COSTELLACEST AJ A NEW SUBGENUS OF LIMA FROM

THE CRETACEOUS OF THE GULF AND

ATLANTIC COAST PROVINCE
ERLE G. KAUFFMAN
DEPARTMEN1' OF PALEOBIOLOGY
UNITED STA1'ES NA1'IONAL MUSEUM

CONTENTS
Page
ABSTRACT________________ --------------------------------------------------------------· ------------------------------------ 89
I. INTRO DU CTI 0 N---------------------------------------- ________---------------------------------------------- ______________ 90
I I. A CKN 0 WLEDG MEN TS____________________________________________________________________________________________ _____ 9 0
II I. BI 0 STRATIGRAPHY______________________________________________________________________________________________________ 9 0
IV. PHYLOGENY AND EVOLUTION__________________ ________________________________________________________________ 92
v. THE Eco LOGY 0 F GIANT LIMAS ___________________________________________ ----------------------------------- 92
VI. SYSTEMATIC DESCRIPTIONS ___________---------------------------------------------------------------------------- 94
Genus Lima ________________________________________________________________________________ .____________ __ ________ ___ ___ 94
Subgenus Cost ellac es ta __________________________________________ ------------------------------------------------__ 95
Lima ( Costellacesta) riddleiJ n_ sp.____________________________ ______________________________________________ 95
Lima ( Costellacesta) sayrei Stephenson__________________________________________________________________ 97
Lima ( Costellac esta) ins olit a Stephenson________________________________________________________________ 99
VII. REFERENCES CITED_____________________________________________________________________________________________________ 99
TABLE 1: Measurements of Lima ( Costellacesta) riddleiJ n_ sp.________________________________ 96
TABLE 2: Measurements of Lima (Costellacesta) sayrei Stephenson ________________________ 97
TABLE 3: Measurements: Holotype of Lima (Costellacesta) insolita Stephenson____ 99
TEXT FIGURE 1: Stratigraphic occurrence and phylogeny of Costellacesta_______________ 91
TEXT FIGURE 2: Depth distribution of living giant Limas ---------·---------------------------- 93
TEXT FIGURE 3: Cardinal area of Lima ( Costellacesta) riddleiJ n. sp.______________________ 96
PLATE ! ____________________________________________ -------------------- ------------------------------------------------------ 101

ABSTRACT Sandstone Member, Mississippi) , L. (C.)

The new subgenus Costellacesta is pro- sayrei Stephenson (Corsicana Marl, Texas),
posed for large Maestrichtian species of and L. ( C.) insolita Stephenson (youngest:
Lima with an opisthocline shell, large lunule upper Peedee Formation, North Carolina).
and byssal gape, single (posterior) auricle, These represent a single line of descent with-
a resilifer situated posterior to the beak, and out significant break. Successively younger
ornamentation consisting of fine costellae species demonstrate the following evolution-
intercalated between prominent, equally de- ary trends: change in marginal outline, par-
veloped costae. Three species of Costellacesta ticularly an increase in concavity of the
are known and described: L. (C.) riddleiJ dorsoanterior margin; increase in the rela-
n. sp., the type species (oldest; Chiwapa tive size of the lunule and byssal gape; re-

EDITORIAL COMMITTEE FOR THIS PAPER:

A. LEE McALESTER, Department of Geology, Yale University, New H aven, Connec-
ticut
NORMAN F. SOHL, United States Geological Survey, Washington, D. C
HAROLD E. VOKES, Department of Geology, Tulane University, New Orleans, Lou-
isiana
89
 90 Tulane Studies in Geology Vol. 2

duction in the strength of the costae and L. ( C.) sayrei Stephenson (Corsicana Marl,
costellae; increase in number of costellae, and Navarro Group, Texas), and L. (C.) insolita
decrease in number of costae per unit length; Stephenson (Upper Peedee Formation, Owl
possible decrease in maximum adult size. Creek - Providence - Prairie Bluff equivalent,
Costellacesta is possibly ancestral to pli- North Carolina) . These species occur in
cate Cenozoic Limas of the subgenus Plic- successive stratigraphic units, without over-
acesta Vokes. The habitat of C ostellacesta lapping ranges, and probably represent a
was probably analogous to that of living continuous evolutionary sequence. Known
giant Limas ( Acesta, Plicacesta)-cold, deep species of Lima (Costellacesta) are exclusive-
waters of the continental slope and abyss, ly Maestrichtian in age, and occur only in
rarely ranging into warm, shallow shelf en- the upper half of the Exogyra costata zone
vironments. This might explain the uncom- (see Sohl, 1960, fig. 3).
mon occurrence of C ostellacesta in shallow
water Cretaceous deposits of the Atlantic II. ACKNOWLEDGMENTS
and Gulf Coastal Plain Province. The sub- Specimens of Lima (Costellacesta) riddlei,
genus is known only from this area, and n. sp., the type species, were collected and
probably represents an endemic group. donated to the Smithsonian Institution by
Mr. W . C. Riddle of Memphis, Tennessee,
I. INTRODUCTION as part of a program undertaken by the Mid-
Lima is a common element of Gulf and South Earth Science Club to provide the
Atlantic Coast Cretaceous pelecypod assem- Institution with research material. To Mr.
blages. Individual species are widely dis- Riddle and his associates I offer my sincere
tributed and are represented in a variety of thanks. Drs. Harold E. Vokes of Tulane
lithologies, rendering them potentially im- University, A. Lee McAlester of Yale Uni-
portant in biostratigraphic zonation and cor- versity, John Pojeta, Jr., and Norman F.
relation. The American species of Lima are Sohl of the U. S. Geological Survey reviewed
divisible into several distinct lineages, some the manuscript and offered valuable criti-
of which are treated as sections or sub- cism. Dr. Sohl was also instrumental in lo-
genera by many workers. One of the most cating specimens and provided valuable as-
unusual Cretaceous lineages, and one which sistance in stratigraphic interpretation.
appears to be endemic to the Gulf and At- Drawings are by 1vt:r. L. B. Isham and Mr.
la?tic Coast Province, is the group of giant L. R. Purnell, and photography by Mr. Jack
Lima comprising a new subgenus, Costell- Scott, all of the U. S. National Museum.
acesta Kauffman. This lineage is possibly
ancestral to the Cenozoic and Recent sub- III. BIOSTRATIGRAPHY
genus Plicacesta Vokes ( 1963). The three known species of Lima (Cos-
C ostellacesta is characterized by its large tellacesta) occur at widely scattered localities ;
size, inequilateral (opisthocline) shell, single no two species have been found in the same
(posterior) auricle, large lunule and byssal area. Nevertheless, the stratigraphic rela-
gape, slightly oblique resilifer situated pos- tionships between the species are well estab-
terior to the beaks, and by the surface orna- lished (fig. 1 ) . The oldest form, L. ( C..) rid-
mentation, which consists of narrow, sparsely dlei, n. sp., occurs exclusively in the Chiwapa
fluted costae and numerous, delicate, inter- Sandstone Member of the Ripley Formation
calated costellae. Unlike many other Cre- ( Mellen, 1958; equivalent to the Keownville
taceous Lima, species of L. (Costellacesta) Limestone Member of Sohl, 1960, p. 18, fig.
are rare and geographically restricted; few 3) near Ingomar and Pontotoc, M-ississippi.
complete specimens are known. Cenozoic The youngest member of the lineage, L. (C. )
and Recent giant Lima of the subgenera insolita Stephenson occurs only in the upper
Plicacesta and Acesta are similarly restricted. part of the Peedee Formation at the New
Three species of Costellacesta are known Rocky Point quarries, Pender County, North
from Upper Cretaceous sediments of the Carolina (upper part of the Exogyra costata
Atlantic and Gulf Coast Province. These zone, Late Maestrichtian: Stephenson, 1927,
are, from oldest to youngest: Lima (Costell- p. 13). Here it is associated with a fauna
acesta) riddlei Kauffman, n. sp. ( Chiwapa of Owl Creek - Prairie Bluff - Providence age
Member, Ripley Formation, Mississippi), (Stephenson, 1927, Sohl, oral communica-
No. 3 C ostellacesta, a new sub genus of Lima 91

