Rochia) niloticus (Linne), Turbo sp. (opercu-

lum), Nerita sp., Rhinoclavis (Rhinoclavis) aspera

(Linne), Clypeomorus sp., Epitonium sp., Cypraea

errones Gmelin, Cypraea sp., Naticarius marochiemis

(Gmelin), Polinices mammilla (Linne), Mitrella ligula

(Duclos) and Conus (Asprella) sulcatus Hwass.

Pelecypoda: Anadara scapha (Gmelin), Lithophaga

sp., Villorita sp., Chama sp., Gafarium dispar (Dill-

wyn), Tridacna maxima (Roding), Hippopus hip-

popus (Linne) and Spondylus ducalis Roding.

Corals (idenfified by Dr. Wells): Acropora sp.,

Stylophora pisstillata (Esper), Platygyra lamellina

(Ehrenberg) forma rustica (Dana), and Acanthocy-

athus grayi Milne Edwards and Haime. The first four

are reefs corals, while the last mentioned suggests a

water depth of at least 50 meters. In this connecfion,

it should be mentioned that H. Digliton Thomas

identified the coral Desmophyllum cf. crista-galli

Milne Edwards and Haime from this locality (32)

which "would suggest a Pleistocene rather than an

earher age for the deposit yielding it." (Bartholomew,

1959 p. 19).

LITERATURE CITED

Bartholomew, R. W. 1959. Geology of the Lautoka

Area, North-West Viti Levu. Geological Survey

Dept., Suva, Fiji, Bulletin 2, 25 pp.

Ladd, Harry S. and others. 1934. Geology of Vitilevu,

Fiji. Bull. Bishop Museum, Honolulu, no. 119:

1-263.

Rodda, Peter. 1967. Outline of the Geology of Viti

82

THE NAUTILUS

November 1972

Vol. 86(2-4)

Levu. New Zealand Jour. Geology and Geophysics

10(5): 1259-1273.

Schofield, J. C. 1970. Notes on Late Quaternary Sea

Levels, Fiji and Rarotonga. New Zealand Jour.

Geology and Geophysics 13(1): 199-206.

Schofield, J. C. 1971. Note on High Sea- Level Evi-

dence from Lau Islands, South-West Pacific. New

Z el and Jour. Geology and Geophysics 14(1):

240-241.

ON THE ALLEGED OCCURRENCE OF

LAMBIS VIOLACEA AND CLITHON

CORONATA IN INDONESIA

Henk K. Mienis

Dept. Zool., Hebrew Univ.. Jerusalem

In his prehminary account of the genus Lambis in

Indonesia, Butot (1955: 79, pi. 2, fig. 6) mentions

Lambis (Millepes) violacea (Swainson, 1821) for the

first and only time from an Indonesian locality:

Ambat, on the east coast of Madura. Quite under-

standably Abbott (1961: 167) was rather puzzled by

this record, as authentic material of this species is

only known from the islands in the western part of

the Indian Ocean. Yet he mentions Butot's locality in

his list of records, but omits it, however, from the

map, which shows the distribution oi Lambis violacea

(loc. cit: p. 168 and pi. 130).

The freshwater gastropod Clithon (Clithon) coro-

nata Leach, 1815, better known by its junior syno-

nym Clithon longispina (Recluz, 1841), is a fairly

common species of Mauritius, Reunion, Rodriguez

and Madagascar, all islands in the same part of the In-

dian Ocean. Van Bcnthem Jutting (1956: 283, fig.

12) mentions, however, Clithon coronata in her crit-

ical revision of the Javanese freshwater gastropods

from a locality in Indonesia: West Java, Muara

Tjibuni, Sukabumi. This record is reason for

Starmiihlner (1969: 56) to consider C coronata an

Indomalaysian stray into the freshwater fauna of

Madagascar.

During our current revision of the family Neritidae

in which we examined more than 120,000 specimens,

of which about a quarter had Indomalaysian locali-

ties, we never found a second specimen of C corona-

ta from a locality in Indonesia.

The similarity in both cases is therefore striking.

The link between both cases is however still much

closer. Butot's record of Lambis violacea from

Madura and van Benthem Jut ting's record oi Clithon

coronata from Java are both based on material in the

collection Ouwens, stored in the Zoological Museum

of Bogor, Indonesia. According to van Benthem Jut-

ting (1956: 283) it is a well-kriown fact that the late

Major P. A. Ouwens (1850-1922) was somewhat care-

less as to localities and labels. The possibility that

both records are based on wrongly labelled material is

thus very great. We propose, therefore, to leave both

records out of any further account until they have

been confirmed by more recently collected material.

