Bulletins of American paleontology

Begun in 1895 ЧУ

VO
LUME 110, NUMBER 351 SEPTEMBER 27, 1996

Neogene Paleontology in the Northern
Dominican Republic

16. The Family Corbulidae (Mollusca: Bivalvia)
by
Laurie C. Anderson
17. The Families Cuspidariidae and Verticordiidae (Mollusca: Bivalvia)
by

Peter Jung

Paleontological Research Institution
1259 Trumansburg Road
Ithaca, New York, 14850 U.S.A.

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fal

Begun in 1895

Yo
LUME 110, NUMBER 351 SEPTEMBER 27, 1996

Neogene Paleontology in the Northern

|
|
|
Dominican Republic
16. The Family Corbulidae (Mollusca: Bivalvia)
by

Laurie C. Anderson

17. The Families Cuspidariidae and Verticordiidae (Mollusca: Bivalvia)

by

Peter Jung

Paleontological Research Institution
1259 Trumansburg Road

9 Ithaca, New York, 14850 U.S.A.

ISSN 0007-5779 |
ISBN 0-87710-442-5
Library of Congress Catalog Card Number: 96-70358

Printed in the United States of America
Allen Press, Inc.
Lawrence, KS 66044 U.S.A.

CONTENTS

16. Тһе Family Corbulidae (Mollusca: Bivalvia)

Laurie C. Anderson Page
ii ЕЕЕ у ао TR STR, 5
EA A и қыса ЧУМ, и 3
INWOUUCHON „тна бе нас се с та зе mene A аный E adito qiiia iud veru decem S 5
Acknowledgmehts e.ri ereas a мори a Өз кеті e uk untra СН te MP quid QUE чак ce. даа 6
Blostratigraphy and Paleoecology: «ici suse A __-____ = о АИ ае da a 2.. 6
Biogeography аа e Eds culata Rs УАН se aan des эле nen 11
Abbreviations of Repository Institutions игл ______--_ de suis ҚЫ» ОЧ н УУЧ ба Geese es 11
Аманат Аны ЕН ене. у en caa сае о rn 11

ee a ad va 11
SYSE er TE A О лы Ақы аа 12
Family Corbulidas Lamarck, 1818 ИЕ ES AR A AA E E COINS DOS RSCG шы ALS + 12
subfamily-Corbülnas Lamarok, 16185 AA ени A A Oe A A Та 12
Genus Corbula ВЕШТ, 1797. VELA A A CES OEA слали WO ea A A ISI ne 12
Subgenus Borkrocorbula Gabb, 18738: ii a nennen a оя 12

Corbula (Bothrocorbula) viminea Guppy, 1866Ь................................................... 12

Subgenus Caryocorbula Gardner 1926 улл... у a са сирка ве 14

Corbula (Caryocorbula) dominicensis Gabb, 18730 ................................................ 14

Corbula (Caryocorbula) sericea Пай, 1898... aa a ER a E a hh ehh нива nee 15

Subgenus Juliacorbula Olsson and Harbison, 1953 .................................................. 17

Corbula (Juliacorbula) fossilis Pilsbry, 1922 .................... eher навек кна кна кна как 17

Subgenus Panamicorbula Pilsbry, 1982: ........................4...6...6 ENE а лан ваа на АА на и ал 18

Corbula (Panamicorbula) canae, new species ........................................2 ........... 19

Corbula (Рапатисотвшаувре, Er ER TE EEE TE ANN IDEEN қ MARET IQ, 19

Subgenus Varicorbula Grant and Gale, 1931 .................................................... ы аа 20

Corbula (Varicorbula) sanctidominici Maury, 1925....................................5..4 nn 20
WNOLETSTOSS САД... оо ом REC Аа a ҰНТАҒЫ ЖА ER PERE: ARNO ХА ALAIOR МАТЕ. нет 22
APPEAR] ic ван 22 A АА IE ost RUN QUEE UIS EUM UO ымаа MIA ы. 26
Appendix 2 senda A AAA de dom HAE A qud en QT. S AI ҚЫМЫЗ ЫЛА RODAR и 26
BIBRUSS На ВАРНА a Ги КО КРЕ ИНИ AR a ME 28
Iron a арии фа сағызды а а А MURAL а A DN ма. E

17. The families Cuspidariidae and Verticordiidae (Mollusca: Bivalvia)

Peter Jung
“ерте e Ue VE DD nu Em За ари A ARE ren
ОИЕ ID A DE ERAS EE оваа енд мо зла x A E
ОО ent Re boa I места
Sei ui ALTI а ЛА OM ESSO ИО ИТ лы ы e acest ОЕР Si d nce rhe
Biostratigraphy and Paleobiogeography . ... serere ан nen hee thesis ааа оао
Abbreviations of Repository Institutions us vice et eet te tt ет nenne rr нан на зина аве
So cn и vey Ga ect кале аа ea атақ O
Men rc ea es in o A AA 15-3 A фанов ER FERN TEE FILE TI II. TR
Sry nl АНА E ТЕТЕ УСАК РЕ er oe SEE AA IT II e
ТАЦУ СОВА ОВ Ci. ura en are ne a же укун тачат да саити а ОНА
Genus Cardiomya A Adams, 1864 ..................:.555 555555555545. rr кана нок аа ee
Subgenus Cardiomya: valuisse eco se OSEE VR а АТ CRY AL ЧО ВВ Ке МИ Ақа USE CUN a a rs
C. (С). islahispaniolas Monty, 1917) .......... pea валка ва као ss Си ҚА бе Зи ии
Subgenus Водена: Пай, 1903.44 за con eo ита un sa ame e a va БАНА ни €
С (В) distra (DAL, 1908). ес лас часна ке а +з an as ж E
Genus Plectodon Carpenter, 1804... 0.0... ¿malo din жаз acest te A oe айне л Б ИА сазы ша,
P. gronulams (ГЕЙ, 1881) ......дезес. зээ аа за ж re,
bue VETO een ets es A ТЕ нА қак окт о а
Genus Ralis Dall, 1896 ооа ee
МН Jamalcensis: ПИШ, 908) .. cele ee rare reo crx esi s ораи
Genus ПУНИ АН OI ТЕЩА c.v A с за Met ee es ДО
T ornate d ОТВЕТУ, ЕН. АНА ОН, IVA кееш, A pen ы ET
T. Расо, Нет Species 2. ооо пана гръцки mcs НА CHO ЕИ ИОВА e ада ори c
T bowdenensis (Dall, 1903) а VOI a ues скокови aaa aa ж OX о» оне
cr Ci сы аа: VA € Ceo a E лае иска a клек A пара
A ыы ий еа a ar У а uA ESL LH DE ышк

LIST OF ILLUSTRATIONS AND TABLES |

16. The Family Corbulidae (Mollusca: Bivalvia)
Laurie C. Anderson
Text-figure Page
І. Index map of areas investigated in the Cibao Valley, Dominican Republic: vec се. nn 6
2. Part of the Rio Cana section showing the stratigraphic distributions of Corbula species ............................ 7
3. Part of the Río Gurabo section showing the stratigraphic distributions of Corbula species .......................... 8
4. Plot of first and second principle component scores (PC1 and РС2) of right valves of Corbula sericea ................. 9
5. Plot of PC1 and PC2 of right valves of Corbula sericea from shallow-marine deposits of the Río Cana and
a o alo 9
6. Plot of PCI versus stratigraphic section (upper) and PCI versus sediment type (lower) for Corbula viminea ............. 10
Table
1. Corbula viminea valves from various sediment types for each stratigraphic section ............................... 11
ШІМ a UA оо ија АД е ООА rr eei dut KI RT URL MI e m EM 13
a ðt Corbula (CANYOLOTDRIA)LUDTUNICENEIS A A TL Mun а оне o sem DOS 15
Ay MDEA E COOU бај дај од ССА Ќе Го ќа ы села A ніл 16
О. MOUSSE OF CODIA (иа сори та) IN n dee € акен pull be нн не Од Wick GU) атана пне 18
A a o lali il ld a UD OMIE: RS ра ннен аты 19
7. Measurements of Corbula (Panamicorbula ) A 1) ви КӨК eis ОИЕ i. Т 20
Se гисивштешеше OL COPLA ui енен көлін ны ние 2 |
17, The Families Cuspidariidae and Verticordiidae (Mollusca: Bivavlia)
Peter Jung
Text-figure
1. Index map showing location of investigated areas in the Cibao Valley, Dominican КӨППЕН ET КЕСЕРІ! 36
2. Columnar section of Río Gurabo showing occurrences of species dealt with herein ............................... 37
3. Río Gurabo: upper part of columnar section showing occurrences of species dealt with herein ....................... 37
4. Columnar section of Río Cana showing occurrences of species dealt with herein ................................ 38
5. Section exposed in Maury's Bluff 2 on Río Mao showing occurrence of Cardiomya (Cardiomya) islahispaniolae
(Мангу, 1917) and stratigraphic positions Of NMB localities. а 00 38
6. Section exposed at mouth of Arroyo Bajón on Río Mao showing occurrence of Cardiomya (Cardiomya) islahispaniolae
(Mary: 1907) АПИ ВАСА ОНИ positions осал... E ио 39
7. Section exposed at the downstream (eastern) end of Maury’s Bluff 3 on Río Mao showing occurrence of Cardiomya
(Cardiomya) islahispaniolae (Maury, 1917) and stratigraphic positions of ММВ localities ....................... 39
8. Length/height diagram of Cardiomya (Cardiomya) islahispaniolae (Maury, 1917) ............................... 42
9; AOS bewi diagram of Caraiomya: (Bowdenia) distira (Dall, 1903) i са isi evs ev nen 43
IO. (engi БӘ ра ои ои oi Pieorodon granuiatus (Wall, 1991). (es err eva cule obi av 04444 ана а ie 45
DE Berl Der Шо 010177 70mutcensis-QDüll, 1903)... ovo hoes о nese 46
e ТОО ТОЙТ ОЛАЙ O e 1842 eis re CR do ra luc ee. об 45
13. Histogram showing rib number distribution of Trigonulina ornata d'Orbigny ..22...2...........22222.22.. .. 2 1 4 4 4Ә4 45
14. Length/height diagram of Trigonulina pacifica, new AI ТУТ” 49
15. Histogram showing rib number distribution of СОПИШТЕ BUCY Da MEN ее E
16. Histogram showing rib number distribution of Trigonulina bowdenensis ЮЕШ IMUS ооо ^
17. Length/height diagram of Trigonulina bowdenensis ШШ ШШ КЕ a ure. 52
Table
1. Numbers of lots and specimens of each of the seven species of Cuspidariidae and Verticordiidae dealt with
АО ЗА И У en cn нын аска 1 Шш
2. Number of specimens available from the Neogene of the Dominican Republic and from the Bowden Formation
a. a
3. Diagnostic features of the three species of Trigonulina dealt with herein ................................ о 50

NEOGENE PALEONTOLOGY IN THE NORTHERN DOMINICAN REPUBLIC 16.
THE FAMILY CORBULIDAE (MOLLUSCA: BIVALVIA)

LAURIE C. ANDERSON

Department of Geology and Geophysics, Louisiana State University,
Baton Rouge, Louisiana, 70803

ABSTRACT

Corbulid bivalves (Myacea: Corbulidae) are an important constituent of fossiliferous Neogene sediments of the Cibao Valley,
northern Dominican Republic. Six corbulid species referred to five subgenera within the genus Corbula are described and figured
(Bothrocorbula, Caryocorbula, Juliacorbula, Panamicorbula, and Varicorbula). One species is new, Corbula (Panamicorbula)
сапае. This 15 the first description of a fossil Panamicorbula species and the first description of a Panamicorbula species outside
of the eastern Pacific. The distribution of Dominican corbulids is strongly influenced by paleoenvironmental conditions such as
salinity, depth, and bioclastic fabric. Corbulids are most common in Miocene sediments that were deposited predominantly in
shallow-marine waters. Corbula (Caryocorbula) sericea Dall, 1898 has the widest stratigraphic and geographic distribution within
the study area and is especially abundant in brackish-water and very shallow-marine deposits. Corbula (Bothrocorbula) viminea
Guppy, 1866b is also common in shallow-marine and brackish-water deposits. Corbula (Varicorbula) sanctidominici Maury,
1925 occurs predominantly in shallow-marine sediments, whereas Corbula (Panamicorbula) canae n. sp. is restricted to brackish-
water deposits of the Upper Miocene Cercado Formation in the Río Cana section. Corbula (Caryocorbula) dominicensis Gabb,
1873b and Corbula (Juliacorbula) Jossilis Pilsbry, 1922 are rare. Corbula dominicensis is apparently restricted to the upper Lower
to lower Middle Miocene Baitoa Formation of the Río Yaque del Norte section, and C. fossilis is found in Upper Miocene
sediments of the Cercado and Gurabo Formations. Dominican corbulid species show close morphologic affinities to species of
the Pliocene Bowden Formation of Jamaica, Neogene units of the Caribbean coast of Central America such as the Gatun
Formation and sediments of the Limón Basin, Miocene and Pliocene deposits of Trinidad, and Miocene to Pleistocene sediments
of Florida.

RESUMEN

Los bivalvos corbúlidos (Myacea: Corbulidae) constituyen una parte importante de los sedimentos fosilíferos del Neógeno en
el Valle de Cibao al norte de la República Dominicana. A continuación, se describe y se dibuja seis especies pertenecientes a
cinco subgéneros dentro del género Corbula, es decir, Bothrocorbula, Caryocorbula, Juliacorbula, Panamicorbula, y Varicorbula.
Corbula (Panamicorbula) canae forma una nueva especie. Por primera vez se describe la especie fósil Panamicorbula y es la
primera vez que se describe la especie Panamicorbula fuera del Pacífico este. La distribución de corbúlidos dominicanos está
influenciada fuertemente por condiciones paleoambientales tales como la salinidad, profundidad de agua y fábrica bioclástica.
Los corbúlidos se encuentran más comúnmente en aquellos sedimentos del Mioceno depositados predominantemente en aguas
Marinas someras. Dentro del la zona de estudio, Corbula (Caryocorbula) sericea Dall, 1898 muestra tener la distribución estra-
tigráfica y geográfica más amplia y abunda sobre todo en agua salobre y en depósitos marinos de agua muy someras. Corbula
(Bothrocorbula) viminea Guppy, 1866b también se encuentra comúnmente en depósitos marinos de agua somera y en depósitos
de agua salobre. Corbula (Varicorbula) sanctidominici Maury, 1925 aparece en sedimentos marinos de poca profundidad mientras
que Corbula (Panamicorbula) canae, п. sp. se encuentra restringida a depósitos de agua salobre en la formación Cercado del
Mioceno superior en la sección del Río Cana. Corbula (Caryocorbula) dominicensis Gabb, 1873b y Corbula (Juliacorbula)
Jossilis Pilsbry, 1922 son poco frecuentes. Aparetmente, C. dominicensis queda restringida entre la parte superior del Mioceno
inferior y la parte inferior del Mioceno intermedio de la formacio Baitoa en la secciön del Rio Yaque del Norte. Corbula fossilis
se encuentra en los sedimentos de las formaciones Cercado y Gurabo con edad del Mioceno superior. Las especies de corbúlidos
dominicanos muestran una afinidad morfológica a las especies de la formación Bowden del Plioceno en Jamaica, a las unidades
Neögenas de la costa caribeña de centroamérica tales como la formación Gatun y los sedimentos de la cuenca Limón, a los
depósitos de edad Miocena y Pliocena en Trinidad y a los sedimentos de Florida que comprenden desde el Mioceno hasta el

Pleistoceno.
INTRODUCTION graphic framework. The units studied include the up-
This k i f: 1 : А рег Lower to lower Middle Miocene Baitoa, Upper
Марн ра арағы анта “ КОДОО MARES Miocene Cercado, Upper Miocene to Lower Pliocene
Northe оп Neogene fossils of the Cibao Valley in the Gurabo, and Lower Pliocene Mao Formations. Al-
et E. Dominican Republic (Text-fig. 1). Saunders though not highly diverse, corbulid bivalves occur
“. (1986) provide detailed information on mea- throughout the stratigraphic section. They are most

8

Т А А Т я
ed sections and samples, and a general biostrati- abundant, however, in brackish-water and shallow-

marine sediments, which are typical of Miocene de-
posits in the study area.

ACKNOWLEDGMENTS

I am especially grateful for the generosity and as-
sistance of Peter Jung, of the Naturhistorisches Mu-
seum Basel, Switzerland and Emily Vokes of Tulane
University who provided long-term loans of extensive
collections. A number of other institutions generously
lent samples and specimens, and I would like to thank
Warren Allmon of the Paleontological Research Insti-
tution; Thomas Waller, Warren Blow, and Jann
Thompson of the United States National Museum of
Natural History, Smithsonian Institution; Gary Rosen-
berg and Elana Benamy of the Academy of Natural
Sciences of Philadelphia; Silvard Kool of the Museum
of Comparative Zoology, Harvard University; Paul
Taylor and Paul Jeffery of The Natural History Mu-
seum, London (British Museum, Natural History); and
Klaus Westphal of the Museum of Geology, University
of Wisconsin-Madison. Thanks also to Juan Lorenzo
for translating the abstract, and to Emily Vokes, Jay
Schneider, and John Kruger for reviewing the manu-
script. This research was supported by the National
Science Foundation (EAR-9316363), Sigma Xi, the
Paleontological Society, and the Geological Society of
America.

BULLETIN 351

BIOSTRATIGRAPHY AND PALEOECOLOGY

Within the study area (Text-fig. 1), it is possible to
trace changes in corbulid species distributions and in-
traspecific variability in a number of stratigraphic sec-
tions that record the interplay of paleoenvironment,
time, and geography. Anderson (1994) documents in
detail the relationship of corbulid species distributions
and intraspecific variability to paleoenvironmental
conditions in these sediments and those findings аге
reviewed here. Distributions of the most common spe-
cies, Corbula (Caryocorbula) sericea, Corbula (Both-
rocorbula) viminea, Corbula (Varicorbula) sanctidom-
inici, апа Corbula (Panamicorbula) canae, track ра-
leoenvironmental conditions, and represent migrations
into and out of the study area over time rather than
speciation and extinction events (Anderson, 1994).
Corbulid abundance and diversity reflect a ргеѓегепсе
for marginal-marine and shallow to intermediate ma-
rine habitats (< 100 m), which is compatible with hab-
itat distributions of living corbulids. This correspon-
dence of species distributions to paleoenvironmental
conditions causes a temporal pattern in species distri-
butions because paleoenvironmental conditions
changed systematically through time in the study are
(Saunders et al., 1986). In the study area, corbulids аге
most abundant and diverse in Miocene marine sedi-
ments deposited primarily in shallow (approximately

GUAYUBIN

Haiti Dominican

Republic

Text-figure 1.—Index map of areas investigated in the Cibao Valley, Dominican Republic (after Saunders et al., 1986).

€—À
{ 1 Rio Cana
[_ JUpper Cenozoic 2 Rio Gurabo
Oligocene-Early Miocene? 3 Rio Mao

4 Rio Amina
5 Cañada Zalaya

6 Rio Yaque del Norte
7 City of Santiago

8 Arroyo Puñal

9 Rio Verde

CORBULID BIVALVES: ANDERSON ~

Е : | | E Río Cana Section
E $ | о Coral Communities
ш =; : Grass flats
8 8 | ІШІ ar Bh e
; E н ige more
| (>10 - 20 n»
E B ВЕН
г с ^ : 8 EZ Уез shallow marine
5 Y 57 (<30 т)
Е p E 5 рд Marine (approximately
© Fy Е | А 30 to < 100 m)
| M
MER
ME
ТІ
| E
5 2::
o | : 3
Y Y а

ау Вазе 2.—Part of the Río Cana section showing the stratigraphic distributions of Corbula species. Coral communities are based on
‚lage linkage cluster analysis of coral assemblages from Budd er al. (1996). Criteria used to distinguish environmental categories are outlined

n Appendix 1.

Slight differences in environmental preferences exist
among Dominican Republic corbulid species. Corbula
sericea is most abundant in brackish-water and shallow-
marine to intermediate-marine deposits, although it is
conspicuously absent from sediments deposited in
grass-flat habitats such as the lower part of the Gurabo

| 20 т) to intermediate depths (approximately 30-

| iver)”, Whereas corbulids are much less common and

га Tse in Pliocene sediments deposited in deeper ma-

| е (> 100 m) waters (Text-figs. 2, 3; see Appendix

| ОГ criteria used to construct these paleoenvironmen-
Categories).