SPECIES OF
TEXAS MISSISSIPPI NORTH
CAROLINA LIMA PHYLOGENY
(COST£LLllCEST.4

; '
t • .l

L. (C.) INSOLITA

z CHIWAPA MBR.
<(

;: KEOWNVILLE LS.
::z:
u
ii: 0..
.... ::>
0
NACATOCH
SAND RIPLEY FM .
"'w a::
<(
::IE
"'0
a::
a:: a:: L (C.) SAYRE!
w <(
0.. >
<(
0..
::> z

L (C .l RIDDLE I

Figure 1-StratigTaphic occurrence, and phylogeny of Costellacesta, showing change

in shape, decrease in overall size, variation in size of the lunule and byssal gape, and
change in ornamentation on progressively younger species of the subgenus. Byssal gape
black where definitely known, otherwise approximated by blank area in lunule. Drawings
for L. (C.) riddlei based on holotype (USNM 132632), for L. (C.) sayrei on paratype
(USNM 132636), and for L. (C.) insolita on holotype (USNM 73431). Outl'ines reduced
14; ornamentation approximately X 1;2.

tion, 1964). Near Ingamar and Pontotoc Bluff strata and those of the Chiwapa Sand-
the Prairie Bluff Chalk disconformably over- stone Member of the Ripley (Sohl, 1960,
lies the Chiwapa Sandstone Member of the fig. 3). This disconformity was recognized
Ripley, and although L. ( C.) insolita does and widely traced by Stephenson and Mun-
not occur here, its age relationship to L. (C.) roe ( 193 7), p. 806), and is present at In-
riddlei, n. sp., is clearly established. gamar and Pontotoc (Sohl, oral communi-
The relative stratigraphic position of L. cation, 1964). Lima (Costellacesta) sayrei
(C.) sayrei from the Corsicana Marl of Stephenson therefore appears to lie strati-
Texas is not as clearly defined. The Corsi- graphically between L. ( C.) riddlei, n. sp.,
cana contains a fauna considered to be tran- and L. (C.) insolita Stephenson. No species
sitional between those of the Chiwapa of Costellacesta are known to have overlap-
Member and Prairie Bluff Chalk. In addi- ping ranges, but they are too rare to be con-
tion, there is a certain amount of faunal sidered important stratigraphic indices.
overlap between the Corsicana and each of The subgenus C ostellacesta is known only
these units. As a whole, the Corsicana fauna from Maestrichtian deposits of the Gulf and
is not duplicated on the eastern Gulf Coast, Atlantic Coast Province (fig. 1). Its range
however, suggesting that no lithologic equiv- lies in the upper part of the E:x:ogyra costata
alent occurs in this area. The Corsicana zone, and is approximately equivalent to the
probably occupies a time span represented range zone of Turritella bilira Stephenson
on the eastern Gulf Coast by a widespread (Chiwapa through Owl Creek: Sohl, 1960,
disconformity between Owl Creek - Prairie fig. 3. Range defined by solid and dashed
92 Tulane Studies in Geology Vol. 2

lines now definitely established; Sohl, 1964, sibly, this reflects a trend toward elimination
oral communication). of the costellae in the lineage. Unfortu-
nately, the only known specimen of L. (C.)
IV. PHYLOGENY AND EVOLUTION insolita is worn and does not permit accurate
determination of the number and total ex-
The Costellacesta lineage appears abruptly
tent of its costellae.
in the American Maestrichtian, and no an-
cestral stock is known. C ostellacesta may The costae exhibit additional changes
have stemmed from some foreign lineage through time, becoming narrower, less
introduced in the Gulf and Atlantic Coast prominent, less numerous, and more widely
Province as a fully differentiated group, but spaced in progressively younger species (fig.
a search of the principal foreign literature 1). The apparent decrease in size of the
has not revealed any members of the sub- adult shell, and variation in size of the lunule
genus, or an obvious ancestral group. Present through time may be additional evolution-
evidence indicates that the lineage is prob- ary trends, but this cannot be substantiated
ably endemic to this area. without more material.
The rarity of Costellacesta in shallow The reduction in size and partial loss of
water Cretaceous Coastal Plain sediments is costellae in the youngest member of the
possibly explained by analogy with the habi- group, coupled with the retention of the
tats of living giant Limas. Vokes (1963a) primary radiating elements of the ornamen-
has shown that living species of Acesta and tation ( costae) , gives rise to a form closely
Plicacesta, which are structurally similar to resembling species of the subgenus Plic-
C ostellacesta (pl. 1, fig. 2) are found pre- acesta, known only from the Cenozoic and
dominantly in deep waters of the outer con- Recent (Vokes, 1963a). Species of Plicacesta
ti~ental shelf, continental slope, and abyss lack costellae and have plicae rather than
( f1g. 2). Only one species is found in less costae. This suggests close relationship or
than 300 feet of water and this species is possible ancestry of Costellacesta to Plic-
rare at this depth. It is reasonable to as- acesta. Additional evidence for the ancestry
sume that species of Costellacesta were simi- of Plicacesta may be provided by future .
larly distributed. If so, the ancestors and studies of certain large, coarsely plicate, In-
early representatives of the subgenus, and dian and European Cretaceous limids, such
perhaps most species of the group, would as Lima ( AcestaJJ) obliquistriata (Forbes)
1