On this basis Lambis violacea and Clithon coronata

have to be considered as species with a rather restrict-

ed range in the western part of the Indian Ocean,

with the possible center of their distribution being

Mauritius and the Cargados Carojos Shoals.

LITERATURE CITED

Abbott, R. Tucker, 1961. The genus Lambis in the

Indo-Pacific. Indo-Pac. Moll., 1(3): 147-174.

Benthem Jutting, W.S.S. van, 1956. Systematic

studies on the non-marine Mollusca of the

Indo-Australian Archipelago, V. Critical revision of

the Javanese Freshwater Gastropods. Treubia

23(2): 259-477.

Butot, L. J. M., 1955. Duivelsklauwen, boksbeugels,

schorpioenen of enterhaken. Penggemar Alam 35:

71-84, 3 pis.

Starmiihlner, F., 1969. Die Gastropoden der Madagas-

sischen Binnengewasser. Malacologia 8(1-2):

1-434.

Vol. 86 (2 - 4)

THE NAUTILUS

83

4NCISTR0LEPIS K A n AMURAI A NEW JAPANESE BUCCINIDAE

Tadashige Habe & Kiyoshi Ito

National Science Museum. Tokyo 160, Japan

We have had a good opportunity to obtain spec-

imens of a new Ancistroleph collected by the whelk

fishing boats of Mr. Shosan Mizutani at the Abashi-

ri-Yamatotai, off Abashiri (about 500 meters deep)

on the northeast coast of Hokkaido, through the

courtesy of Mr. Ryosuke Kawamura. a leading shell

collector in Japan.

These specimens closely resemble A. grammata

(Dalk 1907) and its varietal forms. A. g. hikitai

Kuroda, 1944, and A. g. yamazakii Kuroda, 1944, in

general features, but they have elongated shells with

rather weak spiral cords, except on the shoulder and

the base. Also, Ancistrolepis (Clinopegma) clamon

Dall, which seems to be a subspecies of Ancistrolepis

(ClinopegfJia) unica (Pilsbry, 1905), is a related spe-

cies in having an elongated shell, but has only one

proniinent shoulder cord down from the suture with

an obliquely-placed convex interval between them, in-

stead of the horizontally-placed, flat interval.

Therefore, they are new to science, and we are

pleased to name this interesting species in the honor

of Mr. Ryosuke Kawamura.

Ancistrolepis kawamurai new species

Figs. 1-3

Shell large, elongated and tabulated; rather thick

but fragile; 7-whorled white; covered with a

dark-brown periostracum, marked with finely, axially

lamellose growth Hnes and sculptured with the spiral

cords. Wliorl with proniinent T-rail-shaped, spiral

cords, two at the shoulder and 5 or 6 at the base of

the body whorl. Broad peripheral area between the

shoulder and the base smooth to spirally sculptured

in various degrees of strength. Space between the su-

ture and the shoulder spiral cord is canaliculated hori-

zontally and has a weak spiral cord on its bottom in

some specimens. Aperture roundly ovate, white with-

in. Outer margin gently curved but shouldered and

crenulated by the endings of spiral cords on the sur-

face. Canal rather wide, sinuated backward. Columel-

lar margin white, weakly curved and twisted at the

lower end.

Operculum rather thick, corneus, dark-brown,

ovate in shape. Nucleus situated at the narrowed low-

er end. Radula stenoglossate. Central tooth trans-

versely broad and has three cusps at the hind margin,

the middle one smaller than the other two. Marginal

tooth also has three cusps, the outer one the largest

and the middle one smaller than the inner one.

Heiglit 89.5 mm,. Breadth 42.4 mm. (Type spec-

imen presei"ved in the National Science Museum,

NSMT-Mo 41826) (fig. 1); height 87.4 mm., breadth

40.0 mm. (paratype specimen preserved in the

National Science Museum, NSMT-Mo 41827) (fig. 2);

heiglit 97.5 mm., breadth 48.2 mm. (paratype spec-

imen preserved in the Kawamura's collection); height

77.8 mm., breadth 41,6 mm., (paratype specimen

preserved in the Kawamura's collection); height 90,0

mm., breadth 40.0 mm, (the specimen examined for

its radula).