BULLETIN 351

o
Ф
=
-
€
Е
8
©
Ui
due 8
=
б |воот pee
% id
E ze
£ |
Е?
Ф о 4
а (9
Ф 2
©
9
а
500 m -
©
2 !
+2 ae
E | 400 m 17:
и
о eas "
8 [300m]? | 2
a =
T 5
Ф А o
5 200 m 4 |
Ф a
0 !
= А >
Е 4 !
E [100m d :
3 ЕЕ
с
о
5

= С sericea

Rio Gurabo Section

Coral Communities
Grass flats
ES] efs іп deeper, more
turbid waters (>10-20 m)
Т Transported corals

Environments

=] Brackish
= EA very shallow marine
9 (<30 т)
Е
Е Магі imatel
S |р arine (approximately
Ж 2 30 10 < 100 m)
с
3 Deeper marine (> 100 m)
©
|
2
8
е
4!
|

Text-figure 3.—Part of the Río Gurabo section showing the stratigraphic distributions of Corbula species. Coral communities аге Ба
average linkage cluster analysis of coral assemblages from Budd et al. (1996). Criteria used to distinguish environmental categories are OU

in Appendix 1.

Formation in the Río Cana section (Text-fig. 2; Ander-
son, 1994). Corbula viminea also occurs in brackish-
water and shallow-marine to intermediate-marine de-
posits in the study area, whereas C. sanctidominici ос-
curs primarily in shallow- to intermediate-marine sedi-

sed ОП
lined

ments (Text-figs. 2, 3). Corbula canae only occurs e
brackish-water deposits of the Río Cana section, whit
were probably deposited in or near a mangrove swam
(Text-fig. 2; Anderson, 1994). Corbula sericea and ©

ра ; : ‚тай
sanctidominici also occur in deeper marine sedime”

CORBULID BIVALVES: ANDERSON 9

С. sericea--Right Valves
3 в
а Deeper Marine
o Intermediate Do ig
24 в Shallow Marine a [5]
А Brackish E
о 0969 a

PC2: Valve Elongation
o

PC1: Size and Size-correlated Shape

Text-figure 4.—Plot of first and second principle component
Scores (PC1 and PC2) of right valves of Corbula sericea. Procedures
and results of the principal components analysis (PCA) are outlined
Ш Anderson (1994). PC1 (explaining about 55 % of the variance)
Tepresents size and size-correlated shape variability. PC2 (explaining
about 16 % of the variance) represents valve elongation, more elon-
gate valves have negative principle component scores. Criteria used
to distinguish environmental categories are outlined in Appendix 1.
See Appendix 2 for samples used for each category.

(>100 т deep) that show evidence of downslope
Movement, such as the top of the Río Gurabo section
(Text-fig. 3), the top of the Rio Yaque del Norte sec-
tion, and in the Cañada Zalaya section (Saunders е!
al., 1986; Anderson, 1994).

Intraspecific variability in the two most abundant
Corbulid species, C. sericea and C. viminea, are also
Telated to paleoenvironmental factors. Corbula sericea
Shows continuous morphologic variability along a pa-
leoenvironmental gradient of salinity, depth, and bio-
Clastic fabric (Text-fig. 4). The smallest and least elon-
8ate С. sericea valves are found in intermediate and
deeper marine deposits. Intermediate morphologies oc-
“ur in shallow-marine deposits, and larger (but not
More elongate) valves generally occur in brackish-wa-
ter deposits. A few valves categorized as “brackish-
Water" from the López section of the Río Yaque del

Orte, however, do not fit this trend (triangles with
Negative values on the first principle component axis
ІРСІ < 0] in Text-fig. 4). This anomaly is caused by
difficulties in assigning these sediments to an environ-
Mental category because of conflicting environmental
Nterpretations based on different faunal components.

ven though the ostracode and foraminiferal faunas of

е López section indicate deposition in brackish wa-

37 4
С. зепсеа B
| Right Valves
S Shallow Marine
se 3 u
% а Н. Мао 8 к
dE len
m 7 5 в S о 8 =
0- n Н n Cone кај в
Е u rr zi “У ЈА в
5 а 80%, m E pao i ï
T 26 a ge 5% * в.
ы So мора Oe a ват а
о = 00 о н E =
=
|. ia i * п mua =
E 1
в в
-3 Т Т Т T T T 1
3 2 -1 0 1 2 3

PC1: Size and Size-correlated Shape

Text-figure 5.—Plot of РСТ and PC2 of right valves of Corbula
sericea from shallow-marine deposits of the Río Cana and Río Mao
sections, where this species is locally abundant. Size and shape dif-
ferences are related to differences in bioclastic fabric. This plot
shows a subset of data illustrated in Text-figure 4. See Appendix 2
for samples used for each category. (After Anderson, 1994.)

ters (Saunders et al., 1986; Bold, 1988), these sedi-
ments also are rich in irregular echinoids and massive
reef corals (Saunders et al., 1986) indicating a signif-
icant marine influence. Alternatively, the other brack-
ish-water valves are from sediments with an unambig-
uous brackish-water signal. These sediments of the Río
Cana section are rich in brackish-water mollusks, in-
cluding beds of Anadara patricia (Sowerby, 1850) (—
“Arca” beds of Saunders et al., 1986). This species is
closely related to or is a subspecies of Anadara gran-
dis (Broderip and Sowerby, 1829), a Recent species
inhabiting intertidal mud banks bordering mangrove
swamps in the eastern Pacific (Maury, 1922; Olsson,
1932, 1961; Woodring, 1973; Saunders et al., 1986).
Geographic variation related to bioclastic fabric also
can be discerned within C. sericea from roughly con-
temporaneous Miocene sediments of the Río Cana and
Río Mao sections (Text-fig. 5; Anderson, 1994). Cor-
bula sericea is larger and more elongate in the Río
Cana section, where it occurs mainly in silts with bio-
clasts concentrated in lenses. In the Río Mao section,
C. sericea is smaller and less elongate, and occurs in
lenticular beds rich in bioclasts with a silty matrix.
Variation in C. viminea shows a morphologic cline
from west to east in contemporaneous sediments, with
variability related to sediment type and bioclastic fab-
ric (Text-fig. 6; Table 1; Anderson, 1994). Corbula vi-
minea is relatively small in the west (Río Cana section,
where it occurs in pebbly shell beds), whereas to the

BULLETIN 351

C. viminea

Bg 544]
8
5 Мао
Gurabo в
9 |
oz i
E Yaque
5 Cana | а
à | |
N 04 8
Ф |
о B
= о
©
Ф а А
E
a
а. - 4 Т Т T 1
Section
que
а.
©
с
Ф
о
| в
o
Р 8 0
Ф
ы 0- 8 i
Ф
Lo] 3 n
|--
G
Ф
IN. 27 B
[7] = А n
5
o с
-4 Т Т Т 1
1 2 3 4

Sediment Type

Text-figure 6.—Plot of PC1 versus stratigraphic section (upper) and PC1 versus sediment type (lower) for Corbula viminea. Procedures and
results of PCA are outlined in Anderson (1994). PC1 (explaining about 46 % of the variance) represents size and size-correlated shape
variability. Sediment types are: 1 = silts with bioclasts concentrated into lenticular beds or lenses, 2 = silts with bioclasts concentrated into
burrows, 3 = silts with bioclasts scattered, and 4 = silty sands with bioclasts either scattered or concentrated. Two valves from the Río Yaque
del Norte section are not included in the lower plot due to a lack of information on sediment type. Note clinal variation from west to east
(Río Cana to Río Mao section). Geographic variation is apparently related to differences in sediment type among sections (see Table 1). 5”

Appendix 2 for samples used for each category. (After Anderson, 1994.)

east (Río Gurabo and Río Mao sections, where it oc-
curs in lenticular-, burrow-, or lens-shaped shell con-
centrations within silts) it becomes progressively
larger. Valves from the Río Yaque del Norte section
are from older deposits of the Baitoa Formation and
do not follow this trend. Morphological differences
among the four sediment-type/bioclastic-fabric cate-
gories are significant using the nonparametric Krus-
kal-Wallis analysis of variance (H=103, df=3, p <
0.005).

In summary, both abundance of particular corbulid
species within their stratigraphic ranges and total rang”
es within stratigraphic sections are controlled by P?
leoenvironmental factors. In addition, intraspecill
morphologic variability in C. sericea and C. vimine?
corresponds strongly with specific paleoenvironment@
conditions. Although aspects of intraspecific variabl”
ity in both species are related to bioclastic fabric, n
phonomic processes do not control the morpholog!¢
variability observed (Anderson, 1994). Morpholog!®

CORBULID BIVALVES: ANDERSON 11

Table 1.—Corbula viminea valves from various sediment types
for each stratigraphic section. Sediment type categories are based on
descriptions in Saunders et al. (1986) and on personal observation.
Categories are: 1) silts with bioclasts in lenticular beds or lenses, 2)
silts with bioclasts filling burrows, 3) silts with mollusks dispersed
in matrix, 4) silty sands with bioclasts dispersed or concentrated.
Samples used are listed in Appendix 2.

sediment type

section 1 2 3 4
Cana 14 0 98 0
Gurabo 88 11 0 0
Mao 44 0 0 0
Yaque! 0 0 8 15

1 С. . . . . .
"Hid valves from this section were not included in the analysis
есаизе sediment type was unknown.

Change within these species over time is relatively mi-
Dor or is a side effect of systematic changes in envi-
Tonmental conditions.

BIOGEOGRAPHY

In tropical America (including the Caribbean Sea,
eastern Pacific Ocean, and western Atlantic Ocean),
Miocene to Recent corbulid faunas are nearly identical
at the subgenus level, except for a few apparently en-
demic, low-diversity taxa. Widespread and abundant
Subgenera include Varicorbula Grant and Gale, 1931
(Notocorbula Iredale, 1930 of some authors) and Car-
Jocorbula Gardner, 1926, and to a lesser extent Both-
"ocorbula Gabb, 1873a and Juliacorbula Olsson and

arbison, 1953. Endemic subgenera typically inhabit
Ше Recent Panamic-Pacific province and include Ser-
'acorbula Olsson, 1961 and Tenuicorbula Olsson,
1932 (also found in the Tertiary of Peru, Venezuela,
and Trinidad). Panamicorbula Pilsbry, 1932 was first
described from the Recent Panamic-Pacific province,
but also is reported from the Miocene of the Pacific
Coast of Costa Rica (Punta Judas; Seyfried er al.,
1985), the Miocene Cercado Formation of the Domin-
‘can Republic (Anderson, 1991, 1994), and the Plio-
cene l'Enfer and Springvale Formations of Trinidad
(see p. 19).

Dominican corbulid species show affinities to a
Number of other Miocene to Pleistocene faunas. The
[селе Bowden Formation of Jamaica shares two
е with the Dominican fauna, Corbula viminea
E C. sericea, and contains a species of Varicorbula,
1% bula (Varicorbula) heterogena Guppy (іп Dall,
tid 8), that is morphologically very similar to C. sanc-
a Өтіпісі. Corbula viminea and С. sericea also аге
Ed from Neogene deposits of the Limón Basin,

9sta Rica (Dall, 1898; Olsson, 1922). In addition, the
proving units all contain a number of corbulid spe-

es with strong affinities to Dominican corbulids:

Miocene and Pliocene sediments of Trinidad (includ-
ing the Manzanilla, Springvale, l'Enfer Formations;
Maury, 1925; Jung, 1969), the Miocene Gatun For-
mation of Panama (Woodring, 1982), Upper Miocene
Santa Rosa beds of Veracruz, Mexico (Perrilliat
[1984], who assigns these beds to the Agueguexquite
Formation; E. Vokes [1989] states, however, that this
formation only occurs farther north, near Coatzacoal-
cos, and is Pliocene), and Miocene to Pleistocene sed-
iments of Florida (Dall, 1898; Gardner, 1928; Olsson
and Harbison, 1953).

ABBREVIATIONS OF REPOSITORY
INSTITUTIONS

The following abbreviations for repository institu-
tions are used in this paper:

ANSP: Academy of Natural Sciences, Philadelphia,
PA, U.S.A.

IGM: Instituto de Geología, Ciudad Universitaria
de México, D.E, México

BMNH: The Natural History Museum, London, En-
gland, U.K. (British Museum, Natural His-

tory)

NMB: Naturhistorisches Museum Basel, Switzer-
land

PRI: Paleontological Research Institution, Ithaca,
NY, U.S.A.

TU: Tulane University, New Orleans, LA, U.S.A.

USNM: United States National Museum of Natural
History, Washington, DC, U.S.A.

UW: Museum of Geology, University of Wiscon-
sin-Madison, Madison, WI, U.S.A.

SYSTEMATIC PALEONTOLOGY
INTRODUCTION

All species described here are assigned to the genus
Corbula; other supraspecific taxa are considered sub-
genera, following Vaught (1989). All subgenera share
the same basic hinge structure and differ primarily in
shape, ornamentation, and the degree of disparity in
size, shape, and ornamentation of the left and right
valves. A majority of species described here are abun-
dant enough to incorporate intraspecific variation into
interpretations of species boundaries (see also Ander-
son, 1991, 1994 for quantitative treatments). Variabil-
ity is considered intraspecific if it can be related to
size differences (presumably ontogenetic) and/or to
changes in paleoenvironmental conditions. In compar-
ison to this approach, the taxonomy of living corbulids
has been more typological because taxonomic and sys-
tematic studies have not incorporated potential phe-
notypic variation caused by ontogenetic, environmen-
tal, or geographic factors.

Only forms considered identical to described species
are listed in the synonymy. A question mark (?) before
an item in a synonymy indicates that material was in-
sufficient to confirm conspecific status. The abbrevia-
tion “sp.” indicates that the described species could
be identified to the subgenus but not to the specific
level. Diagnoses are used both for species and for su-
praspecific categories and indicate the diagnostic fea-
tures of a particular taxon. Descriptions are used for
species only, and outline overall species morphology.
The Remarks section provides information on the par-
ticular taxon described, and may include a taxonomic
discussion, taphonomic and paleoenvironmental infor-
mation, and geologic and geographic ranges (ranges
for supraspecific taxa only). The Comparisons section
outlines differences between the species being de-
scribed and morphologically similar species. Detailed
locality information for Dominican Republic samples
is listed in the Occurrence section, whereas the Dis-
tribution section is a more general statement of a spe-
cies’ geographic and stratigraphic distribution inside
and outside the Dominican Republic. A question mark
(?) after a formation indicates that synonymy is un-
certain.

SYSTEMATICS
Family CORBULIDAE Lamarck, 1818

Diagnosis.—Small- to moderate-sized sturdy shells
(length typically 2 cm or less), inequilateral, and gen-
erally strongly inflated. Slightly to strongly inequival-
ved in size and shape; left valves smaller than right
valves. Hinge simple with anterior cardinal tooth in
right valve and socket in left valve. Resilifer present
behind hinge; typically a projecting chondrophore in
left valve that corresponds to socket-like resilifer in
right valve. Pallial sinus small to obsolete. (See H.
Vokes, 1945; Moore, 1969.)

Subfamily CORBULINAE Lamarck, 1818

Diagnosis.—Left valve typically slightly smaller
than right valve. Chondrophore in left valve present in
most species, fitting into socket-like resilifer in right
valve. Posterior typically rostrate. (See H. Vokes,
1945; Moore, 1969.)

Genus CORBULA Bruguiére, 1797

Corbula Bruguière, 1797, pl. 230.
Corbula Lamarck, 1799, p. 89.
Aloidis Megerle von Mühlfeldt, 1811, р; 07.

Type species. —Corbula sulcata Lamarck, 1801; by
subsequent designation, Schmidt ( 1818); Recent, Sen-
egal.

Diagnosis.—Shell sturdy, moderately inflated, ine-

BULLETIN 351

quilateral and inequivalved; left valve smaller. Shells
smooth to concentrically ribbed. Hinge with cardinal
tooth in right valve and cardinal socket in left valve.
In type species, socket-like resilifer present behind
dentition in both valves; in most species, however, left
valve with chondrophore and right valve with socket-
like resilifer. (See H. Vokes, 1945.)

Remarks.—The generic name Corbula was first
used in Вгиршеге (1797) in a plate title, and the name
is generally credited to him, although he did not de-
scribe the genus. Lamarck (1799) first described Cor-
bula but did not specify a type. Schmidt (1818) sub-
sequently designated Corbula sulcata as type (set
Stewart, 1930). Aloidis Megerle von Mühlfeldt (1811)
is a synonym of Corbula (H. Vokes, 1945, 1980). Un-
like most other members of the genus, the type species,
Corbula (Corbula) sulcata, has no chondrophore in
the left valve; the resilium instead is received in 4
socket-like resilifer behind the cardinal socket.

Geologic range of the genus is Cretaceous to Recent
(Moore, 1969). The genus is found worldwide in trop”
ical to temperate waters that range from marine (0
brackish salinities (Boss, 1982). Some species are tol-
erant of waters low in dissolved oxygen (Lewy an
Samtleben, 1979).

Subgenus BOTHROCORBULA Gabb, 1873a
Bothrocorbula Gabb, 1873a, p. 274, pl. 10, figs. 3, 3a.

Туре species.—Corbula viminea Guppy, 18666; БУ
monotypy. Pliocene, Bowden Formation, Jamaica.

Diagnosis. —Shell moderately large and thickened.
Valves subequal, with coarse concentric ribs and fine
radial striations. Characteristic deep lunular depressio”
in front of umbos. Right valve hinge with cardina
tooth in front of socket-like resilifer. Left valve hing?
with cardinal socket in front of chondrophore. (See
Gabb, 1873a; Gardner, 1926; H. Vokes, 1945.)

Remarks.—This subgenus ranges from Miocene 10
Pleistocene in Florida, the West Indies, and eastern
Central America (see p. 13-14). It is found in sedi-
ments deposited in shallow-marine and marginal-M%
rine waters. The Miocene-to-Recent taxon Нехасо“
bula Olsson, 1932 closely resembles Bothrocorbul@
but lacks а lunular pit.

Corbula (Bothrocorbula) viminea Guppy, 1866b
Plate 1, figures 1-8, 10, 11, 13, 14

2,
Corbula viminea Guppy, 1866b, p. 293, pl. 18, fig. 11; Olsson, 192

р- 270, pl. 28, fig. 25.

Corbula (Bothrocorbula) viminea Guppy. Maury, 1917, P-
234, pl. 39, figs. 20, 21; 1925, p. 108-109, pl. 19, fig. 19; 42
bry, 1922, p. 428; Woodring, 1925, р. 189-190, pl. 26, figs: р
8; Ramírez, 1950, р. 38-39, pl. 7, бр. 9; Anderson, 1994, fig
2.1-2.4.

233-
pils-

CORBULID BIVALVES: ANDERSON 13

Diagnosis.—Species characterized by relatively
large size, prominent lunule, thick valves, coarse con-
Centric ribs, fine radial striations, evenly rounded an-
terior margin, and evenly rounded to slightly sinuous
Ventral margin.

Description.—Valves moderately large, subequal in
Size and shape. Right valve slightly larger than left,
Only overlapping left ventrally. Valve thickness widely
Variable, although variability in part preservational be-
Cause valve inner layers tend to slough off. Valves
elongate-ovate with rostrum and arcuate keel. On some
Valves, posterior slope has midline depression parallel
to keel. Continuous variation seen in expression of ros-
tum and ventral margin, ranging from subdued ros-
trum and evenly rounded ventral margin, to short pos-
teriorly-pointing rostrum and evenly rounded ventral
margin, to short ventrally-pointing rostrum and gently
бош ventral margin (concave just anterior of ros-
A Valve ornamented with coarse concentric ribs

M die out before reaching keel. Fine concentric and
тайа] striations superimposed on coarse ribs, and es-
Pecially visible between ribs. Posterior slope and ne-
B ic shell with fine concentric striations, but lacking
adial and coarse concentric ribs.
pon valves with deep lunule anterior of umbo. Lu-
re larger and deeper in left valve, encompassing en-

hinge plate. Right valve hinge with large, trian-
i hooked-shaped, cardinal tooth and posterior
et-like, broadly-open resilifer. Broad short ridge
pus from resilifer roof and extends across part of
nge plate. Hinge plate surface depressed at resilifer.
ЗА valve hinge with deep, triangular, hook-shaped

„епог socket and broad posterior chondrophore.
pos plate strongly sinuous resulting in obliquely-

E cardinal socket. Dorsal surface of chondrop-
Eun anterior and posterior ridges separated by

ii ae trough. Posterior ridge more prominent and
Bis pues Adductor muscle scars relatively large
ate Ickened; posterior scar circular, anterior scar

- Pallial sinus obsolete.
eig Material. —Syntypes: BMNH 64088 (left
D. я BMNH 64099 (right valve). Right valve fig-
ate 6 y Guppy (1866b, Pl. 18, fig. 11) Both valves

T gured here (Pl. 1, figs. 3, 6).