thus be preserved in deep water Cretaceous (India ) , and L. dujardini Deshayes or L.

sediments not presently exposed on the marrotiana d'Orbigny (France). These spe-
emergent portion of the continent. The few cies resemble Plicacesta in gross shell mor-
Cretaceous species that ranged into shallow phology, and are contemporary with and
sublittoral waters are the only ones found older than Costellacesta. They may represent
today in the inner shelf deposits of the At- an extension of Plicacesta into the Creta-
lantic and Gulf Coastal Plain. ceous, or may simply be coarsely ribbed
Although the number of specimens avail- Plagiostoma. Unfortunately, certain critical
able for study is limited, the three known characters of these plicate Cretaceous species,
species of C ostellacesta seem to form an such as the position and nature of the re-
evolvi?g lineage in which some general silifer, and presence or absence of an an-
evolut10nary trends can be noted (fig. 1). terior auricle, are not yet known. Subgeneric
Notably among these is the increase in relationships cannot be determined without
number, but decrease in relative size and this knowledge.
strength of .the costellae in progressively
younger ~pec1~s. The oldest known species, V. THE ECOLOGY OF GIANT LIMAS
L. (C.) riddlei, n. sp., has 1 to 3 prominent Ecologic data for living species of giant
fluted costellae in interspaces between ad- Lima (subgenera Acesta, Plicacesta ) have
jacent costae; the middle member of the recently been assembled by H. E. Vokes
lineage, L. (C.) sayrei has 3 to 6 smaller, ( 1963a, b ). The following comments are
~moo~h costellae per interspace. On L. (C. ) drawn from these and other works on Recent
tnsolita the costellae are even narrower and pelecypods.
fainter, and are visible only on parts of the The giant Limas have wide geographic
shell where the ornament is coarsest. Pos- distribution, occurring from the latitudes of
No. 3 Costellacesta, a new subgenus of Lima 93

II

10
0
>
L

"'c 9

=
CL

~ 8
c
cu
>
.,. 7

.,. 6
c
L
L

...
:3

~ 5

-
0

!;; 3
.D
E
:3
z
2

0 100 200 300 400 500 600 700 800 900 1000 1100 1200 1300 1400 1500
Depth in Fathoms
Figure 2-Depth distribution of living giant Limas (subgenera Acesta, Plicacesta),
Data from Vokes ( 1963a, b).

Norway, Iceland, and Greenland [L. ( Acesta) L. (A.) angolensis Adam and Knudson,
excavata ( Fabricius)} south of Patagonia which occurs from 400 to 500 meters depth
[L. (A.) patagonica Dall}. They are most off Angola. Only one species, L. (A.) goliath
common in the tropical and warm temper- Sowerby occurs uncommonly in water shal-
ate regions of the Indo-Pacific, especially in lower than 300 feet. The two living species
the Philippines, East Indies (Sumatra and of Plicacesta have a combined bathymetric
;Borneo), and Japanese Islands. Plicacesta range of 337 to 2194 feet; the average depth
is unknown outside the Indo-Pacific, while for L, (P.) smithi Sowerby is 1265 feet, and
Acesta has two Atlantic representatives. The for L. (P.) sphoni Hertlein 1650 feet. Since
Cretaceous subgenus Costellacesta, however, dead shells were most commonly dredged in
occurs exclusively in the Atlantic Province, all samples analyzed, and living specimens
and is the only large coarsely ribbed group are not usually differentiated in dredge rec-
of Limas with a posterior resilifer known ords, these depths are at best an approxima-
from this area. It may have occupied an tion of the living range of the various spe-
ecologic niche in the Atlantic during the cies. However, they conclusively show a deep
Cretaceous similar to that presently occupied water habitat preference in the giant Limas
by Plicacesta in the Pacific. (fig. 2).
Most species of Acesta and Plicacesta in- Available temperature records for stations
habit deep water of the outer continental from which Acesta was taken show an aver-
shelf and continental slope (fig. 2 ) . Vokes age water temperature of 48.4 ° F. (Range
cited a bathymetric range of 16 to 1450 35-59.2 ° F.). Eliminating the warm water
fathoms (96 to 8700 feet) and an average form, L. (A.) rathbuni, the average habitat
depth occurrence of 336.4 fathoms (2018.4 temperature for all other species is 45 .7 ° F.
feet) for all living species of Acesta except (Vokes, 1963a, p. 85). In some species,
94 Tulane Studies in Geology Vol. 2