Type locality: Abashiri-Yamato-Tai (about 500 m.

deep), off Abashiri, Hokkaido in the Sea of Okliotsk.

Distribution: Besides the type locality, off the east

coast of Saghahen (46° 40'N., 143°58'E. 690 meters

deep) Collected by Mr. Kinji Takagawa.

Figs. 1, 2. Ancistrolepis kawamurai Habc & Ito, new

species. Fig. 1. Holotype, height 89.^^ nun.: Fig. 2.

Paratype, height 87.4 mm

84

THE NAUTILUS

November 1972

Vol. 86 (2 - 4)

LITERATURE CITED

Dall, W. H. 1925. Illustrations of unfigured types of

shells in the collection of the United States

National Museum. Proc. U. S. Natl. Mus. 66(17):

1-41, pis. 1-36.

Habe, T. & K. Ito 1965. Shells of the World in color,

vol. 1 (The Northern Pacific) 176 pp., 56 pis.

Habe, T. & K. Ito 1968. Buccinid species from Rausu,

Hokkaido. Venus, Jap. Jour. Malac. 27(1): 1-8, pi.

1.

Kuroda, T. 1931. On the Japanese species of the

chrysodonid whelks, Ancistrolepis and Japelion,

Venus 2(5): 221-234, text figs. 1-18.

Kuroda, T. 1944. New Shells from Japan (5), Venus

13(5-8): 237-239, text figs. 1-2.

Shikama, T. 1957. On some gastropoda from Nemuro

Strait, Eastern Hokkaido, Sci. Rep. Yokohama

Natl. Univ., (2)6: 31-36, pis. 10.

Fig. 3. Half a transverse row of the radula of Ancis-

trolepis kawamurai Habe & Ito, new species.

Vol. 86 (2-4)

THE NAUTILUS

85

AN UNDESCRIBED STRUCTURAL FEATURE IN THE MARSUPIUM

OF ELLIPTIO LANCEOLATA (LEA 1828) (UNIONIDAE)

Samuel L. H. Fuller

Department of Limnology

Academy of Natural Sciences of Philadelphia

Philadelpliia, Pa. 19103

ABSTRACT

Some populations of EUiptio lanceolata (Lea 1828) from the Atlantic drainage of the

United States bear a longitudinal rib on the posterior surface of each of a number of

interlamellar septa near the middle of the marsupial (outer female) demibranch. A dis-

cussion of the taxonomic implications of this discovery concludes with the rejection ofE.

arctata (Conrad 1834) as a possible name for such populations. The septal ribs of E.

lanceolata are contrasted to the structurally and evolutionarily very different ones found

in Uniomerus. Possibly the ribs in E. lanceolata developed as a source of additional sup-

port for the marsupium.

Samples oi EUiptio lanceolata (Lea 1828) from the

Cape Fear River, North Carolina, exhibit a curious

structural feature of the marsupial water tubes: a

more or less conspicuous rib runs medially along the

interlamellar septum for most of the height of the

(outer) demibranch. Although plainly visible from the

anterior side, the rib intrudes into the lumen of the

water tube from only the posterior surface of the sep-

tum (why the reverse should not be the case is ob-

scure). The rib is more common and strongly de-

veloped on septa which lie at or near the longitudinal

midpoint of the demibranch - i.e., in the region of its

greatest height. The rib interferes with formation of

the egg mass, which exhibits a corresponding, more or

less straight, shallow groove on its anterior surface.

These EUiptio are referrable to Unio perlatus Lea

1863, whose type locality is "Cape Fear River, Black

Rock Landing [=2 mi. S Kings Bluff, Bladen Co.] ,

North Carolina" (Johnson, 1970: 332). Most of my

material is from points upstream in Bladen and Cum-

berland Counties. Johnson (ibid.) referred this taxon

to EUiptio arctata (Conrad 1 834), originally described

from the Alabama River system in the Gulf drainage

and, in the Atlantic drainage, "known only from the

several type lots [of its synonyms] , and from a rather

large unlocalized series from the Cape Fear River,

North Carohna." An unusually thin and laterally

compressed shell with a somewhat arcuate ventral

margin is common to all this material - and does re-

semble the shell of Gulf drainage E. arctata - but

these characters frequently occur in E. lanceolata.

particularly in the southern one half or so of its

range.