зен (осаййу.--“Міюсепе” (= Bowden Formation,
1 LE Jamaica. The type locality is here restricted
беге) ET type locality of Bowden Formation (Plio-

que owden, Parish of St. Thomas, Jamaica (fide H.

8, 1989),
N p 1141. Меавигей and/or figured specimens:
G 14103, 14104, 14105, 14106, 14107, 14108,
» 14110. Other specimens (over 600 valves) аге
E am by locality, which are listed in the Occur-
Section,

14100
Catalo

Table 2.—Measurements of Corbula (Bothrocorbula) viminea.
Figures are in mm.

valve length height width

BMNH 64088: syntype (Pl. 1, fig.

3) right 19.1 150,63
BMNH 64099: syntype (Pl. 1, fig.

6) left 15.1 ДО 0
USNM 115648: (figured by

Woodring, 1925) right ef) 120- 48
PRI 919: (figured by Maury,

1917) left 14.0 96 . 41
PRI 919: (figured by Maury,

1917) right 1220 106 4.8
ММВ С 14103: (PL 1, fig. 14);

NMB locality 16923 left 15.9 118 49
NMB С 14104: (Pl. 1, fig. 13);

NMB locality 16923 right 17.4 Јана раб
NMB G 14105: (Pl. 1, fig. 11);

NMB locality 15900 left 13.3 11,3 5.4
NMB G 14106: (Pl. 1, fig. 10);

NMB locality 15900 right 16.3 io "бир
ММВ С 14107: (РІ. 1, fig. 8); TU

locality 1230 left 14.1 9.7 5.0
NMB G 14108: (Pl. 1, fig. 7); TU

locality 1230 right 14.2 Sy л
NMB G 14109: (Pl. 1, figs. 2, 5);

TU locality 1364 left I3: 9.3 3.6

NMB С 14110: (PL 1, figs. 1, 4);

TU locality 1364 right 14.0 9.7 4.8

Remarks.—This is а common species in upper Mio-
cene shallow-marine and brackish-water deposits of
the Río Cana, Río Gurabo, Río Mao, Río Yaque del
Norte, and Río Amina sections. It is typically found
in silty shell beds or thin shell-rich stringers interbed-
ded with silts. This species also is found in bioclast-
rich sediment filling burrows, scattered in silts, and in
pebbly and conglomeratic layers. Woodring (1925)
states that Dominican forms of C. viminea are smaller
and slightly more elongate than Jamaican forms.

Comparison.—Corbula (Bothrocorbula) wilcoxii
Dall, 1898, a Pleistocene species from the Caloosa-
hatchee and Bermont formations of southern Florida
(Anderson, unpublished data), shows some morpho-
logic overlap with С. viminea. Corbula viminea tends
to be slightly larger and has a more evenly rounded
anterior margin than С. wilcoxii. The valve margin of
C. wilcoxii typically is flattened into a short horizontal
ledge anterior of the beak, resulting in a projecting
anterior, Although the two species typically are dis-
tinct, intermediate forms can be found in both species.

Corbula (Bothrocorbula) synarmostes Dall, 1898 of
the Miocene Chipola Formation of Florida is smaller
and has a much smaller lunule than C. viminea. Cor-
bula (Bothrocorbula) radiatula Dall, 1898 of the Mio-
cene Oak Grove Formation of Florida is also smaller

14 BULLETIN 351

than C. viminea, has a reduced lunule, and more pro-
nounced radial ornament.

Occurrence.—This species was collected from the
following areas (see Saunders et al., 1986 for locality
information):

Río Cana: Cercado Formation: TU 1230, and NMB
16835, 16836, 16837, 16838, 16839, 16844, 16857,
16988, 16989, 16993, 17005. Lower Gurabo Forma-
tion: NMB 16831, 16832, 16833, 16820. Upper Gur-
abo Formation: TU 1354, and NMB 16817-16819,
16824. Mao Formation (Mao Adentro Limestone
Member): NMB 16873.

Río Gurabo: Cercado Formation: TU 1359, 1373,
1377, 1419, and NMB 15896, 15900, 15901, 15903,
15904, 15906, 15907, 15908, 15910. Lower Gurabo
Formation: TU 1297, 1298, and NMB 15876, 15878,
15882, 15887.

Río Mao: Cercado Formation: TU 1294, and NMB
16913 (all correspond to Maury’s Bluff 3); NMB
16915, 16917, 16918, 16923, 16924, 16926, 16927,
16928 (all Arroyo Bajón); NMB 16914, 16930, 16932
(all Maury’s Bluff 2).

Río Amina: Gurabo Formation: TU 1412.

Río Yaque del Norte: Baitoa Formation: TU 1226,
1363, 1364, and NMB 16935, 16936, 16938, 17286,
17288, 17289 (all López section). Unnamed forma-
tion: TU 1445 (Angostura).

Distribution.—Upper Lower to lower Middle Mio-
cene Baitoa, Upper Miocene Cercado, Upper Miocene
to Lower Pliocene Gurabo, and Lower Pliocene Mao
Formations, Dominican Republic; Miocene Thomonde
and Las Canobas Formations (?), Haiti (see Guppy,
1876; Woodring et al., 1924; Woodring, 1925); Plio-
cene Bowden Formation, Jamaica; Pliocene Río Ban-
ano Formation, Costa Rica.

Subgenus CARYOCORBULA Gardner, 1926

Caryocorbula Gardner, 1926, p. 46.

Type species.—Corbula alabamiensis Lea, 1833, by
original designation. Middle Eocene (Claibornian
Stage) of Alabama.

Diagnosis.—Small- to moderately-sized, moderate-
ly thickened valves. Left valve slightly smaller than
right. Shells typically elongate, with posterior keel and
rostrum. Valves with moderately coarse concentric
ribs; some also with fine radial striations. Right valve
hinge with anterior cardinal tooth and posterior socket-
like resilifer. Left valve hinge with anterior cardinal
socket and posterior chondrophore. (See Gardner,
1926.)

Remarks.—Caryocorbula includes most American
species assigned to Cuneocorbula Cossmann, 1886 by
a number of authors (e.g., Dall, 1898; Maury, 1917).

Its geologic range is Eocene to Recent in North and
South America and East Asia (Moore, 1969). Cary-
ocorbula species inhabit shallow-marine to marginal-
marine environments.

Corbula (Caryocorbula) dominicensis Gabb, 1873b
Plate 1, figures 9, 12; Plate 2, figures 1, 2, 4, 5
Corbula dominicensis Gabb, 1873b, p. 247; Pilsbry, 1922, p. 427,

р!. 46, брз. 12, 13.

?Corbula (Cuneocorbula) dominicensis Gabb. Maury, 1917, p. 232,

pl. 39, figs. 14, 15.

Diagnosis.—Species characterized by elongate
shape and sculpture of concentric, closely-spaced,
moderately coarse ribs and no radial striations. Also
distinguished from other Dominican corbulids by rel-
atively uninflated valves, moderately thickened shell,
and large size.

Description. —Right and left valves subequal іп size
and shape; right valve slightly larger than left. Umbos
slightly anterior of valve midline. Greatest convexity
of ventral margin anterior of umbo in left valve but
even with or anterior of umbo in right valve. Valves
not strongly inflated and shells not greatly thickened.
Sculpture consists of concentric, closely-spaced, flat
topped ribs with steep dorsal and more gradual ventral
slopes. Ribs do not split and double towards posterior
keel as described by Pilsbry (1922). Radial sculpture
absent. Nepionic shell somewhat distinct, with very
fine concentric ribs. Posterior region of valve W!
sharp arcuate keel and small rostrum. Left valve hinge
with anterior, large, triangular socket that opens ven-
trally. Broad chondrophore posterior of cardinal 500%”
et, continuous with dorsal margin, and with midlin®
cleft. Right valve hinge with anterior, large, hook-
shaped, triangular cardinal tooth. Widely open socket-
like resilifer present posterior of cardinal tooth. 1,008
furrows for reception of left valve located anterior an
posterior of, and continuous with, right valve hinge
Adductor muscle scars large and thickened; posterior
scar circular, anterior scar ovate. Very small pallial 51%
nus present.

Type Material.—Lectotype: ANSP 2691 (articulated
shell). Specimens figured by Pilsbry (1922, p. 427, р:
46, figs. 12, 13), who designated the lectotype, whic
is now lost (G. Rosenberg, pers. comm., 1993). ,

Type locality.—None designated. It is here desig-
nated as NMB 17281: Baitoa Formation (upper Lowe!
to lower Middle Miocene), López section, Rio Yaque
del Norte, Dominican Republic. |

Material.—Measured and/or figured specimen"
ММВ С 14111, 14112, 14113. Other specimens (104
right valves, two left valves, three articulated shells:
one internal mold, and two left valve fragments) Р a
cataloged by locality, which are listed in the Oceuf
rence section.

u

CORBULID BIVALVES: ANDERSON 15

| Table 3.—Measurements of Corbula (Caryocorbula) dominicen-
sis. Figures аге іп mm.

specimen valve length height width

ANSP 2691: lectotype (figured

ре Pilsbry, 1922) articulated 14.6 90 6.0!
29033: (figured by Maury,
р 1917; refigured in PL. 2, fig. 3) left 134586 83
КІ 29033: (figured by Maury,
1917; refigured in Pl. 2, fig. 6) right 126 86 32
NMB С 14112: (Pl. 2, figs. 2,
5); TU locality 1226 left BO Sa 51
NMB G 14113: ІРІ. 2, figs. 1,
4); TU locality 1226 right 142 90 40

ММВ G 14111: (РІ. 1, figs. 9,
12); NMB locality 17281

m
diameter of articulated shell.

articulated 15.3 494. GM

Measurements.—See Table 3.

Remarks.—This is a rare species found in conglom-
ү» ог conglomeratic lenses containing mollusks. In
на Study, it was found only in the upper Lower to

er Middle Miocene Baitoa Formation of the Rio

aque del Norte section (see Saunders е! al., 1986).
^ p" (1917), however, reports C. dominicensis from

Upper Miocene Cercado Formation of the Río Cana
ке» (Zone Н of the Río Cana at Caimito; equiva-
ао TU 1230 [Saunders et ns 1986; H. Vokes,
5) p Maury's specimens (refigured in РІ. er figs. By
er from the Baitoa Formation specimens in sev-

Ways. Maury’s specimens are smaller, less elon-
8ate, have a more centrally located umbo, more strong-
иар keel, larger posterior slope, and coarser con-
ш ribs. Smaller specimens from the Baitoa For-

n closely resemble Maury's specimens, but with
үзіне material available, it was not possible to
Пета ог separate the specimens from the two forma-

Comparison.—Corbula (Caryocorbula) democracia
за Odson, in Hodson and Hodson, 1931, from the
nl of Falcón, Venezuela is much larger (holo-
E gn is 22.5 mm), is less elongate, has a more

X ventral margin, and has coarser concentric ribs
E dominicensis. Corbula (Caryocorbula) pren-
E Olsson, 1964 of the lower and middle Gatun
E (Upper Miocene) of Panama and the Upper
Пе Angostura Formation of Ecuador is about the
E" as C. dominicensis but is more strongly ros-
^ En the rostrum located in a more dorsal position
cuate omen margin, and has a more strongly ar-
een giving the rostrum a twisted appearance.
brine (Caryocorbula) dominicensis veracruzana
beds E 1984 from the Upper Miocene Santa Rosa

Bos eracruz, Mexico more closely resembles C. de-

b ста than C. dominicensis s. s., and probably 15

а Subspecies of C. dominicensis.

Sa

Occurrence.—This species was collected from the
following areas (see Saunders ef al., 1986 for locality
information):

Río Yaque del Norte: Baitoa Formation: TU 1226,
and NMB 17281, 17283.

Distribution.—Upper Lower to lower Middle Mio-
cene Baitoa, Upper Miocene Cercado (?) Formations,
Dominican Republic.

Corbula (Caryocorbula) sericea Dall, 1898
Plate 2, figures 7-21

Corbula lavaleana Orbigny. Gabb, 1873b, p. 247; 1881, p. 371.

Corbula (Corbula) sericea Dall, 1898, p. 848-849; 1900, pl. 36, fig.
8; Woodring, 1925, p. 186-187, pl. 25, figs 19-22.

Corbula (Cuneocorbula) cercadica Maury, 1917, p. 232-233, pl.
39, figs. 16, 17.

Corbula (Cuneocorbula) caimitica Maury, 1917, p. 233, pl. 39, figs.
18; 19,

Corbula sericea Dall. Pilsbry, 1922, p. 427.

Corbula (Caryocorbula) cercadica Maury. Anderson, 1994, figs.
1.1-1.5.

Diagnosis.—Species characterized by small, inflat-
ed, elongate-ovate to subtriangular valves with rela-
tively fine, closely and evenly spaced concentric ribs,
and very fine radial striations.

Description.—Shells small to moderately sized;
moderately inflated. Valves thin to relatively thick,
variability in part preservational. Left and right valves
subequal in size and shape; right valve larger and
slightly less elongate than left. Umbo at or slightly
anterior of valve midline. Keel nearly straight to ar-
cuate. Valve shape varies from triangular to elongate
ovate; elongate valves more rostrate. Ornament of
closely spaced concentric ribs. Fine radial ribs present
but variably expressed. Radial striations present, bead-
ed under magnification.

Right valve hinge with anterior, large, triangular,
hook-shaped, cardinal tooth, and posterior, large, wide-
ly open, socket-like resilifer. Ridge bisects roof of re-
silifer, as in С. viminea. Long furrow present on either
side of right valve hinge for reception of left valve’s
thickened dorsal margin. Posterior furrow continuous
with resilifer. Left valve hinge with anterior, large, tri-
angular, cardinal socket and broad posterior chondrop-
hore. Anterior half of chondrophore's dorsal surface
concave, posterior half convex. Small denticle present
at posterior edge of chondrophore. Adductor muscle
scars slightly thickened. Posterior scar circular; ante-
rior scar ovate. Pallial sinus small to obsolete.

Type Material. —Lectotype: USNM 135655 (right
valve). Lectotype designated by Woodring (1925).
Specimen figured by Dall (1900, Pl. 36, fig. 8) and
Woodring (1925, Pl. 25, figs. 19, 20), and refigured
here (Pl. 2, fig. 7).

Type locality.—USGS locality 2692: track ballast

16 BULLETIN 351

Table 4.—Measurements of Corbula (Caryocorbula) sericea.
Figures are in mm.

specimen valve length height width

USNM 135655: lectotype (Pl. 2,
ng. 7) right
PRI 29035": (figured by Maury,
1917; refigured in Pl. 2, fig. 9)
PRI 29035!: (figured by Maury,
1917; refigured in Pl. 2, fig. 8) right 6.6 4.6 2.3
ММВ С 14114: (РІ. 2, бр. 17);
ММВ locality 16928 left 20 4.0 1.8
ММВ С 14115: (Pl. 2, fig. 16);
ММВ locality 16928
NMB G 14116: (Pl. 2, fig. 19);

5.3 4.0 2.0

left 5.6 319 2.0

right 6.1 4.5 2.1

NMB locality 15869 left 4.2 2.9 1.4
ММВ С 14117: (Pl. 2, fig. 18);

NMB locality 15869 right 4.9 2,2 1.6
NMB С 14118: (Pl. 2, figs. 11,

15); NMB locality 16848 left a) 4.7 2.4

NMB G 14119: (Pl. 2, figs. 10,
14); NMB locality 16848
NMB С 14120: (Pl. 2, fig. 13);

right 7,0 5.0 3.0

NMB locality 16855 left gi 4.6 2:3
NMB G 14121: (Pl. 2, fig. 12);

NMB locality 16855 right 8.1 4.8 Zul.
NMB G 14122: (Pl. 2, fig. 21);

TU locality 1227A left 2,0 3.9 2.0

NMB С 14123: (Pl. 2, fig. 20);

TU locality 1227A right 6.0 4.2 2,9

! Mislabeled as cotypes (syntypes) of Corbula (Cuneocorbula) caim-
шса but are Corbula (Cuneocorbula) cercadica illustrated in pl. 39,
figs. 16, 17 of Maury 1917). The syntypes of Corbula (Cuneocor-
bula) caimitica are apparently lost (W. Allmon, pers. comm., 1994),

for railroad 1.5 miles west of Limón, Costa Rica. The
source of the aggregate is USGS locality 2694: in situ
Limón Reef (= Moín Formation, Pliocene).

Material.—Measured and/or figured specimens:
NMB G 14114, 14115, 14116, 14117, 14118, 14119,
14120, 14121, 14122, 14123. Additional material: UW
1863/25, 1863/26. Other specimens (thousands of
valves) are cataloged by locality, which are listed in
the Occurrence section.

Measurements.—See Table 4.

Remarks.—This species is locally abundant in Up-
per Miocene shallow-marine (< 30 m) and brackish-
water deposits of the Río Cana, Río Gurabo and Río
Mao sections. It also occurs in Miocene and Pliocene
intermediate and deeper marine (> 30 to + 100 m)
sediments of the Río Cana, Río Gurabo, Cañada Za-
laya, Río Amina, Río Verde, and Río Yaque del Norte
sections. Corbula sericea is found in a variety of sed-
iment types in the study area although it predominantly
occurs scattered in silts and in shell beds with a silty
matrix.

Corbula sericea is morphologically variable, al-
though most shape variation is highly correlated with
size. Size and size-correlated shape, in turn, are con-

tinuous and are closely related to paleoenvironmental
conditions (Anderson, 1994). Specimens from very
shallow marine deposits of the Rio Mao have a sub-
triangular valve outline (Pl. 2, figs. 16, 17). Average
height to length ratio (H:L) is 0.72 (all height to length
ratios for C. sericea reported here are based on data
from Anderson, 1994). The ventral margin of these
forms is directed upward at a slight angle in both di-
rections from the midline. There is no invagination of
the ventral margin in association with the keel or ros-
trum. The rostrum is very subdued to nearly absent
and is directed downward. Most forms һауе an evenly
rounded dorsal margin, although in some, the umbo is
somewhat set off and projecting.

Specimens from very shallow marine sediments of
the Río Cana section (Pl. 2, figs. 12, 13) tend to be
larger and more elongate (average H:L = 0.63) than
those of the Río Mao section. In addition, the rostruM
in Río Cana specimens is located in a more dorsal
position on the posterior margin, and it tends to point
posteriorly rather than ventrally. As a result, valves
have an ovate outline. Convexity of the keel varies
but most specimens have a slightly sinuous keel. The
dorsoanterior slope is nearly straight, and the highest
point of the valve is located more anteriorly than mn
the Río Mao specimens.

Brackish water forms from the Río Cana section (Pl
2, figs. 10, 11, 14, 15) also are large, ovate, and elon-
gate (average H:L = 0.68), although not as elongate
as valves from very shallow-marine deposits of Ше
Río Cana. In addition, the dorsoanterior slope is more
rounded than in very shallow-marine forms. The ro"
trum is moderately expressed to obsolete, and when
present, is directed ventrally. The ventral margin је
strongly rounded, and the keel less sinuous than 1”
shallow-marine forms of the Río Cana section.

Valves from intermediate and deeper marine de-
posits of the Río Gurabo, Río Mao, Río Cana, an
Río Yaque del Norte sections tend to be smaller thar
those of other paleoenvironments but are very simila
in shape to the Río Mao specimens. For these deep?
water forms (Pl. 2, figs. 18, 19), H:L is 0.72 to 0.!
on average. Pliocene valves of deeper marine deposit?
of the Cañada Zalaya section (Pl. 2, figs. 20, 21) 8“
similar in both size and shape to other deeper water
forms, except that the ventral margin tends to be fiat
ter.