depth distribution is controlled rigidly by "A swimming Lima is a most charming

water temperature (Vokes, 1963a, p. 80). sight. Progress is made with a series of
Temperature may be the primary control on rather languid movements wi th each of
distribution in most species, though this which the long fringe of tentacles slowly
remains to be tested. Costellacesta occurs rises and then gently descends around the
with what have been considered warm, shal- white shell."
low water faunas but it is rare in these as- Although any individual swimming inci-
semblages, as are modern giant Limas, and dent in Lima is short, the animal is capable
was probably more widely distributed in of moving a considerable distance during
cooler, deeper waters during the Upper its lifetime as long as it remains unattached.
Cretaceous. Further, it has the ability, as an adult, to be
Bottom sediment was recorded from 3 3 environment selective, a distinct asset to the
stations yielding species of Acesta. Vokes' survival potential of its offspring. Thus the
analysis of these data (p. 85) shows a individual, and its larvae, are commonly
strong preference for mud, or fine sand and more rapidly and widely distributed than in
mud bottoms (color predominantly green; wholly sessile pelecypods or vagrant ben-
rarely gray, blue) ( 24 localities). Species thonic forms, an important consideration for
of Acesta are more rarely found associated the paleontologist employing Lima in cor-
with sand ( 4 occurrences), Globigerina relation.
ooze, ooze and sand, shell and coral bottom,
rock and shell bottom, and on stones ( 1 oc- VI. SYSTEMATIC DESCRIPTIONS
currence each) . Species of Costellacesta are
found associated with chalky marl (sayrei), Family LIMIDAE
fine argillaceous sand (insolita) and chalky Genus LIMA Bruguiere, 1797
sandstone to arenaceous chalk (riddlei). Lima BRUGUIERE, 1797, Tabl. Encycl. Meth.,
These sediments are well within the range Vers Coq., 2, p. 206. (Valid according to
of those inhabited by Acesta at present. International Code of Zool'ogical N omen-
clature 1961, Art. 16, a, vii; see discussion
Species of Costellacesta probably had habi- of Vokes, 1963a, p. 75, 76).
tat requirements similar to those of living Lima Bruguiere. CUVIER, 1798, Tabl. Elem.,
giant Limas. This would account for the p. 421.
rarity in the shallow shelf sediments that Lima Bruguiere. LAMARCK, 1799, Mem. Soc.
d'Hist. Nat. de Paris, 1, p. 88.
characterize the Cretaceous of the Atlantic Mantellum ROEDING, 1798, Mus. Bolt., p. 160.
and Gulf Coastal Plain. As today, rare Cre- Radula Klein. MORCH, 1853, Cat. Conch.
taceous species lived on the shallow portion Yoldi, 2, p. 57.
of the continental shelf in warmer waters; Radula Klein. ADAMS and ADAMS, 1858,
Genera of Recent Mollusca, 2, p. 556, 557.
L. (C.) riddlei, L. (C.) sayrei, and L. (C) Lima Bruguiere. H. E. VOKES, 1963, Tulane
insolita represent this shallow water element Stud. Geol., 1, No. 2, p . 75, 76.
of Costellacesta. The preservation of com-
Type species: Ostrea lima Linnaeus=Lima
plete fragile shells belonging to the known
squamosa Lamarck; by subsequent tautono-
Cretaceous species probably indicates they
my (Lamarck, 1801) .
lived near their burial site, and were not
transported far after death. Diagnosis: Shell thin, small to moderate-
l'y large; outline ovate, obliquely subovate,
Most species of Lima are sessile forms, or subquadrate; predominantly equivalve,
attaching by byssal threads to rocks and other slightly to greatly inequilateral, with one
hard objects. Some build elaborate nests of (posterior) or two small, unequally devel-
byssal threads in order to protect their non- oped auricles. Anterior byssal gape small
to large; posterior gape uncommonly devel-
retractile mantle tentacles from predators. oped. Lunule commonl'y present. Valves
Vokes ( 1963a) records L. (A.) excavata as nearly smooth, with incised radiating lines,
having been found attached by a byssus; the costellate, costate, or finely plicate. Com-
attachment habits of other living giant Limas missure smooth to crenulate. Hinge line
are unknown. Many species of Lima, includ- short; cardinal area consisting of central
triangular resilifer, erect to oblique, beneath
ing a number of those which normally at- or posterior to beak, bounded laterally by
tach, possess the added ability to swim, and flat cardinal plates for ligament attach-
lie free on the bottom when not swimming. ment. Pallial line entire, faint. Monomy-
arian, posterior adductor muscle scar large,
Yonge (195 8, p. 171) has described the in some cases complex, irregular, common-
swimming: ly situated posterocentrally.
No. 3 Costellacesta, a new subgenus of Lima 95