That Johnson's (ibid.) concept of Atlantic drain-

age EUiptio arctata depends upon scattered specimens

without unique conchological characters suggests that

there is little to be gained by separating this material

from E. lanceolata. The septal ribs of Cape Fear River

lanceolata might be grounds for distinguishing them

taxonomically from other EUiptio had I not observed

(a less developed version of) this phenomenon in

conchologjcally ordinary lanceolata from the Patux-

ent River system on the Western Shore of Maryland

near the northern limit of this species' range. (It must

be emphasized that this point of view does not pre-

clude the possibility that true arctata occurs some-

where in the Atlantic drainage.)

Otherwise than septal ribs, Cape Fear River EUip-

tio lanceolata do not differ anatomically from the

descriptive notes of Ortmann (1912), Reardon

(1929), or Fuller (1971). Therefore, I am inclined to

view these ribs as nothing more than another (and oc-

casional) difference between this species and the

more common and widespread E. complanata (Light-

foot 1786), from whose ancestors the lanceo-

lata-type was probably an offshoot.

The single, longitudinal groove on the anterior

surface of the egg mass in some EUiptio lanceolata is

reminiscent of the numerous, transverse grooves on

both surfaces of the egg mass in Uniomerus. In the

latter genus, egg mass sculpture reflects imperfect sep-

tal fusion (Fuller, 1971), whereas the septum is a per-

86

THE NAUTILUS

November 1972

Vol. 86 (2-4)

fectly fused, surficially smooth structure in EUiptio.

The septal ribs of lanceolata, then, appear to be an

adaptation which has little or nothing to do with the

degree of development of the septum. Perhaps their

role is to provide the marsupium with additional sup-

port against its collapse and the destruction of its

contents. Perhaps these ribs developed during a pro-

longed arid period, when dessication was a threat to

life and reproduction.

Thanks are due G. M. Davis for a critical reading of

the manuscript.

LITERATURE CITED

Fuller. S. L. H. 1971. A Brief Field Guide to the

Fresh-Water Mussels (Mollusca: Bivalvia: Union-

acea) of the Savannah River System. ASB Bulletin

18(4): 137-146. text figures 1-14, 1 plate.

Johnson, R. I. 1970. The Systematics and Zoogeo-

graphy of the Unionidae (Mollusca: Bivalvia) of

the Southern Atlantic Slope Region. Bulletin of

the Museum of Comparative Zoology 140(6):

263-450, tables 1-4, text figures 1-5, plates 1-22.

Ortmann, A. E. 1912. Notes upon the Families and

Genera of the Naiades. Annals of the Carnegie

Museum 8(2): 222-365. text figures 1-28, plates

18-20.

Reardon, L. 1929. A Contribution to Our Knowledge

of the Anatomy of the Fresh-Water Mussels of the

District of Columbia. Proceedings of the United

States National Museum 75 (2782): 1-12, plates

1-5.

Vol. 86 (2-4)

THE NAUTILUS

87

AN UNUSUAL FORM OF LITTORINA UrWREA LINNE

FOUND IN NOVA SCOTIA

Derek S. Davis

Nova Scotia Museum

1 747 Summer Street

Halifax. Nova Scotia, Canada

During a survey ot^ tlie intertidal fauna of

Petpeswick Inlet, Halifax County, Nova Scotia, sam-

ples of Littorina littorea (Linne) were taken. On June

28, 1971, a rather unusual form was collected along

with normal L. littorea in a Spartina marsh at Petpes-

wick Harbour. This specimen and an example of the

normal form are shown in Figures 1 and 2. The popu-

lation density of L. littorea in this habitat was 1 20

per square metre, but only one specimen of the un-

usual form was found. It has a high spire, deeply in-

cised sutures and six whorls. Its shell length is 16.1

mm. and shell breadth 1 2.4 mm. The soft parts were

identical to those of normal forms. This specimen was

a female and did not contain trematode parasites.

The normal form (Fig. 2) was selected at random

from the specimens collected. Its shell length is 18.4

mm. and shell breadth 14.2 mm. There are six whorls.

This specimen was a male, parasitised by the digenetic

trematode, Cr}>ptocotyle lingua (Creplin).