Comparison.—Jung (1969) noted that small сагу,
ocorbulids of the Miocene and Pliocene of Trin! A
were oversplit and considered Corbula smithian
Maury, 1912, and possibly С. caribaea per&'^ |
Maury, 1925 and C. daphnis Maury, 1925, to ђе J"
nior synonyms of C. helenae Maury, 1912. Althouß
it is similar in shape to Rio Cana morphologies оғ

CORBULID BIVALVES: ANDERSON 17

зегісеа, С. helenae has a larger maximum size (up to
13 mm long), is more elongate, is less inflated, can
Бе more coarsely sculptured, can have more promi-
nent radial striations, and tends to have a more bul-
bous and projecting anterior. The species C. manzan-
illensis Maury, 1925 of the Miocene Manzanilla For-
Mation of Trinidad is very small, has coarser ribs at
Comparable sizes, is strongly triangular, and less elon-
gate than C. sericea.

Corbula (Caryocorbula) oropendula Olsson, 1922
has coarser sculpture, has a straighter keel and more
ventrally located rostrum, is more elongate, and has
à more strongly and evenly rounded ventral margin
than C. sericea. Corbula (Caryocorbula) oropendula
dolicha Woodring, 1982 is similar in shape to C. or-
OPendula but has finer sculpture. Corbula (Caryocor-
bula) oropendula stena Woodring (1982) is much
More rounded, tends to be smaller, and its rostrum
Creates a more prominent notch in the ventral margin
than seen in C. sericea.

Occurrence.—This species was collected from the
following areas (see Saunders et al., 1986 for locality
Information):

Río Cana: Cercado Formation: TU 1230, and NMB
16838, 16839, 16841, 16843, 16844, 16845, 16846,
16848, 16850, 16851, 16853, 16854, 16855, 16856,
16986, 16987, 16988, 16989, 16990, 16993, 17001,
17003. Lower Gurabo Formation: NMB 16832. Up-
Per Gurabo Formation: TU 1354, and NMB 16865,
17009.

B Gurabo: Cercado Formation: TU 1277, 1419,
ума 15900, 15904, 15910, 15911, 15912,
ү Lower Gurabo Formation: TU 1211, 1215,
Шақ апа ММВ 15842, 15846, 15854, 15860, 15863,
n 15869, 15871, 15873, 15882, 15936, 15937,
EM 15944, 15945, 15947, 16808, 16809, 16810,
m 15835, 15952. Upper Gurabo Formation: TU
1596, and NMB 15933, 15935, 15939, 15964, 15966,
9, 15805, 15804, 15814, 15815. Mao Formation:
U 1352, and NMB 15822, 15827, 15832, 15833.
p Mao: Cercado Formation: TU 1294, and NMB
2, 16913, 17269 (all correspond to Maury's
E (1917)); NMB 16915, 16916, 16917, 16918,
a 22, 16923, 16924, 16926, 16927, 16928 (all Ar-
1694 Bajón); NMB 16914, 16929, 16930, 16931,
Зи 2 (all Maury's Bluff 2); NMB 16802 (located
En Bluff 1 and 2); TU 1293, and NMB 16910
ü luff 1), and TU 1225 (down stream and up sec-
on from Bluff 1).
E Amina: Gurabo Formation: TU 1219, 1411,
Cañada Zalaya: Gurabo Formation: TU 1227,
27A, 1453, 1453A.
Río Verde: Gurabo Formation: TU 1250.

12

Río Yaque del Norte: Baitoa Formation: TU 1226.
Unnamed Formation: NMB 17273 (Arroyo López),
NMB 17278 (Angostura). Gurabo Formation: TU
1403, 1405, 1448, 1449 (La Barranca), TU 1206
(Santiago).

Distribution.—Upper Lower to lower Middle Mio-
cene Baitoa, Upper Miocene Cercado, Upper Mio-
cene to Lower Pliocene Gurabo, and Lower Pliocene
Mao Formations, Dominican Republic; Pliocene
Bowden Formation, Jamaica; Pliocene Moín Forma-
tion, Costa Rica; Miocene Gatun Formation (7), Pan-
ama (see Brown and Pilsbry, 1911).

Subgenus JULIACORBULA
Olsson and Harbison, 1953

Juliacorbula Olsson and Harbison, 1953, p. 148-149.

Type species. —Corbula aequivalvis Philippi, 1836
(= Corbula cubaniana Orbigny, 1846; = Corbula
knoxiana Adams, 1852b); by original designation.
Recent, West Indies.

Diagnosis.—Shell small- to medium-sized, nearly
equivalved. Subrectangular with strong keel and
sharply truncated posterior. Well-defined escutcheon
located behind beak. Hinge as in Caryocorbula. (See
Olsson and Harbison, 1953; Olsson, 1961.)

Remarks.—Juliacorbula differs from Caryocorbu-
la in shape and in the presence of an escutcheon. The
type species was originally designated as C. cubani-
ana, which generally is agreed to be a junior syn-
onym of C. aequivalvis (see McLean, 1951; Weis-
bord, 1964; Rios, 1975). The geologic range of Ju-
liacorbula is Miocene to Recent in the West Indies,
eastern Pacific, Central and South America, and Flor-
ida (Moore, 1969; Olsson and Harbison, 1953). Mem-
bers of this group inhabit shallow-marine environ-
ments.

Corbula (Juliacorbula) fossilis Pilsbry, 1922
Plate 2, figures 22-26

Corbula contracta Say. Gabb, 1873b, p. 247.

Corbula knoxiana fossilis Pilsbry, 1922, p. 427, pl. 46, fig. 14.

2Corbula (Cuneocorbula) cubaniana Orbigny. Maury, 1925, p.
103-104, pl. 20, figs. 2-4.

2Juliacorbula aequivalvis (Philippi). Jung, 1969, p. 410-411, pl.
39, figs. 11-15.

Corbula (Juliacorbula) aequivalvis Philippi. Perrilliat, 1984, p. 17,
pl. 16, figs. 1-4.

Diagnosis.—Species characterized by straight to
slightly concave ventral margin, strongly angled pos-
terior margin, strong keel and rostrum, and subtrap-
ezoidal shape.

Description.—Valves relatively small, subtrapezo-
idal, and moderately inflated. Ventral margin flattened

18 BULLETIN 351

to slightly concave at midline. Dorsoanterior margin
also slightly concave in front of umbo. Directly pos-
terior of beak, valve margin planar, gently sloping
ventrally (sloping more steeply in left valves) to ros-
trum so that rostrum nearly as high as entire valve.
Posterior of rostrum, valve margin nearly planar and
vertical. Keel sharp and gently sinuous. Valve orna-
ment of relatively coarse ribs with steep dorsal and
gentle ventral slopes. Faint radial striations present on
umbo.

Right valve hinge with moderately projecting, tri-
angular, hook-shaped, cardinal tooth directly beneath
beak. Posterior socket-like resilifer present beneath
umbo. Hinge plate strongly sinuous, making resilifer
nearly obsolete. Posterior of hinge, long L-shaped
furrow present on right valve’s posterior margin for
reception of left valve. Left valve hinge with large,
triangular, cardinal socket beneath beak and posterior
broad chondrophore. Dorsal surface of chondrophore
with anterior, midline and posterior ridges; small den-
ticle present at end of posterior ridge. Adductor mus-
cle scars large and moderately thickened; anterior
scar ovate, posterior scar circular. Раша! sinus ob-
solete.

Type Material. —Holotype: ANSP 2689 (right
valve). Figure by Pilsbry (1922, Pl. 46, fig. 14) and
refigured here (Pl. 2, figs. 22, 23).

Type locality.—None designated. It is here desig-
nated as NMB 15914: Cercado Formation (Upper
Miocene), Río Gurabo, Dominican Republic.

Material. —Measured and/or figured specimens:
NMB G 14124, 14125.

Measurements.—See Table 5.

Remarks.—This rare species is found in shell-rich
sediments with a silty matrix.

Comparison.—Species of Juliacorbula from the
Tertiary of Trinidad (Maury, 1925; Jung, 1969) are

Table 5.—Measurements of Corbula (Juliacorbula) fossilis.
Figures are in mm.

specimen valve length height width
ANSP 2689: holotype (РІ. 2, figs.
22, 29) right 8.7 6.1 Ра
ANSP uncat.: paratype (Pl. 2, fig.
24) left 8.4 5.8 2.8

ANSP uncat.: paratype left 6.7 4.4 1.8
NMB G 14124: (Pl. 2, fig. 26);
NMB locality 15914 right PSI 4.1 1.6
NMB G 14125: (Pl. 2, fig. 25);
ММВ locality 16817
USNM 306431: (figured in Perril-
liat, 1984); IGM locality 2851 left S 6.8 2.2
РКІ 870: (бригед in Maury, 1925) left 6:2 4.3 1.9
РКІ 871: (figured іп Maury, 1925) гірім 7.4 24 #3
PRI 872: (figured in Maury, 1925) right 719 6.0 47]

right 2:0 = 1.4

very similar to the Dominican species, although the
posterior slope is nearly vertical in the Trinidad spec-
imens and more oblique in Dominican forms. It is not
possible at this time to determine whether this differ-
ence is of taxonomic significance because of the pau-
city of material. Corbula (Juliacorbula) scutata Gard-
ner, 1943 of the Florida Pleistocene is very similar 10
C. fossilis but has coarser ribs, a more arched and less
sinuous keel, a less concave dorsoanterior margin, and
can be more elongate. The Pleistocene to Recent С.
aequivalvis Philippi, 1836 (=C. knoxiana C. B. Ad-
ams, =C. cubaniana Orbigny) may be a peramorphic
descendent of C. fossilis. Smaller specimens of C. ае-
quivalvis overlap in morphology with C. fossilis. Cor-
bula aequivalvis differs from С. fossilis, however in
attaining a larger size, having a more rounded ventral
margin that shows its greatest convexity anterior of the
midline, not having a concave dorsoanterior margin,
being more elongate, having a more sinuous keel, and
tending to have finer ribs.

Occurrence.—This species was collected from the
following areas (see Saunders et al., 1986 for locality
information):

Río Cana: Upper Gurabo Formation: NMB 16817.

Río Gurabo: Cercado Formation: NMB 15914.

Distribution.—Upper Miocene Cercado, Upper
Miocene to Lower Pliocene Gurabo Formations, DO-
minican Republic; Upper Miocene Santa Rosa beds,
Veracruz, Mexico; Lower Pliocene Melajo Clay Mem-
ber of the Springvale Formation(?), Pliocene Point
Courbaril Sand and Clay Member of the 1'Enfer Рог
mation (2), Pliocene Matura Sand and Clay Member
of the Talparo Formation (?), Trinidad.

Subgenus PANAMICORBULA Pilsbry, 1932

Panamicorbula Pilsbry, 1932, p. 105.

Type species. —Potamomya inflata Adams, 18524
(=P. aequalis Adams, 1852a, =P. trigonalis Adams
1852a, = Corbula macdonaldi Dall, 1912); by original
designation. Recent, Pacific Coast of Panama.

Diagnosis.—Valves moderately large and inflat
but relatively thin, not rostrate. Surface smooth or wit
very fine concentric ribs. Hinge of right valve with
anterior cardinal tooth and posterior socket-like res!
lifer. Left valve hinge with anterior cardinal socket 20
posterior chondrophore. (See Pilsbry, 1932; H. Vokes»
1945; Olsson, 1961.)

Remarks.—Contrary to the original description,
lateral teeth are present. What Pilsbry (1932) describe
as long laterals in the right valve are actually buttresses
for grooves that receive the dorsal margin of the left
valve (H. Vokes, 1945; Moore, 1969). Living Pan
amicorbula are found in the Panamic Province (easter

ted

no

CORBULID BIVALVES: ANDERSON 19

Pacific; Mazatlán to Peru) in brackish water (Pilsbry,
1932; Olsson, 1961). The subgenus is reported as
abundant but poorly preserved from Middle Miocene
Sediments at Punta Judas, Pacific Coast, Costa Rica
(Seyfried et al., 1985). It is also present in USNM
Collections (with U.S. Geological Survey locality num-
bers) of the Lower Pliocene l’ Enfer Formation (USGS
21842, USGS 20433) and Lower Pliocene Springvale
Formation (USGS 20421, USGS 21083, USGS 20423)
Of Trinidad.

Corbula (Panamicorbula) canae, new species
Plate 3, figures 1-10

Corbula (Panamicorbula) aff. C. inflata (C. B. Adams). Anderson,
1994, fig. 3.1.

Diagnosis.—Species characterized by roughly tri-
Angular shape, thin valves, fine concentric ribs, and left
and right valves subequal in size and shape.

Description.—Valves relatively large (maximum
length about 15 mm), inflated, and thin. Left and right
Valves subequal in size and shape; right valve slightly
larger. Valve height shows positive allometry relative
lo valve length; larger valves more triangular, whereas
Smaller valves more quadrate. Ventral margin most
convex anterior of midline, whereas umbo located at
midline, Ventral margin of right valve rounded, of left
valve slightly sinuous. Valve ornament of very fine,
closely-spaced, concentric ribs. Radial ribs absent. Ar-
Cuate keel and subtle rostrum present. Right valve
hinge with large, anterior, triangular, hook-shaped, car-

Mal tooth. Posterior, socket-like resilifer present be-
Neath beak and opening ventrally. Elongate furrow
Present on both sides of right valve hinge for reception
of left valve. Left valve hinge with large, anterior, tri-
angular, cardinal socket and broad posterior chondro-
Phore, Chondrophore continuous with valve margin
and not strongly projecting. Low anterior, midline, and
жеры ridges present on dorsal surface of chon-
| оте. Adductor muscle scars large but not thick-
Bi ед. Posterior scar circular; anterior scar ovate. Pallial

hus obsolete.

Etymology of name.—Name after Río Cana.
| Туре Material.—Holotype: NMB С 14126 (articu-
ated shell). Figured in Plate 3, figures 1, 2.

Туре locality.—NMB 16845: Cercado Formation
d Miocene), Río Cana, Dominican Republic.
| Tatypes are from NMB 16841, a nearby locality in

© same lithologic unit.

Material.—Measured and/or figured specimens:
G 14126, 14127, 14128, 14129, 14130, 14131,
val 2. Other specimens (11 articulated shells, 32 right

ves, and 20 left valves) are cataloged by locality,

Ich are listed in the Occurrence section.

Table 6.—Measurements of Corbula (Panamicorbula) canae. Fig-
ures are in mm.

specimen valve length height width

ММВ С 14126: holotype (Pl. 3,

figs. 1, 2); NMB locality

16845 articulated 14.4 11.8 9.6!
ММВ С 14127: paratype (РІ. 3,

figs. 6, 7); NMB locality

16841 left 152 99 36
ММВ С 14128: paratype (РІ. 3,

fig. 8); ММВ locality 16841 left 135 104 43
ММВ С 14129: paratype (Pl. 3,

бе. 5); NMB locality 16841 right 15.9. liz «s

NMB G 14130: paratype (Pl. 3,
figs. 3, 4); NMB locality

16841 right 124 102 3.4
NMB G 14131: (PL 3, fig. 9);
NMB locality 16841 left 8.3 74052,2

ММВ С 14132: (РІ. 3, fig. 10);

NMB locality 16841 right T о Tio

! diameter of articulated shell.

Measurements.—See Table 6.

Remarks.—This is the first description of a fossil Pan-
amicorbula species and the first description of a Carib-
bean Panamicorbula. This species is restricted to the
* Arca" beds of the Upper Miocene Cercado Formation
Río Cana section (see Saunders et al., 1986). It is found
in interbedded shelly silts and silty clays, and in shell
beds. Living representatives of Panamicorbula are re-
stricted to the Pacific coast of Central America where
they inhabit mangrove swamps and areas near the
mouths of streams (Pilsbry, 1932; H. Vokes, 1945; Keen,
1971). Corbula canae apparently had similar environ-
mental preferences because it occurs in sediments de-
posited in or near mangrove swamps (Anderson, 1994).

Comparison.—Corbula (Panamicorbula) inflata (Ad-
ams, 1852a) (synonyms: C. aequalis [Adams, 18524], С.
trigonalis [Adams, 1852a], and C. macdonaldi Dall,
1912) is much larger than C. canae and differs somewhat
in shape. The ventral margin of C. inflata (Adams,
1852a) is flatter and is most convex near the valve mid-
line, the keel is less arcuate, and the rostrum is absent.

Occurrence.—This species was collected from the
following areas (see Saunders et al., 1986 for locality
information):

Río Cana: Cercado Formation: NMB 16843, 16845,
16990, 16841, 16993, 16987, 16989, 16840, 16844,
16852, and 16842.

Distribution.—Upper Miocene Cercado Formation,
Dominican Republic.

Corbula (Panamicorbula) sp.
Plate 3, figure 11

Diagnosis.—Small uninflated valves (— 6.5 mm) of
Panamicorbula with rectangular shape.

Table 7.—Measurements of Corbula (Panamicorbula) sp. Figures
are in mm.

specimen valve length height width
NMB С 14133: (Pl. 3, fig. 11);
NMB locality 16990 right 5.4 4.0 1.8

Material.—Measured and/or figured specimens:
NMB G 14113. Other specimens (15 right valves, and
4 left valves) are cataloged by locality, which are listed
in the Occurrence section.

Measurements.—See Table 7.

Remarks.—These are small, apparently immature
valves of a Panamicorbula species. Although these
valves may be small individuals of Corbula (Panam-
icorbula) canae, the small number of valves for com-
parison and lack of intermediates makes assignment
uncertain. These valves co-occur with C. canae in in-
terbedded shelly silts and silty clays, and shell beds
dominated by Anadara patricia in the upper part of
the Upper Miocene Cercado Formation of the Río
Cana section.

Comparison.—Valve shape of Corbula (Panamicor-
bula) sp. resembles Caryocorbula, but these valves are
not as inflated as Caryocorbula, and the resilifer of the
right valve opens ventrally (as in Panamicorbula).
These valves differ from Corbula (Panamicorbula)
canae in that they are small, much less inflated, and
rectangular (rather than triangular) in shape.

Occurrence.—This species was collected from the
following areas (see Saunders et al., 1986 for locality
information):

Río Cana: Cercado Formation: NMB 16845, 16990,
16993, 16987, 16989, 16846, 16852, 16986.

Distribution.—Upper Miocene Cercado Formation,
Dominican Republic.

Subgenus VARICORBULA Grant and Gale, 1931
Varicorbula Grant and Gale, 1931, p. 420, footnote 1.

Type species.—Tellina gibba Olivi, 1792; by orig-
inal designation. Recent, west coast of Europe and
Mediterranean Sea.

Diagnosis.—Valves small to moderate in size with
right and left valves strongly unequal in size, shape,
and ornamentation. Right valve larger, more inflated
and relatively higher; left valve smaller, more elongate,
and much less inflated. Right valves with strongly ex-
pressed concentric ribs. Left valves with fine, widely
spaced, radial ribs and fine concentric growth-lines;
thicker concentric ribs may be present on beak and
umbo. Right valve hinge with anterior cardinal tooth
and posterior socket-like resilifer. Left valve hinge
with anterior cardinal socket and posterior chondro-
phore.

BULLETIN 351

Remarks.—Some workers (Stenzel et al., 1957;
Weisbord, 1964; Jung, 1969) consider Varicorbula to
be a junior synonym of Notocorbula Iredale, 1930.
Woodring (1982), however, advocated continued use
of Varicorbula until the type species of Notocorbula
was better known. Stenzel et al. (1957) state that the
nepionic shells evident in both valves of the type spe-
cies, Corbula (Notocorbula) vicaria (Iredale, 1930),
can be seen in a more subdued form in С. gibba (Olivi,
1792), the type species of Varicorbula, and use this
trait as a basis for synonymy. Corbula vicaria, how-
ever, does apparently differ from C. gibba in that left
and right valves are subequal in size and shape, where-
as Varicorbula is strongly inequivalved. Therefore,
Varicorbula 15 used here.

The geologic range of Varicorbula is Еосепе to Re-
cent in eastern North America, Europe, and the eastern
and western Pacific (Moore, 1969). Varicorbula spe-
cies inhabit marine waters and can locally dominate
the benthic fauna (е.о., Yonge, 1946).

Corbula (Varicorbula) sanctidominici Maury, 1925
Plate 3, figures 12-18

Corbula disparilis Orbigny. Gabb, 1873b, p. 247.

Corbula vieta Guppy. Pilsbry, 1922, p. 427.

Corbula (Aloidis) vieta Guppy. Maury, 1917, p. 231-232, pl. 39,
fig. 13.