Subgenus COSTELLACESTA Kauffman, logic features suggestive of the giant Creta-

n. subgen. ceous, Cenozoic, and Recent Limas, in par-
Type species: Lima (Costellacesta) riddlei ticular the valve outline, single (posterior)
Kauffman, n. sp. auricle, large lunule and byssal gape. Regali-
Diagnosis: Large, equivalve, moderately lima is distinct in having radial sculpture
inequilateral opisthocline, slightl'y prosogyre consisting of incised lines, and a resilifer
to pisthogyre. Outline obliquely subovate, which appears to lie directly beneath the
with long straight dorso.anterior edge. Lun- beak. Nevertheless the similarities are so
ule large, well defined. Byssal gape lanceo-
late, large, dorsoanterior to mid-anterior. striking that it is not difficult to envision
Posterior auricle small, dorsoposteriorly sit- the Acesta - Costellacesta - Plicacesta complex
uated, separated from main body of shell by as having arisen from such a group.
well defined auricular sulcus. No anterior
auricle. Surface covered by prominent, The morphologic criteria and the magni-
evenly developed, widely and subequally tude of structural differences which distin-
spaced costae, smooth to finely fluted or guish C ostellacesta from other subgenera of
spinose at intersection with major growth Lima are equivalent to those which separate
lines. Adult costae narrower than inter-
s paces between them, 1-6 fine, evenly far better known limid subgenera such as
spaced, subequally developed costellae com- Acesta and Plicacesta. These well studied
monly intercalated between adjacent costae. subgenera probably arose from a common
Commissure flat to very faintly undulating. Mesozoic stock, and are known to have
Hinge line short, straight, situated mainly
below and posterior to beaks. Resilifer evolved as distinct lineages through the
slightly oblique to hinge line, triangular, Cenozoic. Costellacesta appears to have had
moderately large, situated just posterior to a similar history. The restricted number of
beak. Lateral' cardinal plates flat, triangu- specimens available for study, and lack of
lar, unequal. Pallial line and musculature
unknown. Shell thin. knowledge concerning its ancestors and de-
scendants should not detract from the recog-
Remarks: Costellacesta is distinct from nition of Costellacesta as a distinct subgenus.
the subgenus Acesta Adams and Adams in
lacking an anterior auricle, having a more LIMA (COSTELLACESTA) RIDDLE!, n. sp.
projecting, beak and narrower umbone, in Pl. I, figs. 3, 4, 6
its larger lunule and byssal gape, more promi- Material: A single, nearly complete, large
nent auricular sukus, and in having promi- right valve showing the hinge characters;
nent radial sculpture, consisting of narrow two moderately well preserved, medium size
costae and intercalated costellae, covering val'ves (right and left) ; a large shell frag-
the entire valve. Plicacesta is even more ment.
similar to Costellacesta, but can be distin- Description: Summary of measurements
presented in Table 1. Shell attaining large
guished by its radial plicae of only one size size, equivalve, height greater than length;
which involve the entire thickness of the moderately inequilateral, opisthocline, opis-
shell and are reflected internally. In addi- thogyre. Outline subtriangular to trigonal-
subovate; valves oblique (pl. 1, fig. 4).
tion, the plicae are broader than the inter- Posterior and ventral margins moderately
spaces between them, and are coarser, broad- and subevenly rounded; anteroventral' mar-
er, and more crowded than those on species gin more narrowly rounded; anterior mar-
of Costellacesta. Finally, Plicacesta appears gin straight to very slightly curved (convex
outward), steeply inclined to hinge axis.
to have at least a remnant anterior auricle, Valves moderately convex dorsocentrally,
a more oblique resilifer, and commonly a on um bone; ventral, ventrolateral, posterior
smaller lunule and byssal gape than found flanks slightly convex, gradually flatten-
on Costellacesta. The subgenus Costellacesta ing to·w ard margin. Anterior sl'ope, espe-
cially dorsal half, gently convex centrally,
is uncommon, and is known only from the abruptly truncated and sharply incurved at
Maestrichtian deposits of the Gulf and At- edge of lunule. Lunule poorly known, ap-
lantic Coast Province, from Texas to North parently long, moderately broad, excavated,
and largely fill'ed with a lanceolate byssal
Carolina. gape equal to one-third or one-half of the
Many Jurassic species of Lima attain a anterior margin. Rim of lunule at perime-
size equivalent to that of Costellacesta, Aces- ter of gape apparently narrow.
ta, and Plicacesta. Most of these belong to Beak pointed, somewhat projecting, api-
cal angle slightly greater than 90 ° (pl. 1,
the subgenus Plagiostoma and are not closely fig. 4) · umbone poorly defined, moderately
comparable. One group, however, the sub- convex' costate. Umbonal midline straight,
genus Regalilima Cox, has many morpho- sl'ightly opisthogyre. Beak situated about
96 Tulane Studies in Geology Vol. 2

five-eights the length from the anterior fine, crowded growth lines. Growth lines
margin. Anterior auricle not developed. coarser on ventral one-quarter of valve, at
Posterior auricle small (pl. 1, fig. 4), out- widely spaced intervals crowded and prom-
line subcrescentic, with straight dorsal inent, associated with small concentric con-
margin, gently rounded posterior margin. strictions, particularly prominent near an-
Auricle fiat to slightly convex, separated terior and posterior margins.
from main body of valve by narrow, mod- Hinge line short, straight to slightly
erately deep, well defined auricular sulcus. curved. Cardinal area broad, short, thick,
Entire surface of valve covered with situated below and posterior to beaks, con-
evenl'y spaced, moderate size, subequally de- sisting of a central triangular resilifer, pos-
veloped, straight, primary costae (pl. 1, terior to beak and somewhat oblique to
fig. 3, 4) with steep flanks and flat to gen- hinge axis, bounded by fiat, subtriangul'ar,
tly rounded crests over early part of valve, lateral cardinal plates (fig. 3). Muscle scar,
becoming fluted and more narro·w ly rounded pallial line unknown. Shell thin, smooth
ventrally and laterally near valve margins. internally.
Fluting produced at intersections with
coarsest growth lines. Costae slightly more
crowded posteriorly and centrally than an-
teriorly. Costal interspaces shallow, flat-
based, slightly to moderately broader than
costae on ventral two-thirds of valve, nar-
row dorsally. 1 to 3 costellae intercalcated
in interspaces between adjacent costae on
ventral one-half to two-thirds of valve (pl. Figure 3-Cardinal area of Lima, (Cos-
1, fig. 3) ; 2 or 3 costell'ae per interspace tella,cesta,) riddlei Kauffman, n. sp. showing
most common near margin. First formed position of resilifer (R) posterior to beak.
costellae appear about one-third the height Beak and apex of resilifer reconstructed
below the dorsal margin; second and third (dashed lines) . Ventral margin of cardinal
costellae for each interspace appear at or area buried in matrix. Holotype, USNM
ventral to midvalve, expanding slightly to- 132632; enlarged X 2 % .
ward margins. Ornament of auricle similar,
but with more crowded costae and fewer Remarks: Lima (Costellacesta) riddlei, n.
costellae. Commissure flat to gently undu- sp., is distinct from L. ( C.) sayrei Stephen-
lating at points where costae intersect mar- son, the most closely comparable Cretaceous
gin of valve.
Concentric ornamentation on dorsal three- species, in having a subtriangular outline, a
quarters of val've consisting of very faint, straight to slightly curved (convex outward)
1 TABLE
MEASUREMENTS OF LIMA (COSTELLACESTA) RIDDLE!, N. SP.
(E = ESTIMATED ON BASIS OF RECONSTRUCTION OF VALVE)
Holotype Para type Paratype
USNM 132632 USNM 132633 USNM 132634
Height (mm) 91.5 75 E 59 E
Length (mm) 75.5 62.3
Width (mm) 17.4 11.2 11
Distance, anterior margin to beak,
along line parallel to hinge axis (mm) 41.4 38.4
Length of posterior auricl'e (mm) 18
Height of posterior auricle (mm) 11
Angle of inclination: entire valve 95° 105° E
Angle of inclination: beak, umbo 94° 98° E
Angle between hinge axis and dorso
anterior margin 48° 55° E
Angle between hinge axis and auricular
sulcus 37°
Auricular angle (between posterior, dorsal
margins) 36°
Number of costae 10 mm below beak 41 36 38
Number of costae in 20 mm length, 20 mm
below beak 21 19 21
Terminal number of costae 45 41
Number of costellae in 20 mm length,
20 mm bel'ow beak 5 23 25
Terminal number of costellae '78 54 E
Length of hinge line (mm) 22.9
Angle of inclination of resilifer to hinge
axis 81°
No. 3 Costellacesta, a new subgenus of Lima 97
dorsoanterior margin, a straighter midline, Route 6, 3.5 miles east of Pontotoc, NEl / 4
slightly more convex valves, a smaller, nar- SWl/ 4 sec. 35, T. 9 S., R. 3 E., Pontotoc
rower lunule and byssal gape, fewer but County, Mississippi, in the Chiwapa Lime-
coarser costellae per interspace, arising at a stone Member.
later developmental stage, and straighter more
Types: Holotype, a nearly complete right
numerous, more crowded, and more promi-
valve, USNM 132632. Largest paratype, a
nent costae (compare pl. I, figs. 3, 4 with
medium size right valve, USNM 132633; a
figs. 7, 9). Lima ( Costellacesta) insolita
smaller paratype, about two-thirds of a
Stephenson is smaller, having faint, greatly
crushed left valve, USNM 132634; a para-
reduced costellae, and smooth, low, rounded,
type, a shell fragment sectioned to observe
more widely spaced costae.
thickness and inner valve surface, USNM
Lima (Costellacesta) riddlei is chosen as 132635.
the type species of Costellacesta because it
is the best known of the three species placed LIMA ( COSTELLACESTA ) SAYRE I
in this group; the holotype exhibits most of Stephenson
the essential characters of the subgenus, in- (Pl. I, figs. 5, 7, 9)
cluding the cardinal area. The species is Lima ? sayrei STEPHENSON, 1941, The larger
named in honor of Mr. William C. Riddle invertebrate fossils of the Navarro Group
of Memphis, Tennessee, who collected and of Texas: Univ. Texas Puhl. no. 4101, p.
contributed the types, and many other speci- 146, 147, pl. 23, figs. 12, 13.
mens of Cretaceous mollusks, to the study Material: The primary types of the spe-
cies: 2 nearl'y complete, articulated pairs
collections of the Smithsonian Institution. o.f adult valves (USNM 76476, 132636); 1
Stratigraphic and geographic position: The fragment of a large valve (USNM 76477).
holotype and two paratypes were collected Description: Summary of measurements
from the Chiwapa Sandstone Member ( = presented in Table 2. Shell large, thin, equi-
valve moderately inequilateral, opisthocline.
Keown ville Limestone Member) of the Rip- Outli~e subovate; dorsal margin apparently
ley Formation near Ingomar, Union County, short, straight over posterior auricle; dor-
Mississippi. The third paratype (USNM soanterior margin long, straight to slightly
132634) comes from USGS locality 25418 concave, moderatel'y inclined to hinge axis;
ventroanterior, ventral, posterior margins
( locality 38 of Sohl, 1960, p. 36, 3 7), moderately and unevenly rounded greatest
from roadcuts on new Mississippi State curvature ventroanteriorly, decreasing pos-