Since marked variations in the form of L. littorea

are rather uncommon, the occurrence of this unusual

specimen is of interest. Jeffreys (1865) briefly de-

scribes four varieties and some other abnormalities in

L. littorea. Among these, var. turrita is described as

"spire turreted, the whorls being divided by a deep

and channelled suture". Jeffreys also mentions a

monstrosity with the spire much elongated. Without a

full description for reference it is difficult to know

whether the unusual form from Petpeswick is the

named variety or a mutant.

Read (1965 and 1966) has described shell-less

specimens of L. littorea from Maine, and suggested

that these may be mutants or victims of disease. The

unusual form from Petpeswick was examined for

larval trematode parasites, which sometimes cause

shell deformations in prosobranchs (Fretter and

Graham, 1962), but none were found.

Bequaert (1943) in his description of L. littorea

from the western Atlantic, states that the species

shows little variation on the coast of North America.

It may be that the varieties do occur more frequently

among the offspring, but that their survival would de-

pend upon the habitat in which they settled. The high

spire and small aperture would be a disadvantage on a

shore exposed to wave action, but not so in a salt

marsh. The sui-vival of the shell-less forms in the

eel-grass pond habitat has been attributed by Read

(loc. cit) to the low possibility of predation. L. lit-

torea are more commonly observed and sampled on

rocky shores and not in salt marshes, and so it is pos-

sible that unusual varieties have remained largely un-

detected.

The Petpeswick specimens described are now in

the collection of the Nova Scotia Museum under ac-

cession number 1 971 -Z- 180.

LITERATURE CITED

Bequaert, J. C. 1943. The genus Littorina in the west-

ern Atlantic. Johnsonia 1(7): 1-27.

Fretter, V. and A. Graliam 1962. British prosobranch

molluscs. London, Ray Society, 755 pp.

Jeffreys, J. G. 1865. British conchology, 3: 1-393.

London, van Voorst.

Read, K. R. H. 1965. Littorina littorea L., without

shell. Proc. Malac. Soc. Lond. 36: 307.

Read, K. R. H. 1966. Littorina littorea L., without

shells; a further note. Proc. Malac. Soc. London

37: 127.

» «3l^

1

Figs. 1 and 2. Littorina littorea from a salt marsh,

Petpeswick Harbour, Nova Scotia. L An unusual, tur-

reted form, length 16.1 mm. 2. A normal form,

length 18.4 mm (Photos by R. E. Merrick).

88

THE NAUTILUS

Vol. 86 (2-4)

NEWS

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of Malacologjsts will be held at the Asilomar Confer-

ence Grounds in Pacific Grove, California, July 11-14,

1973, and will feature contributed papers, symposia,

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Inquiries may be sent to the Secretary, Mrs. Edith

Abbott, 1264 W. Cienega Ave., San Dimas, CA

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Executive Board Members for the year are: Presi-

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Nybakken; Secretary Mrs. Edith Abbott; Treasurer

Mr. Glenn Burghardt; Members-at- Large Dr. George

Radwin and Dr. George Davis; the three most recent

Past Presidents are Dr. A. Myra Keen, Dr. Eugene V.

Coan and Mrs. Beatrice L. Burch.

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WILMINGTON SHELL CLUB

The Wilmington Shell Club was founded on Octo-

ber 2, 1972. It meets the first Monday of the month,

October througli May, at 8:00 p.m. in the auditorium

of the Delaware Museum of Natural History. Officers

elected: President, R. Tucker Abbott; Vice-President.

C. John Finlay; Secretary-Treasurer. Sophie (Mrs.

Eldon) Homsey; Archivist, Russell H. Jensen; Coun-

cillors: Mrs. Edmond du Pont and Paul Leach. Annual

dues, $5.00 ($1.00 for each additional member of a

family); corresponding membership, $2.00.

R. E. PETIT SHELL BUSINESS

In order to devote more of his time to research

and other matters, Richard E. Petit of South Carolina

has sold his shell and book business to The Shell Cabi-

net, Box 29, Falls Church, Va. 22044, owned and

operated by the Misses Mary E. Young and Isabelle E.

Welch. The International Directory of Conchologists

will now be published by The Shell Cabinet.

IN SATISFACTION 01- POSTAL REGULATION 132-622:

OWNERS: Mrs. Horace B. Baker. Dr. R. Tucker Abbott,

Dr. Charles B. Wurtz. Printing: 1000 copies. Paid sub-

scribers: 596, sliipped by mail.