Corbula (Aloidis) sancti-dominici Maury, 1925, p. 98-99, pl. 19,
n2 2

Corbula (Varicorbula) vieta Guppy. Anderson, 1994, fig. 3.2.

Diagnosis.—Species characterized by highly inflat-
ed right valve with concave dorsoanterior slope, even“
ly rounded ventral margin, relatively narrow and an-
teriorly directed umbo, and near absence of rostrum
and keel. Left valve characterized by quadrate shap®
and uninflated umbo.

Description. —Shells of moderate size. Valves
strongly inequivalved in size, shape, and ornamenta-
tion. Right valve larger, greatly inflated, and subtrian“
gular. Right valve also with very subtle to obsolete
keel and rostrum, evenly spaced moderately coarse
flat-topped concentric ribs, and evenly convex ventra
margin. Valve shape varies with size. Small right
valves inflated with concave dorsoanterior slope, umbo
directed anteriorly, and obsolete keel. Inflation increas-
es, dorsoanterior slope becomes less concave, ап
umbo becomes more dorsally oriented with increasing
size. Subtle keel and slight rostrum also develop и
size increases.

Left valves more elongate and quadrate, and much
less inflated than right. Left valves with faint, concen”
tric growth lines and widely spaced and slightly more
distinct radial striations. Radial striations absent 9
posterior slope. Small left and right valves (2-3 mm

CORBULID BIVALVES: ANDERSON 21

In length) very similar in shape, but radial ribs more
Prominent on left valves and сопсепшс ribs more
Prominent on right valves. As size increases, left
Valves become more quadrate and keel become more
Strongly expressed.

Hinge of right valve with small, blunt, anterior car-
dinal tooth directly beneath beak and posterior, broad,
Socket-like resilifer that extends slightly beneath beak.
Resilifer, appears reduced because hinge plate con-
Cave. Subsidiary denticle may occur above cardinal
tooth, formed by projection of valve margin beneath
beak. Furrows present on either side of right valve
hinge for reception of left valve. Left valve hinge with
ап anterior small subtriangular socket, and posterior
broad chondrophore. Dorsal surface of chondrophore
with prominent anterior ridge, more subdued midline
паре, and posterior ridge with small denticle. Posterior
Portion of chondrophore projects strongly upward and
May be visible externally. Muscle scars not thickened.

Osterior scar circular; anterior scar ovate. Pallial sinus
Obsolete,

Type Material. —Holotype: PRI 903 (right valve).
Figured by Maury (1917, РІ. 39, fig. 13; 1925, РІ. 19,
fig. 2), and refigured here (Pl. 3, fig. 12).

Type locality.—Río Cana at Caimito, Dominican
Republic (designated by Maury, 1925). The type lo-
Cality is here restricted to TU 1230: Cercado Forma-
tion (Upper Miocene), Río Cana, Dominican Republic
(fide Jung, 1986).

Material.—Measured and/or figured specimens:
NMB с 14134, 14135, 14136, 14137. Other speci-
Mens (over 1,100) are cataloged by locality, which are
‘Sted in the Occurrence section.

Measurements.—See Table 8.

1 Remarks.—Corbula sanctidominici typically occurs

Cattered in silts and іп bioclastic beds with a silty
E This species also occurs in bioclast-rich lenses

burrow-fills within silts.
p literature for Caribbean Neogene and Quater-
У ГУ Varicorbula is complex. Of primary concern here
Те whether Corbula (Varicorbula) sanctidominici is а

Та j DAT
ici эн 8.—Measurements of Corbula (Varicorbula) sanctidomin-
: Figures are in mm.

specimen valve length height width

p
RI 903: holotype (Pl. 3, fig. 12) right 95 96 5.
ВО 14135: (PL 3, figs. 13,

8); NMB locality 16837 Msht | ТЕ 7775-50
чм С 14134: (Pl. 3, figs. 16,
No NMB locality 16837 left 46 40 17

m О 14136: (PL 3, fig. 14);
N U locality 1250

В С 14137: (PI. 3, fig. 15);
U locality 1250 right 44 41 20

right Ta 5.4 2.6

junior synonym of 1) C. vieta Guppy, 1866a, a species
originally described from the Miocene Manzanilla For-
mation of Trinidad, or 2) C. operculata Philippi, 1848
(=C. disparilis Orbigny, 1846?), a Recent Atlantic
species. Gabb (1873b) placed Varicorbula from the
Dominican Republic in C. disparilis and also consid-
ered C. vieta to be a junior synonym of C. disparilis.
Recent specimens of Varicorbula of the western At-
lantic represent a taxonomic quandary, and they have
been variously assigned to Corbula disparilis Orbigny,
1846?; C. operculata Philippi, 1848; and C. philippii
Smith, 1885 (see Weisbord, 1964 and Woodring, 1982
for discussion). The synonymy of these names is not
controversial, although authors differ on which is se-
nior. Woodring (1982) states that Orbigny’s illustration
is poor and the dates of publication controversial; the
imprinted date is 1846, although the probable publi-
cation date is 1853 (Keen, 1971). Those who doubt
the 1846 publication date use C. operculata Philippi,
1848 for this species. However named, the Recent spe-
cies appears distinct from C. vieta s. s. and C. sanc-
tidominici, because it attains a larger maximum size,
is more elongate in shape with gentler anterior and
posterior dorsal slopes, has coarser ribs, and has a
stronger rostrum and keel.

Pilsbry (1922), in his revision of Gabb’s Dominican
species, concurs with Dall (1898) that C. vieta and C.
disparilis are not synonymous, and assigned Gabb’s
specimens to C. vieta. Maury (1917) first placed her
Dominican Varicorbula in C. vieta, but later (Maury,
1925) considered her Dominican form to be distinct,
and named it C. sanctidominici. Maury (1925) stated
that her Dominican specimen was larger (10 mm long,
10 mm high compared to 6 mm X 6 mm for C. vieta
from the type area), more inflated, and had more nu-
merous (about 26 rather than about 25) and more
closely spaced ribs. Maury (1925), however, did not
place Gabb’s (1873b) Dominican Varicorbula speci-
mens in synonymy with C. sanctidominici, stating they
are either C. vieta or C. sanctidominici.

The type material of C. vieta (USNM 115650; left
valves of C. vieta were assigned to Erycina tensa Gup-
py, 1866a, USNM 115652) are not well preserved, and
the margins of the larger right valve are not complete,
making comparison difficult. Corbula vieta from the
type locality shows the same size range as Dominican
varicorbulids, but tends to be more elongate, have a
wider umbo, less concave dorsoanterior slope, and
stronger keel.

The material from this study agrees well with
Gabb’s (1873b) material (ANSP 2690) from the Do-
minican Republic. Maury’s type (PRI 903) of C. sanc-
tidominici is, however, somewhat unusual for Domin-
ican Varicorbula. Her specimen is a right valve that is

22 BULLETIN 351

unusually large and coarsely ribbed. Unfortunately, the
ventral margin is not intact so it is difficult to deter-
mine original valve shape, a diagnostic feature. Nev-
ertheless, the traits that Maury noted as diagnostic for
C. sanctidominici seem to be correlated with size in
Dominican Varicorbula. Therefore, I place both my
material and Gabb's Dominican material in C. sancti-
dominici.

Comparison.—Corbula (Varicorbula) sanctiander-
aea Maury (1925) of the Miocene Manzanilla For-
mation of Trinidad has a left valve with a very convex
beak that is subequal in size to the right valve's beak.
In addition, the right valve of C. sanctianderaea is
relatively more elongate and subequilateral with a wid-
er umbo and stronger rostrum than C. sanctidominici.

Corbula heterogena Guppy (in Dall, 1898) of the
Pliocene Bowden Formation of Jamaica is very similar
in size and shape to C. sanctidominici. In C. hetero-
gena, however, the umbos of the right valve are less
elevated and prominent, and are not as strongly di-
rected anteriorly because the dorsoanterior margin is
less concave. Therefore, in C. sanctidominici the right
valve umbo appears narrower and more set off from
the ventral portion of the valve. In addition, the right
valve of C. heterogena is more elongate and has a
stronger rostrum and keel. The left valves of C. het-
erogena are more inflated and have more prominent
beaks.

Woodring (1982) assigns specimens of the Miocene
and Pliocene of Panama and Costa Rica to Corbula
(Varicorbula) disparilis. He also considered C. wal-
tonensis Gardner, 1928 from the Miocene of Florida a
junior synonym of C. disparilis, thus allowing much
morphologic variation in the species. Woodring's
(1982) figured specimens are very similar to C. het-
erogena. 'These Central American forms differ from C.

sanctidominici in having a stronger rostrum and keel,
beaks directed dorsally rather than anteriorly, and à
somewhat sinuous rather than straight ventral margin.

Occurrence.—This species was collected from the
following areas (see Saunders et al., 1986 for locality
information):

Río Cana: Cercado Formation: TU 1230, and NMB
16835, 16838, 16837, 16842, 16857, 17005. Lower
Gurabo Formation: NMB 16828, 16832, 16833,
16834. Upper Gurabo Formation: TU 1354, and NMB
16817, 16818, 16959, 16824.

Arroyo Ballaco: Cercado Formation: TU 1420.

Río Gurabo: Cercado Formation: NMB 15895,
15896, 15906, 15900. Lower Gurabo Formation: TU
1215, 1277, 1211, and NMB 15846, 15860, 15863,
15864, 16810, 15869, 15871, 15873, 15874, 15878.
15882, 15836, 15944, 15952, 15954. Upper Gurabo
Formation: TU 1210, and NMB 15804, 15805.

Río Mao: Cercado Formation: NMB 16927, 16915,
16926, 16924 (all correspond to Arroyo Вајбп); NMB
16929, 16914 (all Maury’s Bluff 2); ТО 1410, and
NMB 16802 (between Maury’s Bluff 1 and 2); TU
1293, NMB 16910 (Maury’s Bluff 1); TU 1225
(downstream of Maury’s Bluff 1).

Cañada Zalaya: Gurabo Formation: TU 1227,
1227a, 1453, 1453a.

Río Yaque del Norte: Baitoa Formation: ТІ) 1363,
and NMB 16938 (all López section). Unnamed Еог
mation: ММВ 17273 (Arroyo López). Gurabo For
mation: TU 1449, 1448, 1405, 1403 (La Barranca); TU
1206 (Santiago).

Rio Verde: Gurabo Formation: TU 1250.

Distribution. —Upper Lower to lower Middle Mio
cene Baitoa, Upper Miocene Cercado, Upper Miocen®
to Lower Pliocene Gurabo Formations, Dominican Re
public.

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1848. Beschreibung zweier neuer Conchylien aus der Sam-
mlung des Herrn Consul Gruner in Bremen. Zeitschrift
für Malakozoologie, vol. 5, pp. 13-27.

Pilsbry, H. A.

1922. Revision of W. M. Gabb's Tertiary Mollusca of Santo Do-
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Proceedings, vol. 73, pp. 305-435, pls. 16-47.

1932. Notes on a Panamic corbulid clam. The Nautilus, vol. 45,

PALOS
Ramírez, R.

1950. Descripción de algunos moluscos del Mioceno del Valle
del Cibao de la República Dominicana. Publicaciones de
la Universidad de Santo Domingo, series 4, vol. 1, pp- 22
39, pls. 1-7.

Rios, E. de C.

1975. Brazilian marine mollusks iconography. Museu Oceano

gráfico, Río Grande-RS, XII, Brazil, 331 pp., 91 pls.
Saunders, J. B., Jung, P., and Biju-Duval, B.

1986. Neogene paleontology in the northern Dominican Repub-
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letins of American Paleontology, vol. 89, pp. 1-79, pls
1-9.

Schmidt, F. C.

1818. Versuch über die beste Einrichtung zur Aufstellung, Be”
handlung und Aufbewahrung der verschieden Naturkórper
und Geganstünde der Kunst, verzüglich der Conchylien”
Sammlungen, nebst kerzer Beurtheilung der conchyliol-
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Zusammenstellung und Vergleichung der sechs beste?
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Verzeichniss der am meisten bekannten Conchylien 21"
gehänge ist, wie solche nach dem Lamarkischen System
geordnet werden kónnen. Gotha. Justs Perthes, pp. Th
ША

Seyfried, H., Sprechmann, Р., and Aguilar, Т.

1985. Sedimentología y paleoecología de un estuario del lito
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Smith, E. A.

1885. Lamellibrachiata, Report on the scientific results
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Zoology. vol. 13, pp. 1-341, pls. 1-25.

Sowerby, G. B. |

1850. іп Moore, 7. С., Descriptions of new species ОЁ fossil
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Stenzel, H. B., Krause, E. K., and Twining, J. T.

1957. Pelecypoda from the type locality of the Stone City Beds
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Stewart, R. B.

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Vokes, H. E.

1945. Supraspecific groups of the pelecypod family Corbulida®
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ral

of the

Ше
ра,

pub”

phi
4

CORBULID BIVALVES: ANDERSON 25

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Weisbord, N. E.

1964. Late Cenozoic pelecypods from northern Venezuela. Bul-
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26 BULLETIN 351

APPENDIX 1

Criteria used to distinguish four environmental cat-
egories, based on the paleoenvironmental interpreta-
tions of Saunders et al. (1986) and Bold (1988) pri-

30 left or right valves, 30 of each were randomly selected. All se-
lected valves were measured but due to missing data on less than
pristine valves, not all may have been incorporated into the principal
components analyses (PCA).

marily using macrofossil and microfossil assemblages.

Brackish:

Shallow:

Intermediate:

Characterized by mollusks such as
Larkinia, Mytilus, and Melongena,
Anadara patricia, and brackish-wa-
ter ostracodes.

Depths approximately 0-30 m.
Characterized by shallow-marine
benthic foraminifera (Amphistegi-
na, soritids, miliolids), shallow-ma-
rine mollusks (Pachycrommium,
Stigmaulux, Anadara, Tellina,
Strombina) and shallow-marine os-
tracodes (Cytherella, Radimella,
Caudites, Proteoconcha, Loxocon-
cha, Paracytheridea). Planktonic
foraminifera and calcareous nan-
nofossils are absent.

Depths approximately 30-100 m.

Characterized by abundant and di-
verse assemblage of shallow-water
ostracodes, benthic foraminifera,
corals, mollusks (including Strom-
bus, Oliva, Polystira, and Lyria),
and a few planktonic foraminiferal
species.

Depths > 100 m. Characterized by
rich assemblage of planktonic fo-
raminifera, high ratios of plankton-
1c to benthic foraminifera, deep wa-
ter ostracodes (e.g., Krithe), and
pteropods.

Deep:

APPENDIX 2

Samples used for analyses illustrated in Text-figures 4-6. Samples
selected to provide broad stratigraphic coverage. Abbreviations are
as follows:

Section = Stratigraphic section; Cana = Río Cana; Gurabo = Río
Gurabo; Mao = Río Mao; Yaque = Río Yaque del Norte; Zalaya =
Cañada Zalaya.

Formation = Geologic Formation. If blank, Saunders er al. (1986)

did not assign.

Age = Geologic age.

Environment = Environmental category (for Corbula sericea). See
Appendix 1 for criteria used to distinguish categories.

Sed. Type = Sediment type (for Corbula viminea). Categories are:
1 = silts with bioclasts in lenticular beds or lenses; 2 = silts with
bioclasts filling burrows; 3 = silts with mollusks dispersed in matrix;
4 = silty sands with bioclasts dispersed or concentrated,

Valves = Number of valves used in each sample. If a sample
contained less than 30 valves (left or right) all were measured, For
С. sericea, if a sample contained more than 30 right valves, 30 were
randomly selected. For C. viminea, if a sample contained more than

Corbula (Caryocorbula) sericea

environment valves

section sample formation age

Cana NMB 16865 Gurabo Pliocene Intermediate 3
Cana NMB 17009 Gurabo Pliocene Intermediate 4
Cana NMB 16848 Cercado Miocene Brackish 22
Сапа TU 1230 Cercado Miocene Shallow 19
Сала UW 1863/27 Cercado Miocene Shallow 22
Cana NMB 16855 Cercado Miocene Shallow 21
Gurabo ММВ 15833 Мао Pliocene Deep 7
Gurabo TU 1352 Mao Pliocene Deep 10
Gurabo NMB 15805 Gurabo Pliocene Deep 24
Gurabo ММВ 15842 Gurabo Miocene Intermediate 10
Gurabo NMB 15846 Gurabo Miocene Intermediate 10
Gurabo ММВ 15863 Gurabo Miocene Intermediate 12
Gurabo ММВ 16810 Gurabo Miocene Intermediate 17
Gurabo ММВ 15869 Gurabo Miocene Intermediate 21
Gurabo NMB 15911 Cercado Miocene Shallow 1
Gurabo ММВ 15912 Cercado Міосепе Shallow 1
Мао NMB 16910 Miocene Intermediate 5
Mao NMB 16802 Miocene Intermediate 7
Mao NMB 16929 Miocene Shallow 24
Mao NMB 16928 Miocene Shallow 25
Mao UW 1863/25 Miocene Shallow 25
Mao UW 1863/26 Miocene Shallow 23
Mao NMB 16913 Miocene Shallow 23
Yaque ТО 1206 Pliocene Deep 16
Zalaya TU 1227A Pliocene Deep 27
Yaque TU 1403 Pliocene Deep 16
Yaque ММВ 17278 Miocene Shallow 2
Yaque ММВ 17273 Miocene Brackish 3

Corbula (Bothrocorbula) viminea

section sample formation age sed. type valves
Cana NMB 16833 Gurabo Miocene 2 28
Cana ММВ 16832 Gurabo Miocene 3 5
Cana NMB 16837 Cercado Miocene a 9
Cana NMB 16838 Cercado Miocene E 3
Cana TU 1230 Cercado Miocene 2 53
Cana NMB 16857 Cercado Miocene І 10
Cana ММВ 17005 Cercado Miocene 1 4
Gurabo NMB 15878 Gurabo Miocene І 4
Gurabo TU 1297 Gurabo Miocene 2 7
Gurabo ММВ 15882 Gurabo Miocene 2 4
Gurabo ММВ 15906 Cercado Miocene 1 47
Gurabo ММВ 15900 Cercado Miocene 1 37
Mao ММВ 16915 Міосепе | 2
Mao NMB 16917 Miocene 1 „a
Mao ММВ 16923 Міосепе l 21
Mao NMB 16927 Miocene 1 1
Yaque TU 1445 Miocene — 2
Yaque ММВ 16938 Baitoa Miocene 3 5
Yaque ММВ 16936 Вайоа Miocene 3
Yaque ММВ 16935 Baitoa Miocene 4 1
Yaque ММВ 17289 Baitoa Miocene 4 2
Yaque ТО 1364 Baitoa Міосепе 4 9
xaque ІШ 1363 Вайоа Міосепе 4 2
Yaque ММВ 17288 Baitoa Miocene 3 2
Yaque ММВ 17286 Baitoa Miocene 4 |

PLATES

28 BULLETIN 351

EXPLANATION OF PLATE 1

Figure Page
1-5, 10, ГІ, 15, 14, Corba (БОШОСО УШ а) viminea Guppy, 18660 ie cue ea 12
1, 4. Hypotype. NMB G 14110; from locality TU 1364: Rfo Yaque del Norte, upper Lower to lower Middle Miocene Baitoa
Formation. Right valve: 1. interior, X 4%; 4. exterior, X 4. Length, 14.0 mm; height, 9.7 mm; width, 4.8 mm. |
2, 5. Hypotype. NMB G 14109; from locality TU 1364: Rfo Yaque del Norte, upper Lower to lower Middle Miocene Baitoa
Formation. Left valve: 2. interior, Х 4%; 5. exterior, Х 4%. Length, 13.3 mm; height, 9.3 mm; width, 3.8 mm.
3. Syntype. BMNH 64088. Bowden Formation, Jamaica. Right valve exterior. Length, 19.1 mm; height, 13.9 mm; width, 6.3

mm; х 3.
6. Syntype. BMNH 64099. Bowden Formation, Jamaica. Left valve exterior. Length, 15.1 mm; height, 10.3 тт; width, 5.0
mm; X 3.
7. Hypotype. NMB G 14108; from locality TU 1230: Río Cana, Upper Miocene Cercado Formation. Right valve exterior.
Length, 14.2 mm; height, 9.9 тт; width, 5.2 mm; X 4. |
8. Hypotype. ММВ С 14107; from locality TU 1230: Río Cana, Upper Miocene Cercado Formation. Left valve exterior. |

Length, 14.1 mm; height, 9.7 mm; width, 5.0 mm; X 4%. |

10. Hypotype. NMB G 14106; from locality NMB 15900: Río Gurabo, Upper Miocene Cercado Formation. Right valve exterior. |
Length, 16.3 тт; height, 11.5 mm; width, 6.0 mm; х 3. |

11. Hypotype. NMB G 14105; from locality ММВ 15900: Río Gurabo, Upper Miocene Cercado Formation. Left valve exterior.
Length, 15.3 тт; height, 11.3 mm; width, 5.4 mm; X 3. |

13. Hypotype. ММВ О 14104; from locality NMB 16923: Río Мао, Upper Miocene Cercado Formation (Maury's Bluff 3).
Right valve exterior. Length, 17.4 mm; height, 12.3 mm; width, 6.0 mm; Х 4. |

14. Hypotype. ММВ С 14103; from locality NMB 16923: Río Mao, Upper Miocene Cercado Formation (Maury’s Bluff 3). |
Left valve exterior. Length, 15.9 mm; height, 11.8 mm; width, 4.9 mm; X 4. |

9, 12. Corbula (Caryocorbula) dominicensis Gabb, 1873b |

9, 12. Hypotype. ММВ G 14111; from locality NMB 17821: Río Yaque del Norte, upper Lower to lower Middle Miocene Baitoa

Formation. Articulated shell: 9. right side; 12. left side. Length, 15.3 mm; height, 9.2 mm; width, 6.9 mm; X 4%.