TABLE 2
MEASUREMENTS OF LIMA (COSTELLACESTA) SAYRE! STEPHENSON
(L = MEASURED ON LEFT VALVE; R = MEASURED ON RIGHT VALVE;
C = MEASURED ON BOTH ARTICULATED VALVES AND VALVES SAME;
E = ESTIMATED)
Holotype Para type
USNM 76476 USNM 76477
Height (mm) 95 c 84 CE
Length (mm) 84 c 79.2 c
Width (mm) 16.6 R 14.9 R
32.6 c 30 L
Distance beak to anterior margin, along Tine
parallel' to hinge axis (mm) 32.5 c 49 .2 LE
Length of lunule, along margin (mm) 55.5 c 54.6 c
Width of lunule; maximum (mm) 14.4 c 15.7 c
Length of byssal gape, alon~ margin (mm) 40.3 CE
Width of Byssal gape; maximum (mm) 6.1 c
Angle of inclination, entire valve 110 c 108 c
Angle of inclination, beak and umbone . . 88 CE 86 CE
Angle between hinge axis and dorsoanteri?r marg1"!1 40 c 42 L
Angle between hinge axis and dorsopostenor margm 57 c 52 L
Number of costae 10 mm below beak 29 L
Number of costae in 20 mm l'ength, 20 mm below beak 14 L 19 L
Total number of costae at margin 35 L 39 L
Number of costellae in 20 mm length ,
20 mm below beak 30 L 44 L
Total number of costellae at margin 130 L 165 L
98 Tulane Studies in Geology Vol. 2