INFORMATION FOR SUBSCRIBERS

The annual subscription rate for The Nautilus is

$7.00 for individuals and $12.00 for institutions

(domestic or foreign). Subscriptions may begin either

in January or, preferably, in July when a new volume

begins. Send check or money order to "The Nautilus"

to Mrs. Horace B, Baker, Business Manager, 11

Chelten Road, Havertown, Pa. 19083.

Back issues from volume 72 to date are obtainable

from the Business Manager. Volumes 1 through 71 (if

available) may be obtained in reprint or original form

from Kraus Periodicals, Inc., 16 East 46th Street,

NewYork,N.Y. 10017.

Advertising rates may be obtained from the

Business Manager or Editor.

CONTRIBUTORS

Manuscripts: Authors are requested to follow the

recommendations of the Style Manual for Biological

Journals, which may be purchased from the American

Institute of Biological Sciences, 2000 "P" Street,

N.W. Washington, D.C. 20036. Manuscripts should be

typewritten and doublespaced; original and one copy

are required, to facilitate reviews. Tables, numbered

in arable, should be on separate pages, with the title

at the top. Legends to photographs should be typed

on separate sheets. Explanatory terms and symbols

within a drawing should be neatly printed, or they

may be pencilled in on a translucent overlay, so that

the printer may set them in 8 pt. type. There is a

charge of 50 cents per word for this extra service. All

authors or their institutions will be charged 50 cents

per line of tabular material and taxonomic keys. The

pubUshers reserve the right, seldom exercised, to

charge $32 per printed page.

An abstract should accompany each paper.

Reprints and covers are available at cost to

authors. When proof is returned to authors,

information about ordering reprints will be given.

They are obtained from the Economy Printing Co.,

Inc., R. D. 3, Box 169, Easton, Maryland 21601.

[ULY, 1972

THE

NAUTILUS

Vol. 86

No. 1

A quarterly

devoted to

malacology and

the interests of

conchologists

Founded 1889 by Henry A. Pilsbry. Continued by H. Burrington Baker.

Editors: R. Tucker Abbott and Charles B. Wurtz

-c

EDITORIAL COMMITTEE

CONSULTING EDITORS

Dr. Arthur H. Clarke, Jr.

Department of MoUusks

National Museum of Canada

Ottawa, Ontario, Canada K1A-0M8

Dr. William J. Clench

Curator Emeritus

Museum of Comparative Zoology

Cambridge, Mass. 02138

Dr. William K. Emerson

Department of Living Invertebrates

The American Museum of Natural History

New York, New York 1 0024

Mr. Morris K. Jacobson

Department of Living Invertebrates

The American Museum of Natural History

New York, New York 10024

Dr. Aurele La Rocque

Department of Geology

The Ohio State University

Columbus, Ohio 43210

Dr. James H. McLean

Los Angeles County Museum of Natural History

900 Exposition Boulevard

Los Angeles, CaUfornia 90007

Dr. Arthur S. Merrill

Biological Laboratory

National Marine Fisheries Service

Oxford, Maryland 21654

Dr. Donald R. Moore

Division of Marine Geology

School of Marine and Atmospheric Science

1 0 Rickenbacker Causeway

Miami, Florida 33149

Dr. Joseph Rosewater

Division of Mollusks

U. S. National Museum

Washington, D.C. 20560

Dr. G. Alan Solem

Department of Invertebrates

Field Museum of Natural History

Chicago, Illinois 60605

Dr. David H. Stansbery

Museum of Zoology

The Ohio State University

Columbus, Ohio 43210

Dr. Ruth D. Turner

Department of Mollusks

Museum of Comparative Zoology

Cambridge, Mass. 02138

Dr. Gilbert L. Voss

Division of Biology

School of Marine and Atmospheric Science

1 0 Rickenbacker Causeway

Miami, Florida 33149

EDITORS

Dr. R. Tucker Abbott

Delaware Museum of Natural History

Box 3937, Greenville, Delaware 19807

Dr. Charles B. Wurtz

3220 Penn Street

Philadelphia, Pennsylvania 19129

Mrs. Horace B. Baker

Business and Subscription Manager

1 1 Chelten Road

Havertown, Pennsylvania 19083

Second Class Postage paid at Wilmington, Delaware

MBl WHOI LIBRARY

lilH 17XV Z

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