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 110 PLATE

|

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 110 PLATE 2

CORBULID BIVALVES: ANDERSON 29

EXPLANATION OF PLATE 2

Figure Page
1,2, 4, 5, Согрша (Garyocorbula) dominicensis Gabh, 1875555 e е e onse e eR узор. 14
1, 4. Hypotype. ММВ G 14113; from locality TU 1226: Río Yaque del Norte, upper Lower to lower Middle Miocene
Baitoa Formation. Right valve: 1. interior; 4. exterior. Length, 14.2 mm; height, 9.0 mm; width, 4.0 mm; X 4%.
2, 5. Hypotype. NMB G 14112; from locality TU 1226: Río Yaque del Norte, upper Lower to lower Middle Miocene
Baitoa Formation. Left valve: 2. interior; 5. exterior. Length, 13.6 mm; height, 8.2 mm; width, 3.1 mm; x 4%,
3, 6. ?Corbula (Caryocorbula) dominicensis Gabb, 1873b ................................................ 14
3. Hypotype. PRI 29033. Río Cana, Upper Miocene Cercado Formation; figured in Maury (1917). Left valve exterior.
Length, 13.4 mm; height, 8.6 mm; width, 3.3 mm; X 4%.
6. Hypotype. PRI 29033. Rio Cana, Upper Miocene Cercado Formation; figured in Maury (1917). Right valve exterior.
Length, 12.6 mm; height, 8.6 mm; width, 3.2 mm; X 4%.
ИЗД. Corbula (Caryocorbula) sericea АЛ... .....,....... 00 15
7. Lectotype. USNM 135655. Moín Formation, Limón, Costa Rica. Right valve exterior. Length, 5.3 тт; height, 4.0
mm; width, 2.0 mm, X 4%.
8. Syntype of Corbula (Cuneocorbula) cercadica Maury, 1917 (mislabeled as Corbula (Cuneocorbula) caimitica Mau-
гу, 1917). PRI 29035. Río Mao, Upper Miocene Cercado Formation (Maury's Bluff'3). Right valve exterior. Length,
6.6 mm; height, 4.6 mm; width, 2.3 mm; Х 4%.
9. Syntype of Corbula (Cuneocorbula) cercadica Maury, 1917 (mislabeled as Corbula (Cuneocorbula) саітііса Mau-
ry, 1917). PRI 29035. Río Mao, Upper Miocene Cercado Formation (Maury's Bluff 3). Left valve exterior. Length,
5.6 mm; height, 3.9 mm; width, 2.0 mm; X 5%.
10, 14. Hypotype. NMB G 14119; from locality NMB 16848: Río Cana, Upper Miocene Cercado Formation. Right valve:
10. interior, X 4%; 14. exterior, X 5%. Length, 7.0 mm; height, 5.0 mm; width, 3.0 mm.
11, 15, Hypotype. NMB G 14118; from locality NMB 16848: Río Cana, Upper Miocene Cercado Formation. Left valve:
11. interior, X 4%; 15. exterior, X 5%. Length, 7.5 mm; height, 4.7 mm; width, 2.4 mm.
12. Hypotype. NMB G 14121; from locality NMB 16855: Río Cana, Upper Miocene Cercado Formation. Right valve
exterior. Length, 8.1 mm; height, 4.8 mm; width, 2.7 mm; X 5%.
13. Hypotype. NMB G 14120; from locality NMB 16855: Río Cana, Upper Miocene Cercado Formation. Left valve
exterior. Length, 7.1 тт; height, 4.6 mm; width, 2.3 mm; Х 5%.
16. Hypotype. NMB G 14115; from locality NMB 16928: Río Mao, Upper Miocene Cercado Formation (Arroyo Bajón).
Right valve exterior. Length, 6.1 mm; height, 4.5 mm; width, 2.1 mm; Х 524.
17. Hypotype. NMB G 14114; from locality NMB 16928: Río Mao, Upper Miocene Cercado Formation (Arroyo Bajón).
Left valve exterior. Length, 5.6 mm; height, 4.0 mm; width, 1.8 mm; X 6%.
18. Hypotype. NMB G 14117; from locality NMB 15869: Río Gurabo, Upper Miocene part of Gurabo Formation. Right
valve exterior. Length, 4.9 mm; height, 3.2 mm; width, 1.6 mm; X 6%.
19. Hypotype. NMB G 14116; from locality NMB 15869: Río Gurabo, Upper Miocene part of Gurabo Formation. Left
valve exterior. Length, 4.2 mm; height, 2.9 mm; width, 1.4 mm; х 6%.
20. Hypotype. NMB G 14123; from locality TU 1227A: Cañada Zalaya, Lower Pliocene part of Gurabo Formation.
Right valve exterior. Length, 6.0 mm; height, 4.2 mm; width 2.3 mm; X 5%.
Hypotype. NMB G 14122; from locality TU 1227A: Cañada Zalaya, Lower Pliocene part of Gurabo Formation. Left
valve exterior. Length, 5.6 mm; height, 3.9 mm; width 2.0 mm; X 5%,
#926, Corbula (Jullacorbula) fossilis Pllabry, 1922... EAR eene rer ne о с.
22, 23. Holotype. ANSP 2689; Dominican Republic (locality unknown). Right valve: 1. exterior, 2. interior. Length, 8.7 mm;
height, 6.1 mm; width, 2.7 mm; X 4.
24. Paratype. ANSP uncataloged; Dominican Republic (locality unknown). Left valve exterior. Length, 8.4 тт; height,
5.8 mm; width, 2,8 mm; x 5%.
25. Hypotype. NMB G 14125; from locality NMB 16817: Río Cana, Lower Pliocene part of Gurabo Formation. Right
valve exterior. Length, 5.6 mm; height, 3.7 mm; width, 1.4 mm; X 7%.
26. Hypotype. ММВ С 14124; from locality NMB 15914: Río Gurabo, Upper Miocene Cercado Formation. Right valve
exterior. Length, 5.7 mm; height, 4.1 mm; width, 1.6 mm; X 6%.

21.

30 BULLETIN 351

|
EXPLANATION OF PLATE 3 |

Figure Page
1-10. Corbuls (Езпаписогота) Cande new species пъ ка bins ни e es кава ee ne COE E YO Y YF YET ығ) 19
1, 2. Holotype. NMB G 14126; from locality NMB 16845: Rfo Cana, Upper Miocene Cercado Formation. Articulated shell: |
1. right side; 2. left side. Length, 14.4 mm; height, 11.8 тт; width, 9.6 mm; х 3%. |
3, 4. Paratype. NMB G 14130; from locality NMB 16841: Río Cana, Upper Miocene Cercado Formation. Right valve: 3. |
exterior, 4. interior. Length, 12.4 тт; height, 10.2 тт; width, 3.4 mm; X 4%. |
5. Paratype. NMB G 14129; from NMB 16841: Río Cana, Upper Miocene Cercado Formation. Right valve exterior. Length, |
13.9 mm; height, 11.2 mm; width 4.8 mm, X 4%. |
6, 7. Paratype. ММВ G 14127; from locality NMB 16841: Río Cana, Upper Miocene Cercado Formation. Left valve: 6.
exterior, 7. interior. Length, 12.2 mm; height, 9.9 mm; width, 3.6 mm; X 4%.
8. Paratype. NMB G 14128; from NMB 16841: Rio Cana, Upper Miocene Cercado Formation. Left valve exterior. Length,
13.5 mm; height, 10.4 mm; width 4.3 mm; X 4%.
9. Hypotype. NMB G 14131; from NMB 16841: Rio Cana, Upper Miocene Cercado Formation. Left valve exterior. Length,
8.3 mm; height, 7.0 mm; width, 2.3 mm; X 4%.
10. Hypotype. NMB G 14132; from NMB 16841: Rio Cana, Upper Miocene Cercado Formation. Right valve exterior. Length,
7.1 mm; height, 5.5 mm; width, 1.5 mm; Х 4%. 19

11. Corbula (Panamicorbula) sp. ..¿. rose a ndo Yr eon э ж Ран ЧЕНА |
11. Hypotype. NMB G 14133; from locality NMB 16990: Río Cana, Upper Miocene Cercado Formation. Length, 5.4 mm; |
height, 4.0 mm; width, 1.3 mm; Х 4%. |
12-18. Corbula Cy aricorudla) sancuaominicn Maury, 1923 ....:..... ra ey etr nde arr ee 19
12. Holotype. PRI 903. Río Cana, Cercado Formation. Right valve exterior. Length, 9.5 mm; height, 9.6 mm; width, 5.1
mm, X 4%,
13, 18. Hypotype. NMB G 14135; from locality NMB 16837: Río Cana, Upper Miocene Cercado Formation. Right valve: 13.
exterior, X 4%; 18. interior, X 4%. Length, 7.8 тт; height, 7.9 тт; width, 4.6 mm.
14. Hypotype. NMB G 14136; from locality TU 1250: Río Verde, Lower Pliocene part of Gurabo Formation. Right valve
exterior. Length, 5.5 mm; height, 5.4 mm; width, 2.6 mm, Х 4%.
15. Hypotype. NMB G 14137; from locality TU 1250: Río Verde, Lower Pliocene part of Gurabo Formation. Right valve
exterior. Length, 4.4 mm; height, 4.1 mm; width, 2.0 mm, x 4%.
16, 17. Hypotype. NMB G 14134; from locality NMB 16837: Rio Cana, Upper Miocene Cercado Formation. Left valve: 16.
exterior, X 4.7, 17. interior, X 6%. Length, 4.6 mm; height, 4.0 mm; width, 1.7 mm.

Фе
os
бә

CORBULID BIVALVES: ANDERSON 31

INDEX

Note: Page numbers are in light face; plate numbers are in bold face type; pages on which principal discussion occur are in italics.

EB 00002202020“ ы 18,19
RA A вада где е 17
aequalis,

Corbula era e ФБ) екен трен 19

РО УНА C Iota qo TUE OS 18
аедиіраіуіѕ,

"Сената О ае 17

Corbula (ПИЯ ОТЫН ПАКТ КН 17,18

шы НО нины р 17
aff. С. inflata,

Corbula ЧОНАН ЛОО A Са СН c Ee КТІ ТЕСТІНІ 19

Ardo нов ен eS TS 11
o REM et 14
alabamiensis.

MEN ы 00-0 BUNT 14

Штоп, Ме О readline yon О 6,16
Aloidis ОТЕП von: Mühlteldt, IB stir 12
ёл, Orbigny, 1826... Ло 26
MEN Gin (MT оо сата 26

8randis (Broderip and Sowerby, 1829) ........................... 9

Patricia a ВЈ до ms wees A ан 9,20,26

ШЕ ЕШ Ән dert E 11
к= КОНАК АНЫ Е 6,8-12,15,16,19,20
Eu INIURIIS АА 15

SP (Academy of Natural Sciences of Philadelphia,

ч Philadelphia, LIA A e Lia а БРЕ ој ДК 6,11,14,15,18,21

ға” beds 9,19

Baitoa Formation

И оо е ое" 6
ШО ШШ ДН S E 19
В MB OS a ниви 6
ca (Natural History Museum, London, U.K.) ........ 677415
ШЕ ұн аа ан Ты о а 9,26
NEM тте к ит О едно е және 12

Owden Formation

B a їл. зды.
Te AA DD rasane ae 9
EMEN TREE eee erret rete 17
ПИ Кл ааа ат 12
ШЕК... 7,8

“dimitica,

со ща ШЕЛЛЕОСОТОИЦИ) боса ddd tele падане EA 19,10

6 RANA o ени coercere erect AERE Ls d 19
апае,

(еј

ша WESRADTCC DUI: са сна ње a OO ИО
два pergrata,

ES Wa all dr По EEE 16
Dun Sea... 11
КО

сар ОТЕЦ АНЕ Pielan, 1987 аон ces а 26

И на оне 5,12,17,19,22

rcadica,
C m
Orbula ПА СИР) nenn о AA AIN 13
ро ща AA A НИ 15,16
| айо ШЕ ЦӨЛ. A dud 5,7,8,11,14,15,17-22,26
a ИВ РИИ СО И ea OSEE 13
Ornian St;

с Ontracta ED ЗЕН EEN E O ЦИТ 0-Е а 14

co Pula ал. 17
а аас, 1818... сеен аА 5,12

Gorbulmse ТООТОН NE 12
Corbulu ТОО То eer ERROR MEN lle
дебри РЫШрр! 1030 (o Ое. 17
ИРИНЕ LOS алаа ы сан силтеу лыы 14
СОКЕ МОЛА e CIO Me SH isha Не 17
aan IN e Е EAN пи
КРАМАЎ ОТЕУ об ала ыл at e s 20
е УЧ ро sera 14
kadang AGKS, OOR vetet ТЫ ТТ НЫ 17
nane Joss ВизОгу 922 Аа 17
Rita ОЛ ЕТУ MI SH оа аты ne 15
тасар о DAA VE aie deecuxven eda а enn REIN 18
ОЛОВО Della a San ere CE Too боја T HRS ELIT 13
САИ ДА ОТТЕ ОРА 12
ООУР во еее on ЕТ 20
ео UB USO ина EI 12
Corbula (Aloidis) Megerle von Mühlfeldt, 1811
WE GUY, LODO een 20
КОНОНКО NIBUDY TIRS einen 20
Corbula (Boihrocorbula) Gabb, 15734... „М у а
PFARRER DS еее 13
а OO А ЕВА EEUU TCU ee 13
Vinea GUPPY; 19500 nannten 1,5,6,7-14,15,26
ШІ БОЛАРЫН! S SUC АЛ АҒ АЛ Ы а 13
Сою ийа (Caryocorbula) Gardner, 1926 .............. 5,11,1417,20
caribasa реготата Maury, 1982 coa каа irse кізе 16
СЕТО МУНИ У | a сноси иа eros ends ела 18
daphnis. Maury, 1925. e cedere eren ren nece ett ires 16
GOMOCTAGE Коооп, ТОЧ (erreur ee а ун стента 15
dominicensis GEOG, ISTID аена анаан 1,2,5,14-15
dominicensis veracruzana Perrilliat, 1984 ...................... 15
helenae Maury, 1912 о.е нна вае 55% t mares у лз eh 16
manzanlliensis:Maury, 1928. „urn 17
oropenduia Olsson, 1922 ...... c eee erexerunt 17
oropendula dolicha (Woodring, 1982) ....................... 17
oropenduia stena (Woodring, 1982) ...... «азоонун 17
prenasuta Olsson, 1966. «cere eren tt) тла крану ce 15
ҚТ АСА һа ИВА е ааа сее тег стаен ее 2,5,6,7-11,15-17,26
ЭБИ ИКАН MUL, dU IRE nennen ira 16
Corbula (Corbula) Bruguiere, 1797
SEHasa DL. AA nn E E T 15
A Сто! A ae 12
Corbula (Cuneocorbula) Cossmann, 1886 ........................ 14
CARICA: MAUS, 1917. nern OP LG ens 15,16
cercadion Maury, 1917 i. ees recettes ortho p FE 15,16
cubaniana Orbigny, 1946 „он нон nen iri re 17
dominicensis Gabo 1130. nn en 14
Corbula (Hexacorbula) Olsson, 1932 ..2.....55559 4)... 12
Corbula (Juliacorbula) Olsson and Harbison, 1953 ....... а.
aequivalvis Philippi, 1836 ...................... 2... 17,18
cubaniana Orbigny, 1846 .....................ЦШ..4.4 а а ы 17,18
fossilis Pilsbry, 1922 ...............а ұр. 2,5,7,8,17-18
knoxiana Adams, 1852Б лани ин винова 18
scutata Gardner, 1943 ...................2.5445% eene three 18
Corbula (Notocorbula) Iredale, 1930 ........................ 11,20
vicaria (Iredale, 1930) .......................... үл ее. 20
Corbula (Panamicorbula) Pilsbry, 1932 ............. 5,11,18-19,20
aequalis (Adams, 18522) ................ ӨӨ 19
aff. С. inflata (Adams, 18524) .........................21 1... 19
cande M Sp. esee нан 3,5-8,19,20

јака (Adams, 18228) i.i o eei eere rore еккен torear кек Ее дей 19
ОСЛОНЕ ТОА ШИ око кетке кесекке nort Eye nh кебек еее теі бүй 19
Вр: A нар ERAS AR vive RA ee 3,19-20
trigonalis (Adams, 18528) |... iier rennen 19
Corbula (Serracorbula) Olsson, 1961 „ог, tnr rmn 11
Corbula (Tenuicorbula) Olsson, 1932 .... és ertet 11
Corbula (Varicorbula) Grant and Gale, 1931 ....... 5,11,20,21,22
disparilis Orbigny, 1846. ..... eee eterne 2122
gibba (ӨШ, 1192) колено zen ira cono ПРЕ пева ените 20
heterogena Guppy (in Dall, 1898) ........................... 11,42
operculata Philippi, 1848 ....................... 5. өш». 21
philippii Smith, 1883... ere nam Eee re er nn mann 21
sanctianderaea Maury, 1925 .......... оро ener enn rin 22
sanctidominici Maury, 1925 iiaeie rii вав 3,5-8,11,20-22
viera Сирру, LODOS „х.е... HIA e ent d праски 20,21
waltonensis Gardner, 1928 ............ ver an d
Cossmang (1980) каго кеенен ви Рес Fee eda io 14
A A R c qu M сес ады» 11,14,15,17,19,22
LU ons cp ика edt кек FIP) Lenni ken Си ove еке EMEN 15,16
Limon BOB oro prensa nenn AE Бак que DRE braves 5,11
Ponts Judas. rare ee £n rd доне Бр 11,19
cubaniana,
ОТОРИ T ии 17
Corbula (Сипеосотриіа) «sinn еке к, баклажка бара 17.
Corbulo. (ПИО ВО? ВИИ) .›.......... ‚5+ эки» бабыз» цени 17,18
буйлтеЙа Jones, 2949 ,;.,,.,..,.....-1..»»< pe e eor rh навики 26
МАП ТИЙИ иеа санке н rennen етике eph А А Зе 2,2122
КАН Ос A кн ТТ” 18,19
daphnis,
COPD: (CATVOCOTDUIA) у сна reve erret nera narra nen arena ne 16
democracia,
Corbula ссатуосотоша). r.esrrireresrserirerizinrg ici Tens raves 15
disparilis,
де A O A CO 20
Corbula (Varleörbula). ..eoeceroresiroireerrosracenor rh барға. 21,22
Dominican Repüblie .....„..... еее 5,6,11,12,14,15,17-22
Cibao Valley MM 5,6
Arroyo Bajón ..........................%....4...... 14,17,22
Arroyo ВаПасо ...ioveoonssororscrosconcarecarnacioagrcanonoss 22
JAMTOVO Оре mora rro n nr ir ron rear ios eren ри ОЧИ 17:22
Сайайа ZalayB „етот entro tnnt 6,9,16,17,22,26
A OL EH иа ивици ДИ ќа 6,13,14,16,17
RIS мөс шш Ао ја E ТН АН ѓа 5-10,13-22,26
іле не A ER тету ни ol
ОООО арыс 6,8-11,13,14,16-18,22,26
ОИН, 6,9-11,13,14,16,17,22,26
alla A AN ONO ri 17300
D E OF затне, ОД НЕ 14,17,22
ИГ oO NBI ТОТУ A cvs bree te en 14,17
ООСО ООО Т Зое
Rio Yague del None о 5,6,9-11,13-17,22,26
АМИОАИШЕ wenden ҮРУ девети ЕУ ERA а 14,17
IM TOI etn HERR EET 9,14,22
Bone aV os o РЕ ыы УНЫ ТЕН 17,22
O RE EHE O 722
dominicensis,
COV OWE А а а ҮЛТ” 14
Соғрша(Сатуосотр иа) са ван грива ок ES 1,2,5,14-15
COTOMIANGUTCOCONOUIG) аи ee 14
dominicensis veracruzana,
(ОТРОВАО) е ONE рми) 19
ЕВО MBIA РТ ОР енен Г, СИПА, BOB ГУ 14