teriorly; margin concave and notched dor- costae like those on main body of shell, sep-
soposteriorly beneath auricle. Central dor- arated by equally wide or narrower, round-
sal parts of valve slightly to moderately ed interspaces rarely bearing a single, inter-
convex, ventral, ventrolateral, posterior calated, fl'uted costella.
flanks slightly flattened, dorsoanterior Remarks: Lima (Costellacesta) sayrei
flanks steeply inclined into lunule. Lunule
poorly preserved on all specimens (pl. 1, Stephenson is the largest and most ornate
fig. 5); lanceolate, l'ong, occupying dorsal species of the lineage. It is known only
one-half of anterior margin, excavated, mod- from Stephenson's 3 type specimens. Al-
erately deep, apparently sharply defined though he partially exca\'ated them, Stephen-
and partially over hung by abruptly trun-
cated dorsoanterior margin. Anterior byssal son did not note the lunule with its large
gape long, moderately narrow, lanceolate byssal gape. Consequently, he reversed an-
(pl. 1, fig. 5) ; commissure slightl'y upturned terior and posterior throughout his descrip-
around gape, flat to very faintly undulat- tion, considering the left valve the right,
ing over remainder of valve.
Beaks not known. Umbone slightly to and the auricle as being anterior. Compari-
moderately convex, apparently erect to son of the species of Costellacesta with the
slightly prosogyre. Anterior auricle absent; similar, living, Lima (Plicacesta) smithi (pl.
anterior edge of cardinal area rounded, I, figs. 1, 2) clearly shows that the major
slightly projecting and incurved at dorsal
extremity of byssal gape, similar to ill'us- byssal gape is dorsoanterior and situated in
trated specimen of L. (Plicacesta) smithi a prominent lunule, the anterior auricle is
(pl. 1, fig. 1). Posterior auricle small, flat, absent, and the valves are opisthocline. An-
incompletely known on both specimens (see terior and posterior are determined on L. (P.)
Stephenson, 1941, pl. 23, fig. 12) apparently
straight dorsally, rounded posteriorly, with smithi by the resilifer (oblique and pos-
distinct ornamentation. Posterior auricular terior to the beak) and the well developed
sulcus narrow, moderately deep, well' de- posterior adductor muscle scar. A parallel
fined, an enlarged and much deepened in- orientation in Costellacesta is determined by
terspace between costae. Hinge line not pre-
served, assumed similar to L. (C.) riddlei, the position of the lunule and byssal gape
n. sp. Pallial line, musculature, unknown. on all species, and by the position of the
Ornamentation consisting of 34 to 38 resilifer in L, ( C.) riddlei, n. sp.
simple primary costae separated by 3 to 6
fine intercalated costellae per interspace The distinction between L. ( C.) sayrei and
(pl. 1, fig. 7). Costae, prominent, subevenl'y L. (C.) riddlei have been discussed under the
spaced, equally developed, narrow, steep- "Remarks" section for the latter species.
sided, with rounded crests, originating- on
early part of umbone, their trace straight L. (C.) sayrei is distinguished from L. (C.)
to slightly curved over most of shell, gently insolita Stephenson by its larger size, more
sinuous ventrally and ventrolaterally at in- rounded outline, more prominent and slight-
tersections with major concentric elements. ly more numerous fluted costae, and espe-
Costae slightly more crowded posteriorly
than anteriorly; 2 to 3 costae cl'osely crowd- cially by its more numerous, more extensive
ed on inturned edges of lunule and on steep beaded costellae. L. (C.) sayrei also appears
part of dorsoposterior flank descending into to have a larger byssal gape, and better de-
auricular sulcus. Costae fluted to subspi- fined lunule. Stephenson illustrated the holo-
nose at somewhat regular intervals where
intersected by major growth lines (pl. 1, type ( 1941, pl. 23, fig. 12).
fig. 7) ; 4-10 flutes per centimeter, best de- Stratigraphic and geographic distribution:
veloped ventrally. Costellae fine, thread- Lima (Costellacesta) sayrei is known only
like, evenly spaced and equall'y developed in
broad flat inters paces between costae; from the Corsicana Marl, Navarro Group,
straight to slightly sinuous, beaded due to at U.S.G.S. locality 15621, in a ravine east
irregularities produced at intersections with of Medio Creek, 0.8 mile south of Castro-
majority of growth lines (pl. 1, fig. 7). 1
to 2 costellae present between each pair of ville road, near Castroville, Texas; at U.S.G.S.
costae on earliest part of umbone preserved; locality 15622, from a bluff on Medio Creek
number increasing by intercalation through- about 0.8 mile "below" (south of) Castro-
out growth of shell, with 5 to 6 commonly ville road; and from 6 miles east of Castro-
found in interspaces near margin of adult
shell (pl. 1, fig. 7, 9). Concentric ornament ville, Texas (Texas Bureau 53; locality of
consisting of faint to well defined but small, holotype).
crowded growth lines over entire shell, and Types: Holotype, the largest set of rela-
2 to 3 scattered, narrow zones of moderate tively complete articulated valves USNM
to coarse growth lines, and more rarely
lamellae, near margin ( <;mter 1 inch or less). 76476. Paratypes, a well preserved set of
Ornamentation of posterior auricle dis- articulated valves (U.S.G.S. loc. 15621),
tinct, consisting of crowded, coarsely fluted USNM 132636 (pl. I, figs. 5,9) (formerly
No. 3 Costellacesta, a new subgenus of Lima 99
a part of USNM 76477, unfigured paratype TABLE 3
lot); a shell fragment showing the orna- MEASUREMENTS: HOLOTYPE OF
ment (USGS loc. 15621, USNM 76477). LIMA (COSTELLACESTA) INSOLITA
STEPHENSON
LIMA ( COSTELLACESTA,.) INSOLITA Height, including reconstruction
Stephenson of beak (mm) · 63
Length (mm) 53.3
Pl. I, figs, 8, 10 Width (mm) 9.8
Lima inso.Zita STEPHENSON' 1927' Additions Distance, anterior margin to
to the Upper Cretaceous Invertebrate fau- beak, along line parallel to
nas o.f the Carolinas: Proc. U. S. Natl. hinge axis (mm) 26.4
Mus., v. 72, art. 10, p. 13, pl. 5, fig. 10. Angle of inclination, entire valve 105°
Material: A single, nearly complete, worn Angl'e of inclination; beak, umbo 85°
adult left valve, the holotype (USNM Angle between hinge axis
73431). and dorsoposterior slope 42°
Description: Summary of' measurements Angle between hinge axis and
presented in Table 3. Shell moderately dorsoanterior slope 46°
large, thin, presumably equivalve; inequilat- Number of costae 10 mm
eral', moderately opisthocline; height greater below beak 28-30
than length. Outline subovate, with long, Number of costae in 20 mm
slightly concave, dorsoanterior margin. Ven- length, 20 mm below beak 17
troanterior margin strongly curved; ven- Total terminal number of costae 28-30
tral, posterior margins moderately, and sub-
evenly rounded (pl. 1, fig. 10) . Dorsal mar- of costae, fainter discontinuous costellae, and
gin not preserved. Umbone slightly to mod-
erately convex; ventral, ventrolateral, pos- the smaller, less rounded valve are sufficient
terior flanks slightly convex medially, be- to distinguish the species from L. (C.) sayrei
coming flatter toward margin. Dorsoan- and L. (C.) riddlei, n. sp. The species ap-
terior flank abruptly truncated, moderately pears to be most closely related to, and de-
to narrowly rounded at edge of lunule.
Lunule poorly defined, depressed, occupying rived from L. (C.) sayrei. It is the youngest
two-thirds of dorsoanterior margin. Byssal known member of the lineage. Apparent
gape unknown, probably long and narrow. reduction of the costellae, and retention of
Beaks unl•nown. Umbo suberect, broad, well defined costae, may reflect ancestry to
not well defined. No anterior auricle devel-
oped. Posterior auricle, if present, not pre- the Cenozoic - Recent subgenus Plicacesta
served. Hinge l'ine, internal structures un- Vokes.
known. Stratigraphic and geographic position: The
Radial ornamentation consisting of 28 to only known specimen is from the upper part
30 straight simple primary costae extend-
ing from early part of umbone to margin. of the Peedee Formation, in sediments bear-
Costae narrow, low, rounded, smooth equally ing an Owl Creek-Prairie Bluff fauna (upper
developed, subevenly spaced (more crowded part of the zone of Exogyra costata Say: late
anteriorly), separated by much larger, flat Maestrichtian). It was found at U. S. Geo-
interspaces, a few of' which bear very faint,
fine, scattered, discontinuous costellae (pl. logical Survey locality 13585, in the New
1, fig. 8), 1 to 2 per interspace preserved, Rocky Point quarries, one mile northeast of
but probably much more extensive on un- Rocky Point Station, Pender County, North
worn shells of species. Costellae smooth, Carolina.
straight, very low, rounded. Concentric or- Type: Holotype, USNM 73431; a nearly
namentation consisting of numerous, crowd-
ed, very faint growth l'ines over entire valve, complete left valve.
and a few, scattered, moderately prominent
growth lines near margin. Commissure VII. REFERENCES CITED
smooth to very broadly undulating. ADAMS, HENRY, and ARTHUR ADAMS, 1858,
The genera of Recent Mollusca: v. 2, p.
Remarks: As in L. (C.) sayrei, Stephenson 556-558, London, J'ohn Van V oorst.
did not note the position of the lunule and BRUGUIEREI, J. G., 1797, Tableau encyclope-
main byssal gape in this specimen, and con- die methodique, OU par ordre de matiere:
sidered it a right valve; anterior and pos- Histoire naturelle des Vers: v. 2, p. 206,
Paris & Leige.
terior are reversed throughout his descrip- CUVIER GEORGES, 1798, Tableau elemen-
tion. taire' de l'histoire naturelle des animaux:
The holotype is worn, and thus the precise p. 421, Paris.
LAMARCK, J. B. P. A. DE, 1709, Prodrome
nature of the delicate surface ornamentation d'une nouvelle classification des coquilles:
cannot be determined. In spite of this, the Mem. Soc. d'Hist. Nat. de Paris, v. 1, p.
wide spacing of the costae, lower number 88.
100 Tulane Studies in Geology Vol. 2