BULLETIN 351

РАНИМ ТУШОО оо ee 11,18
ШЕН ee 15
Erycina Lamarck, 1805

Tensa Опору 18068 rra сете ке бо себични ТУ 21
Оре o aterra ку ечат cala Т ВНА QUA ec 20
Kimi naa laa a НСО ee 5,11-13,17,18,22
fossilis,

Corbula ПОШ... 2,5,17-18
CAD ne сити TEE. 11,12
GEB (18730) уко ожау елла сы тко,

AUDE terna кке лкен кетті бекеті T кейін
Gardner (1926)
Gardner (1928)
Gardner (1943)

ССОО ОЙ nern СЈАЕ 15,0
Geological Society of America ...............4 9.9994... аа, 6
gibba,

Corbula (Уалсо? Юй) төк кекті сөзден кеүек yes 20

ТЕЛІМІ, оу ек (3940 GOL AR cre cron re E 20
grandis,

ААИ Ls A RO ви Нета ве PY 19 еке bere: nono OER 9
Grant md Gale (1931). v. eta cr An ро TM diu TAFE 11,20
ЕО От 21
ТОБУ СОВ ao o 5,12,13
A A нш 14
A ТТС” 11,22
Gurabo A RR вредни 5,7,8,14,17,18,22,26
ee TENA qup mattis 6,14
helenae,

Corbula (Сағуосот ша) „ооо sc 16
heterogena,

AA id A MD ТЕКТЕН ККЕ TROU et 11,22
Hodson and Hodson (1931) |. cce eee rne carre eer ит 15
IGM (Intituto de Geología, Ciudad Universitaria de México, БЕ,

Мара IECUR ДЕЗ с че М Аа 11,18
inflata,

Corbula (POnamicorDula) Үл сука ака voran 19

POIDHOTU scr a reas СЕ НН И 18
die N nen A ee 11,20
ИА ЕТЕНЕ TEE, Si 1-14,17,2
Jettery, PAUL „еее тивне етене зевает неее көзі ге evi НИЦИ 6
Juliacorbula Olsson апа Harbison, 1953

aequivalvis (Philippi, 1836) ............... 04 еее, 17
AAA A мын 11,16-182
MU ONE ica ta rs to OG 4
SS TE ONE
E A 1921
knoxiana fossilis, i

COFDUIMM „узин esa esa trahat Fr A
knoxiana, 1

LOPDIBIB. ЗН Id AAA ТУВА Ма кіт тік сер OA 18

Corbula (WIullacorbüla). «es conn one vet ИО SECO ЕЛА 6
ОСІ; ӨПА... en a И %
Krithe Brady, Crosskey, and Robertson, 1874 .................."“
РЕШЕ MOTEL ciis нө куйаас стаж «раза еее к REOR І 1,18%

Courbaril Sand and Clay Member ..................Д.Д0Д 444440" 2
Таша (1709) акел» кеке текте ren nern AA E 12
Lunarck СТВОТО“ „евон өекееекк кке ааа нека ТЫН

——

CORBULID BIVALVES: ANDERSON 33
ds (1818) A ТЕКТЕ т. 12 ЈРВ А ФУ 22 11,15,17,18
UND Tos Relnhert; 1958 ia tra ЫЫ са 26 Р ШЕ 55 ынасы ИЧ. 11415
E KORAN ON ererevecor е 14 Philippi КАКО Крка ыы колу CU d ENTE. 17,18
na, Philippi: (948): onda er rod ка 21
E o ee. O EUREN rubi; 15 philippii,
Б. E T RNC LU E 14 ОГОО СУАТ) een nen ee BED ND 21
"о barrer РАЦЕ UD НУ 12 Pilsbry ММТ Т 5,12,14,15,17,18,20,21
а o. ALS LS OW наи Суве бае E EEA 26 Pilsbry (1932) ЕЕЕ 11,18,19
СЕУ МАРИ a ГО, Ў СМИ 26 A een 26
Macdonaidi Potamomya Sowerby, 1839
Corbula b aegualis Adams. 18028 ra cus eee ee cec сна e PRA REQUE ќе ПОРИ 18
Corbula (P ne аа A ААА 18 SN A ie cueste eiat suoni cutie vidt ie «ӘЛИДІ 18
Manzanilla ne ША) никне 19 ТОЛО Adams, Поза ан o cete оте DUNT 18
o уе ее ns Ly Је renasuta
Manzanillensis, T 1394 ск C bul
Corbula (Caryocorbula) .... 17 Оша. жы дао а Merten Li ела ты и 15
ША fs Сай соо сасу Der эю айла PRI (Paleontological Research Institution, Ithaca, МУ, U.S.A.)
ОТИТ 5,7,8,14,17,26
По ива nes осо не 59,15, se et duh ЖТА, 6,22,13,15,16;18,21
Mau stone Member ............................... Proteoconcha Plusquellec and Sandberg, 1969 .................. 26
DECIDE) c eere rev ved НИМ соге ИНАТ ССА ООСО КК 16 pteropod 26
MENS soe A INI EMEN rs ант Local
Әнеу Ене ДЕ c were АР ВА he ants a ase 9
“Ам oneri d 5,11,12,16-18,20-22 radiatula,
CLean MUS rede ce ao cr, 1? СЕГО СВОГО) OLOR ONERE 13
ОЕ И ае TOT СЕ. 20 ПО ПЕН POLI LOS Cue И I E EO 26
aane МИД ПЕЛ РА ка а ete a 12 АЕО) О eee teet eve d OR те 12
ШО Sohümaoheh ТУ ee ДЖЕЛ NILUS 26 МӨ ШЕШШ SUO C E ERN 14
E MERI IE Ане PIE O we е 111518 RIOSCISTO) дек ек кен тты et ken re е 17
BEEN ooo от осо: ic MEN 11 Rosenberg, Gary |... 6,14
eracruz
miliolig ко EIU T wa
Moin A Пи ~ бы sanctianderaea,
Oore (1969) ay ANS NT 12 1417 1820 (ОТОНА СУВИ НС И) ssai irs керка are 22
e nee ce ЧО
Cambridge, МА, USA... ' "Б у Corbula (Aloidis) een 20
Mytilus a Қ: Sanehklominich
bin етра Сола CVaritorbula) man асан 3,5,6,8,11,20-22
National При аи реда dias. ага Беше 6 A Ore ive еее т 11,15,18
В (Naturhistorisches Museum, Basel, Switzerland) .. 6,/1,13- Заро АР ВР). re 359111613, шы ыг
22 a ГЫ A АИИС 12
North EL c 14.20 ШЕП ДУ УОИ Т 6
{ scutata,
Oak DOW Formation ДО ЫШ 13 (ОПРОСТИ тоа) wre en GRIN sanken 18
PME ME еее 2
ШАП «олие а бр рако зика Ти 20 sericea;
МЕНИ > <= а: Peur 11,12,17 MOP Соза ARA sow оС аланда ар Ren 14
Sson С с тек 9.1 1,12 Corbula (CANVOCOTOWG) „еконо, 2,5,6,7-11,15-17,26
Isson NEN o en en бағ) 9.11.17.18 (ОБРНИ бабаны лы ту чуе орудие ент 15
A ОТ CRN ФИНСКУ Па ТОН | E 15 Seyfried et al; (1985) i eee ао: 11,19
E НАЧИНА СОЗУ EO ES e exo er а 11,17 Sigma Xl ii E ЛЫ eee Od EUG акни ni re ec CA NS ER e 6
culata, Smith (1S) sense E hr on nar nde 21
Corbula (О ЧООР) “Қат Колев nd een 21 smithiana,
e igny ШШ o s. o esci c M ве UN 17.21 Cordula (COPYOCOF BULA) escort iia 16
"орепаша, SOL pipis ЛІК o 26
„КИШ (САКОО КИД. л у. т 17 шй АЙ ИО ое eH ide a на Сее 14,17
Opendula dolicha, Sowerby (1850) | iv e eer nat аиа, 9
s Orbula (ЕРУ NR. SU Ae NN He CN IS 17 Springvale Богшайов ОИ СРРР en 11,18,19
Орепаша stena, Melajo Clay Member «oie nennen nene ns 18
Corbula E un a a ia чиа 17 Stenzel et al. (1957) «iiec eee een innen 20
Pach Stewart (1930) хә. элуу coro sens елек menu sn sans sn 12
e Мы С 26 Stigmaulux Mörch, 1852 ......................... ааа 26
ШШК aet r a o пи ин TON 6 Strombina Mórch, 1852 .................... 0... 26
jm —————— 11,15,17,158/22 Strombus Linnaeus, 1758 ............а... 1.1.11 2. 26
onm а МШЕ; LOWER A PO 26 sulcata,
^ Cla, СОР позора O EH ANNI HE ENERGY ан 12
Nadara 9,20,26 СТЕНА СОРТИ) ои С НА ТН een 12

34 BULLETIN 351

synarmostes,

ОФИС (ШОШ ООР) an Ree chu а hatt a nth trait е 13
Talparo Formation ...errcorororcoricnsciónerarassanerero rre eani 18

Malum Sand and Clay МЕШШӨГ енене еуин nun mns 18
Батор РАШ. э... у НИ РРА ВАРИРА РИМА ПО eH IIR ch PURI M [6]
TOO БЛОА 6 сеннен ВИ ИМ РВ КАМ РАСА 26

ООО Olivi, 1/92. irs esere een rennen hn harm hn nnn suber heen t ey 20
tensa,

РУСА ‚у... уу,» кк кн, кәжә к>, ТЫ ERA I ire POE 21
Thomonde Forman .................., rt ben ts беа no erroe 14
THOMPSON, Jani ......................Ӛ.......99..0..../40000 etree 6
trigonalis,

Cor bula СРИПДРИСОУВША) ъръарежквикикрккав vagy ка реки ER 19

POLIUS rien anta Ke erhebt PA ЕИО RUD Мі де 18
TIERE роба жола имам О ЕР 5,11,16-19,21,22

TU (Tulane University, New Orleans, LA, U.S.A.) .. 6,/7,13-17,21,22

USNM (United States National Museum of Natural History,

Washington, DC, USA) ............... 6,71,19,15,10,19,19;24
UW (Museum of Geology, University of Wisconsin-Madison,
Madison, WEL USA) neueren rte etn 6,/1,16
Маш (1989). A cree des ве i URE кекті» 11
SE A Р T 11,15
A кеуек е Ско thre CERAM ES e 19
vicaria,
Corbula (Мотосотрша) узук, тикове кезекке кебе ғат б) 20
vieta,

(Aaa A ao Y ДО A NR 20

СОЛИ а САТО ука en ива ano НИНА ОТОТ ЕД 20

COP TULA CV ATICOTDILLGL) 6 re mmn нене ПИК NEEL УН MOON 20,21
viminea,

ОРОШ oo eb ON 12

Corbula (Bothrocorbula) ......... tisrisnersais 1,5,6,8-14, 15,20
VON В. C989) ааа рала Со ces o US UE NE НАНЕ E
МЕ ВИНОТО e eee sectione dare acia ка ки ИІСІ» ТЕТІ дм 12,18,19
ОВНА СОВО teeth e tee ie SI PON 12
Volkes ОИ (10920) aae делу ллы е el 13,15
VOLS EMG О ОЛ УОЛТ. 6
Waller, THOMAS: i viua e add 6
waltonensis,

Corbula (Уайсогви я) cc. кин 22
WEA (ДОБИЈЕ аа oia PYTH Y 17,20,21
КӨМІП ШЕКЕСІ... уал оза а EG UC UO HO WM 12,17
estem Atlantic Ocan ОЛЕ moses M n vengo one 11,2]
western: Расе СФваб eec ree си cerent ӨК EHE 20
Westphal, Klaus... eee eere entente e o one la 6
wilcoxii,

Corbula Bothrocorbula) serrer i c eec oc ССИ M EHE 13
WIDER: einsamer Mae 12-15
WOSAHHE (1073) V4 U. ove re f ON 9
МУЗВЕНАВ (1982) 5 ico d wenn e MEO ub DU 11,17,20-22
жынын (1920 cose Edi улан 14
Yonga (1040): nennen нина ка eoe сан мб ија 20
Zone Н of Maury (1917) ........ оснивана ввкика тов ЕЕ 15

NEOGENE PALEONTOLOGY IN THE NORTHERN DOMINICAN REPUBLIC. 17. THE FAMILIES
CUSPIDARIIDAE AND VERTICORDUDAE (MOLLUSCA: BIVALVIA)

PETER JUNG

Naturhistorisches Museum Augustinergasse 2, CH-4051, Basel, Switzerland

ABSTRACT

Seven species belonging to two genera of Cuspidariidae and two genera of Verticordiidae are described and figured. Five of
them occur in the Neogene sections of the Dominican Republic and their stratigraphic occurrences are given. Two species of the
verticordiid genus Trigonulina are known from the Recent fauna only: one from the Western Atlantic, the other from the Eastern
Pacific. The latter is described as new (7. pacifica п. sp.). The two living species are discussed һегеіп for comparative purposes.
Out of the five fossil species, only two are well represented (Cardiomya islahispaniolae and Trigonulina bowdenensis). The

remaining three species are known from very few specimens.

RESUMEN

Se describen siete especies pertenecientes a dos géneros de la familia Cuspidariidae y a dos géneros de la familia Verticordiidae.
Cinco de ellas se encuentran representadas en las secciones del Neogeno de la República Dominicana; se indican sus ámbitos
estratigráficos. Dos especies de verticordidos del género Trigonulina solamente se conocen de la fauna actual. 7. pacifica habita
en el Pacífico Oriental y se describe aqui por primera vez; la otra habita en el Atlántico Occidental. Ambas se mencionan con
Propósitos comparativos. De las cinco especies fósiles, apenas dos estan bien representadas (Cardiomya islahispaniolae y Tri-

gonulina bowdenensis). El material existente de las restantes tres especies es muy escaso.

INTRODUCTION

E paper is a further contribution to the series of
B * studies dealing with Neogene fossils from
lons situated in the Cibao Valley of the northern

| оно Republic (Text-fig. 1). The project and the
| within which these studies are being car-
a LE have been outlined by Saunders ef al. (1982)
aunders et al. (1986). Jung (1986, p. 5) listed the

Er Important early collections of molluscs from this
avail The material of Cuspidariidae and Verticordiidae
thon able for this paper is not rich d nevertheless
ERN it worthwhile studying these two families es-
t Dey considering the facts that (1) illustrations of
(2) a groups in literature are generally rather poor, and
à е method of scanning electron microscopy allows

Produce good illustrations.

AS is the case for all the contributions to this series,
а studied has been collected from measured
es. The geographic location of the investigated
таб 15 shown іп Text-figure 1. For detailed infor-
si E as to geographic locations and stratigraphic po-
Bene of all the collecting stations, as well as to the
teade, ostratigraphic framework and the ages, the
(1986 is referred to the paper by Saunders е! al.

еса ). Formational names have been used with care,
Use correlations of the sections are not certain.

T ACKNOWLEDGMENTS
ES material on which this paper is based was col-
ed during field work carried out in the years 1978,

le

1979, and 1980 as part of the project referred to above.
The field work was made possible by a grant from the
Swiss National Science Foundation (Grant 2.646-
0.76). The financial help and the assistance in the field
provided by Institut Français du Pétrole are gratefully
acknowledged.

I am indebted to the following persons for the loan
of specimens under their care: Gary Rosenberg, Acad-
emy of Natural Sciences, Philadelphia, PA, U.S.A.;
Alan Kabat, Mark Florence, Jann Thompson, Warren
Blow, all of the National Museum of Natural History,
Washington, DC, U.S.A.; Kathie Way, The Natural His-
tory Museum, London, England; James McLean, Los
Angeles County Museum, Los Angeles, CA, U.S.A.

Eugene Coan, Palo Alto, CA, U.S.A., offered useful
comments and information on parts of the manuscript,
which are gratefully acknowledged.

In addition, I am grateful to Richard Guggenheim,
Marcel Düggelin, and Daniel Mathys, all of the Scan-
ning Electron Microscope Laboratory, University of
Basel as well as to Severino Dahint, photographer at
the Naturhistorisches Museum Basel, Switzerland.

BIOSTRATIGRAPHY AND
PALEOBIOGEOGRAPHY

A total of seven species belonging to two genera of
Cuspidariidae and two genera of Verticordiidae is dis-
cussed in this paper. Two species of the verticordiid
genus Trigonulina are known from the recent fauna

BULLETIN 351

Q 10 20km 1 Rio Cano
2 Rio Gurobo
3 Rio Moo
AYUBIN С | В С] Upper Cenotoic 4 Rio Amina
4 О [77] Oligocene - Early Miocene ? 5 Coñodo Zoloyo
5 V ; 6 Rio Yoque del Norte
3 4 е 7 City of Santiago
" 4% 4 4 8 Arroyo Puñol
% 9 3) & Y 9 Rio Verde
9
$ 2 5
- 1 9 VALVERDE ESPERANZA NAVARRETE
%% ZAMBA 2 MAO p
КА Ж
Eu Ф (у € А? шр,
mods LOSQUEMADOS E
Fi e SANTIAGO ir
2 RODRIGUEZ) 4
И 75, | SANTIAGO
BULLA
MOCA
6 8
BAITOA
Text-figure 1.—Index map showing location of investigated areas in the Cibao Valley, Dominican Republic (after Jung, 1986, Text-fig. р).

only, one from the Western Atlantic, the other from
the Eastern Pacific. They have been included here for
comparative purposes.

The stratigraphic occurrences of the remaining five
species аге plotted in Text-figures 2-7. The species are
not continuously present through a given sequence of
sediments, but their occurrences are spotty (Jung and
Petit, 1990, p. 88; Jung, 1994, p. 6). The five species
occur in the following sections:

Río Gurabo section (Text-figs. 2, 3):

Cardiomya islahispaniolae (Maury, 1917)
Cardiomya distira (Dall, 1903)

Haliris jamaicensis (Dall, 1903)
Trigonulina bowdenensis (Dall, 1903)

Río Cana section (Text-fig. 4):

Cardiomya islahispaniolae (Maury, 1917)
Plectodon granulatus (Dall, 1881)
Trigonulina bowdenensis (Dall, 1903)

Río Mao section (Text-figs. 5-7):
Cardiomya islahispaniolae (Maury, 1917)
Arroyo Zalaya:

Cardiomya distira (Dall, 1903)
Haliris jamaicensis (Dall, 1903)

Arroyo Babosico near La Barranca, Río Yaque del
Norte:

Cardiomya distira (Dall, 1903)
Río Verde:
Trigonulina bowdenensis

As сап be seen from the above lists Cardiomya 15
lahispaniolae occurs in the sections of Río Gurabo»
Río Cana, and Río Mao. Trigonulina bowdenensis has
been found in the sections of Río Gurabo, Río Can?
and Río Verde. Cardiomya distira 15 recorded from the
sections of Río Gurabo, Arroyo Zalaya, and Arroyo
Babosico; Haliris jamaicensis from the sections of RI?
Gurabo and Arroyo Zalaya, whereas Plectodon gral”
ulatus is restricted to the Río Cana section.