LAMARCK, J. B. P. A. DE, 1801, Systeme des the Carolinas: Proc. United States Natl.
animaux sans vertebres: v. 1-8, p. 1-432, Mus., v. 72, art. 10, p. 13, pl. 5, fig. 10.
Paris. STEPHENSON' L. w., 1941, The larger inver-
MELLEN, F. F., 1958, Chiwapa Sandstone tebrate fossils of the Navarro Group of
Member of Ripl'ey Formation in Creta- Texas: Univ. Texas Publ. No. 4101, p.
ceous shelf sediments of Mississippi: Mis- 146, 147, pl. 23, figs. 12, 13.
sissippi State Geol. Surv., Bull. 85, p. STEPHENSON, L. w., and w. H. MUNROE,
49-54, figs. 28-33. 1937, Prairie Bluff and Owl Creek For-
MORCH, 0. A. L., 1853, Catalogus conchyli- mations of eastern Gulf region: Ameri-
orum quae reliquit D. Alphonso d'Aguirva can Asscc. Petrol. Geol. Bull., v. 21, no.
& Gadea Comes de Yoldi: Pt. 2, Acephala, 6, p. 806.
Echinodermata Hafniae, p. 57.
VOKES, H. E., 1963a, Studies on Tertiary
ROEDING, P. F., 1798, Museum Boltenianum and Recent Giant Limidae: Tulane Stud.
sive catalogus cimeliorum e tribus regnis Geol., v. 1, no. 2, p. 73-92, pls. 1, 2.
naturqe: pt. 2, p. 160, Hamburg.
SOHL, N. F., 1960, Archeogastropoda, Meso- VOKES, H. E., 1963b, Additions to a cata-
gastropoda and Stratigraphy of the Rip- logue of the described Recent and Terti-
ley, Owl Creek, and Prairie Bluff Forma- ary species of Acesta and Plicacesta : Tu-
tions: United States Geol. Surv. Prof. lane Stud. Geol., v. 2, no. 1, p. 18-20.
Ppr. 331-A, p. 18-20, 22, fig. 3. YONGE, C. M., 1958 (2nd printing), The sea
STEPHENSON, L. w., 1927, Additions to the shore: p. 171, London, Collir.s, St. James
Upper Cretaceous invertebrate faunas of Place.

June 30, I964

EXPLANATION OF PLATE I
Figures Page
I, 2, - Lima ( Plicacesta) smithi Sowerby_______________________________________________________________________ 98
I, Interior and 2, exterior views (XI) of a left valve from Echigo, Japan;
USNM 344887; Hirase Collection, Division of Mollusks, USNM. Illustrated
for comparison with members of L. (Costellacesta).

3, 4, 6 - Lima ( Costellacesta) riddlei Kauffman, n. sp. ____ ·-----------------------------·------------ ____ 95

3, Portion of right valve (fig. 4) showing adult ornamentation ( X2 ) .
4, Lateral view (XI), holotype (USNM I32632), a right valve.
6, Anterior view (XI) of holotype showing margin of lunule.

5, 7, 9 - Lima ( Costellacesta) sayrei Stephenson___________________________________________________________ 97

5, Anterior view (XI ) of Stephenson's unfigured paratype (USNM I32636)
showing limits of lunule, position of byssal gape (space inside byssal gape
retouched to improve contrast), infolded anterior margin of cardinal area.
7, Portion of the same adult shell near margin (X2) showing details of orna-
mentation: left valve.
9, Lateral view of left valve (paratype, USNM 132636, of figs. 5, 7) with
auricle broken off.

8, I 0 - Lima ( Costellacesta) insolita Stephenson ----------------------------------------------------------- 99

8, Portion of left valve shown in fig. I 0 ( X2) , showing costae and faint, dis-
continuous, intercalated costellae.
IO, Lateral view (XI ) of left valve, the holotype (USNM 7343I) and only
known specimen.
No. 3 Costellacesta, a new subgenus of Lima 101

PLATE I