The representation of these five species in the VA"
ious sections is rather uneven. Four species occur i
the Río Gurabo section, three in the Río Cana section
two in Arroyo Zalaya, and a single species in the sec“
tions of Rio Mao, Arroyo Babosico, and Rio Verde:

The numerical representation of the five species !
the Dominican deposits is uneven as well. Trigonú 0
bowdenensis is represented by 80, Cardiomya isla
ispaniolae by 52 Dominican specimens. On the още!
hand Cardiomya distira is represented by only eight
Haliris jamaicensis by three, and Plectodon gran" =
tus by a single specimen. The three latter species there"
fore are very rare.

CUSPIDARIID AND VERTICORDIID BIVALVES: JUNG 37

К
4
os
u
HOO мат | Ф
4 In `@
“ лч @
пар | []
o [x le
ма = | 5
x z
> 8
= 9
с Е
ч. @ б
E e
5 E
© Р”
Ду
>
Е
2
о
x
©
o Ф
a
с |
па
2
Ф
= &
9
с
[uj
Q
Q
с
5
[7]
јо 1
Е |
5 !
©
=
©
©
2004
= [1004
о
к
ч
>
а
о
u
o
o
<
o
x
ш
о
A

ten *Xt-figure 2.—Columnar section of Río Gurabo showing occur-
Ces af o А > А е: į
65 of species dealt with herein (after Saunders ег al., 1986, Text-

lg. 6 A Е qr қ à
). Numbers in second column from left refer to thickness in m.

E ve five species mentioned above, only Cardi-
' islahispaniolae is endemic to Ше Neogene of the

B e Republic, Three. species, namely Cardi-
distira, Haliris jamaicensis, and Trigonulina

on enensis, also occur in the early Pliocene Bowden
ation of Jamaica, and Plectodon granulatus is
nown from the middle Miocene Shoal River Forma-

=a Haliris jamaicensis
Cardiomya distira

wm Trigonulina bowdenensis

FORMATION

MAO

Text-figure 3.—Río Gurabo: upper part of columnar section show-
ing occurrences of species dealt with herein (after Saunders er al.,
1986, Text-fig. 6). Numbers in second column from left refer to

thickness in m.

tion of Florida, the Pliocene of Florida, and from the
Recent fauna of the Western Atlantic.

ABBREVIATIONS OF REPOSITORY
INSTITUTIONS
ANSP: Academy of Natural Sciences, Philadelphia,
PA, U.S.A.

38 BULLETIN 351

FORMA
== Cardiomya islahispaniolae
Trigonulina bowdenensis

GURABO

=== Plectodon granulatus

FORMATION

CERCADO

Text-figure 4.—Columnar section of Río Cana showing occur-
rences of species dealt with herein (after Saunders et al., 1986, Text-
fig. 16). Numbers in second column from left refer to thickness in m.

BMNH: British Museum (Natural History), London,
England, now The Natural History Museum,
London.

LACM: Los Angeles County Museum of Natural
History, Los Angeles, CA, U.S.A.
ММВ: > Naturhistorisches Museum Basel, Switzer-

Pleistocene/Holocene gravel terrace

169300

==oda_- 173298
17176 m

М^, Cum 16932 e

169290
17179 ш

Ñ——— Cardiomya islahispaniolae

Text-figure 5.—Section exposed in Maury's Bluff 2 on Río Мао

showing occurrence of Cardiomya (Cardiomya) islahispan

jola®

(Maury, 1917) and stratigraphic positions of NMB localities: blac

squares represent localities collected for microfossils and lith

olog?

А dat А 26115
analyses; black circles represent localities collected for macrofo$*

(after Saunders et al., 1986, Text-fig. 31).

CUSPIDARIID AND VERTICORDIID BIVALVES: JUNG 39
gravel terrace Ф
5
9
[n
©
o
Q
Ф Le
8 s
9 2
с
y S.
TE =
[2]
= 9
9 =
0 G
169280 © о
Е
____- 16918 ө 16924 e 16925 e E 16912 е is fallen rubble thought to be |
| —— 169260 17177 ш 17178860 Б зен ои NH
О
169150 16923 ө 169270 |
\
EEN
roms
+: |-—169130 172690) 4—17307 m
169220

Text-figure 6.— Section exposed at mouth of Arroyo Bajön on Río
пи Showing occurrence of Cardiomya (Cardiomya) islahispani-
ку 1917) апа stratigraphic positions of NMB localities:
Mba Squares represent localities collected for microfossils and lith-
teh, analyses; black circles represent localities collected for ma-

Ssils (after Saunders et al., 1986, Text-fig. 32).

land (the letter G after NMB stands for bi-
р valves).
RI: Paleontological Research Institution, Ithaca,
T NY, U.S.A.
Dt Tulane University, New Orleans, LA, U.S.A.
SNM: United States National Museum of Natural
History, Smithsonian Institution, Washing-
ton, DC, U.S.A.

SYSTEMATIC PALEONTOLOGY
INTRODUCTION

The basis for the preparation of this paper has been
ТМ Combined collections of the Naturhistorisches Mu-
i. Basel and Tulane University. All of the figured
E of Cuspidariidae and Verticordiidae derived
lonis these collections are deposited in the Naturhis-
iia ches Museum Basel. It was originally planned to
е апа тей gure the type specimens of all the spe-

Of Cuspidariidae and Verticordiidae occurring in
Spee отіпісап Republic Neogene as well as the type
8 Imens of species which were most important for

Mparative purposes.

Owever, this plan has not been carried out for var-

Text-figure 7.—Section exposed at the downstream (eastern) end
of Maury's Bluff 3 on Río Mao showing occurrence of Cardiomya
(Cardiomya) islahispaniolae (Maury, 1917) and stratigraphic posi-
tions of NMB localities: black square represents a locality collected
for microfossils and lithologic analyses; black circles represent lo-
calities collected for macrofossils (after Saunders et al., 1986, Text-
fig. 33).

ious reasons. As mentioned below under Cardiomya
islahispaniolae, the holotype of that species is badly
broken (Fast, 1978, p. 80) and therefore could not be
refigured. In addition the type specimens of four spe-
cies of Cuspidariidae and Verticordiidae described by
Dall (1903) from the early Pliocene Bowden Forma-
tion of Jamaica have been available to me: Cardiomya
craspedonia, which does not occur in the Dominican
Republic, and Cardiomya distira, Haliris jamaicensis,
and Trigonulina bowdenensis, which occur in the Do-
minican Republic as well. The type lots of all these
four species consist of three specimens each. All those
specimens are glued to a piece of black paper. I tried
to remove the specimens from the black paper using
various chemicals in an attempt to dissolve the glue
but without success. Lectotypes of all four species
have been selected, but they are not refigured herein.

Although the amount of material available for this
study is mentioned under each species, a summary of
the number of lots and specimens of each species is
given in Table 1.

As mentioned above, three of the species discussed
herein are also known from the Bowden Formation of

Table 1.—Numbers of lots and specimens of each of the seven
species of Cuspidariidae and Verticordiidae dealt with in this paper.

number number of

taxon of lots specimens

Cardiomya (Cardiomya) islahispaniolae 20 50
Cardiomya (Bowdenia) distira 8 47
Plectodon granulatus 14 22
Haliris jamaicensis 4 10
Trigonulina ornata 6 1049
Trigonulina pacifica 22 1840
Trigonulina bowdenensis 32 hz

Total 104 3130

Jamaica, but their numerical representation differs
greatly from that in the Dominican Republic. Table 2
gives the number of specimens of the five fossil spe-
cies available from the Dominican Republic and from
Bowden, Jamaica.

A short discussion of species concepts has been giv-
en by Jung (1986, p. 9; 1989, p. 37), and definitions
of the headings used in the following systematic part
may be found in Jung (1989, p. 35) and in Jung and
Petit (1990, p. 93). They are not repeated here.

SYSTEMATICS
Family CUSPIDARIIDAE Dall, 1886

Genus CARDIOMYA A. Adams, 1864
Cardiomya A. Adams, 1864, p. 208

Type species (by monotypy).—Neaera gouldiana
Hinds, 1843. Recent, seas of Japan.

Diagnosis.—Shell of small to medium size, rostrate.
Sculpture consisting of radial ribs. Radial sculpture of-
ten restricted to main shell disc or continuing over the
rostrum as well. There may be secondary radial ribs.
Left hinge with a subumbonal chondrophore but no
teeth. Right hinge with a subumbonal chondrophore
and one or two lateral teeth. Posterior lateral tooth usu-
ally prominent.

Remarks.—Cardiomya not only includes Neogene,
Pleistocene (Grant and Gale, 1931), and Recent spe-
cies but also a number of species from deposits of
Eocene and Oligocene age (Durham, 1944; Gardner,
1945; Harris, 1919; Meyer and Aldrich, 1886; Turner,
1938; Vokes, 1939). According to the Treatise on In-
vertebrate Paleontology (p. N854) the oldest record
dates back to the late Cretaceous. The stratigraphic

range of Cardiomya is therefore late Cretaceous to Re-
cent.

BULLETIN 351

Table 2.—Number of specimens available from the Neogene of
the Dominican Republic and from the Bowden Formation of Ja-
maica.

specimens
from specimens
Dominican from Bowden,
species Republic Jamaica
Cardiomya islahispaniolae 50 0
Cardiomya distira 8 39
Plectodon granulatus 1 0
Haliris jamaicensis D 7
Trigonulina bowdenensis 80 32

Subgenus CARDIOMYA sensu stricto

Cardiomya (Cardiomya) islahispaniolae
(Maury, 1917)

Plate 1, figures 1—6; Plate 2, figures 1—4;
Text-figure 8

Neaera alternata d'Orbigny. Gabb, 1873, p. 248.

Neaera ornatissima d'Orbigny. Gabb, 1873, p. 248; Guppy, 1876,
p 330,

Cuspidaria islahispaniolae Maury, 1917, p. 196, pl. 26, fig. 20.

Cuspidaria ornatior Pilsbry and Johnson, 1917, p. 195; Pilsbry
1922, р. 414, р!. 38, брз. 11, 12.

Cuspidaria gabbi Pilsbry and Johnson, 1917, р. 195; Pilsbry, 1922.
р. 415, pl. 38, fig. 10.

Description.—Shell of medium size (up to 9 mm іп
length), delicate, rostrate. Umbos prosogyrate, place
almost centrally. Sculpture consisting of numerous 127
dial ribs. Posteriormost rib more prominent than the
others, forming a small carina marking the boundary
between main shell disc and rostrum. In addition 19
this small carina there may be one or more ribs just
anterior to it which are more prominent than all the
other ribs on the main shell disc and have wider 11”
terspaces. There may be a few secondary radial ribs.
Except for the more prominent posterior ribs the radia
ribs are usually well developed only on the ventral part
of the main shell disc. Dorsal part of the main she
disc is smooth or sculptured by concentric grow!
lines. The rostrum is smooth or sculptured by grow!
lines. There may be an indication of a radial rib пейї
its postero-dorsal margin, where the growth lines are
coarser and more prominent. Left hinge with 4 s :
umbonal chondrophore but no teeth; margin bent ІЛ :
dorsal direction anteriorly and posteriorly. Right hing?
with a subumbonal chondrophore and a promine?
posterior lateral tooth; antero-dorsal margin ben
slightly upwards.

Holotype of C. islahispaniolae.

PRI 28904. This 8

а right valve which is badly broken according 10
(1978, p. 80) and Warren Allmon (written comm
cation, September 22, 1993).

uni

CUSPIDARIID AND VERTICORDIID BIVALVES: JUNG 41

Dimensions of holotype of С. islahispaniolae.—
Length 9 mm; height 6 mm (Maury, 1917, p. 196).

Type locality of C. islahispaniolae.—Bluff 3 of

aury on Río Mao, Dominican Republic. Cercado
Formation (late Miocene). This includes NMB locali-
lies 16912, 16913, 17269, and 17307 (Saunders еі al.
1986, text-figs. 29, 33).

Holotype ој С. ornatior.—ANSP 2790. This 15 the
Specimen (a left valve) figured by Pilsbry (1922, pl.
38, fig. 12).

Dimensions of holotype of C. ornatior.—Length 5.0
Mm; height 2.9 mm.

Type locality of C. ornatior.—‘‘Santo Domingo”.
No further details are available.

Paratype of C. ornatior.—ANSP 79015. This is the
Specimen (a left valve) figured by Pilsbry (1922, pl.
38, fig. 11).

Dimensions of paratype of C. ornatior.—Length 4.8
Mm; height 2.7 mm.

Holotype of С. gabbi.—ANSP 2791. This is the
‘Pecimen (a left valve) figured by Pilsbry (1922, pl.
38, fig. 10).

Dimensions of holotype of C. gabbi.—Length 8.0
Mm; height 5.3 mm.

Type locality of C. gabbi.—''Santo Domingo”. No
further details are available.

Paratype of C. gabbi.—ANSP 79016.

Dimensions of paratype of C. gabbi.—Length 7.4
Mm; height 5.2 mm.
une marks. — The 40 available specimens (some of
h ich are incomplete) show variability in rostrum
MI and radial sculpture. The rostrum may be rela-
Е ely long and narrow, or it may be shorter, thus giv-
Ти Ше Impression of greater width. In some specimens

У the posteriormost radial rib is more prominent
w the others. In other specimens there may be up
ifs our more prominent radial ribs with wider inter-
E This variability of the radial sculpture obvi-

Y has been the reason for the introduction of the
‘Mes ornatior and gabbi by Pilsbry and Johnson
(1917, р. 195).

Dalen mentioned in the introduction to the systematic
à ontology, the type specimens of four species of
( подигнаа and Verticordiidae described Бу Dall
iim 3) from the early Pliocene Bowden Formation of
b are available. Three of them occur in the Do-
E Republic as well: Cardiomya (Bowdenia) dis-
" , ација Jamaicensis, and Trigonulina bowdenen-
Selec Or each of these species a lectotype has been
for За (see under those species). The same is done
E и fourth species, Cardiomya (Cardiomya) cras-
и ~ which does not occur in the Dominican Re-
з ІС but is compared with С. islahispaniolae. As is

Case for the other three species the type lot of C.

craspedonia consists of three specimens, which had
been glued to a piece of black paper and cannot be
removed from it without risk of damage. The type
specimens of C. craspedonia (all left valves) are:

1. Lectotype: USNM 135691. Length 4.5 mm; height
2.7 mm. This is the specimen figured by Dall
(1903, pl. 57, fig. 17) and Woodring (1925, pl. 10,
fig. 20).

2. Paralectotype: USNM 482409. Length 3.6 mm;
height 2.3 mm. This is the specimen figured by
Woodring (1925, pl. 10, fig. 22).

3. Paralectotype: USNM 482410. Length 4.3 mm;
height 2.8 mm. This is the specimen figured by
Woodring (1925, pl. 10, fig. 21).

Comparisons.—C. islahispaniolae is similar but
nevertheless clearly distinct from C. craspedonia Dall
(1903, p. 1506, pl. 57, fig. 17) from the early Pliocene
Bowden Formation of Bowden, Jamaica. It is not only
considerably larger (practically twice as large) than C.
craspedonia, but C. craspedonia has more numerous
secondary radial ribs on the main shell disc. In addi-
tion the rostrum of C. craspedonia is proportionately
shorter.

Material.—20 lots with a total of 50 specimens as
listed below:

1. 1 spec., ANSP 2790: holotype of C. ornatior;
“Santo Domingo”.

2. | spec., ANSP 79015: paratype of C. ornatior;
“Santo Domingo”.

3. 1 spec., ANSP 2791: holotype of С. gabbi; “Santo
Domingo”.

4. 1 spec., ANSP 79016: paratype of C. gabbi; ““San-
to Domingo”.

5. 1 spec., NMB locality 16912: Río Mao, Bluff 3
of Maury; Cercado Formation (late Miocene).

6. 4 spec., NMB locality 16913: Río Mao, Bluff 3
of Maury; Cercado Formation (late Miocene).

7. 2 spec., NMB locality 16915: Río Mao, Arroyo
Bajón; Cercado Formation (late Miocene).

8. | spec., NMB locality 16917: Río Mao, Arroyo
Bajón; Cercado Formation (late Miocene).

9. 1 зрес., NMB locality 16922: Río Mao, Arroyo
Bajón; Cercado Formation (late Miocene).

10. 1 spec., NMB locality 16923: Río Mao, Arroyo
Bajón; Cercado Formation (late Miocene).

11. 2 spec., NMB locality 16929: Río Mao; Cercado
Formation (late Miocene).

12. 1 spec., NMB locality 16930: Río Mao, Bluff 2
of Maury; Cercado Formation (late Miocene).

13. 1 spec., NMB locality 15878: Río Gurabo; lower
part of Gurabo Formation (late Miocene).

42 BULLETIN 351

length in mm

= T T
0 2 3 4 5
height in mm

Text-figure 8.—Length/height diagram of Cardiomya (Cardi-
отуа) islahispaniolae (Maury, 1917).

14. 5 spec., NMB locality 15903: Río Gurabo; upper
part of Cercado Formation (late Miocene).

15. 1 зрес., NMB locality 15906: Río Gurabo; upper-
most part of Cercado Formation (late Miocene).

16. 3 spec., NMB locality 15907: Río Gurabo; upper-
most part of Cercado Formation (late Miocene).

17. 2 spec., NMB locality 16817: Río Cana, Cañada
de Zamba; lower part of Gurabo Formation (early
Pliocene).

18. 12 зрес., NMB locality 16837: Río Cana; upper-
most part of Cercado Formation (late Miocene).

19. 1 зрес., NMB locality 16838: Río Cana; upper-
most part of Cercado Formation (late Miocene).

20. 10 spec., TU locality 1294 (= NMB locality
18556): Río Mao, Bluff 3 of Maury; Cercado For-
mation (late Miocene).

Measurements.—Plotted in Text-figure 8.
Occurrence.—This species is recorded from the fol-
lowing areas: Río Mao: Cercado Formation (late Mio-

cene): NMB localities 16912, 16913, TU locality 1294
(= Bluff 3 of Maury); 16915, 16917, 16922, 16923
(all Arroyo Bajón); 16929, 16930 (= Bluff 2 of Mau-
ry) (Saunders et al., 1986, text-figs. 29, 30, table 3):
Río Gurabo; lower part of Gurabo Formation (late
Miocene): NMB locality 15878. Upper part of Cercado
Formation (late Miocene): NMB localities 15903,
15906, 15907 (Saunders ег al., 1986, text-figs. 4, 6).

Río Cana; lower part of Gurabo Formation (early
Pliocene): ММВ locality 16817. Uppermost part of
Cercado Formation (late Miocene): NMB localities
16837, 16838 (Saunders et al., 1986, text-figs. 15, 16).

Distribution.—Not known from outside the Domin-
ican Republic.

Subgenus BOWDENIA Dall, 1903
Bowdenia Dall, 1903, p. 1504.

Type species (by original designation and mono“
typy).—Cuspidaria (Bowdenia) distira Dall. Bowden»
Jamaica. Bowden Formation (Pliocene).

Diagnosis.—Shell small (around 3 mm in length)
rostrate. Umbos prosogyrate, almost centrally placed:
Sculpture of fine, more or less well developed radia
ribs. Rostrum well set off from main shell disc by ||
more prominent radial rib. Ventral margin of таш
shell disc evenly rounded. Hinge of left valve consist
ing of an inconspicuous posterior lateral tooth situate
just behind a cavity to receive the posterior cardinal
tooth of the right valve, which in turn is situated be
hind the subumbonal chondrophore. Hinge of right
valve consisting of а subumbonal chondrophor®, 8
weakly developed anterior cardinal tooth, and a prom
inent posterior cardinal tooth. Postero-dorsal margin of
left valve somewhat thickened to fit the groove of Ше
postero-dorsal margin of the right valve.

Remarks.—So far the type species of Bowdenia, В.
distira Dall, is the only species assigned to this sub”
genus. Bowdenia is therefore known only from t
Pliocene Bowden Formation of Bowden, Jamaica, an
the late Miocene part of the Gurabo Formation, рој
minican Republic (see below).

Cardiomya (Bowdenia) distira (Dall, 1903)

Plate 3, figures 1-5; Plate 4, figures 1-5;
Text-figure 9

ll
Cuspidaria (Bowdenia) distira Пай, 1903, р. 1506, pl. 57, fig. !
Woodring, 1925, p. 91, pl. 11, figs. 1—5.

Description.—Shell small (around 3 mm in length)
delicate, rostrate. Umbos prosogyrate, placed almo?
centrally. Sculpture consisting of fine radial та
which may Бе fairly well developed, but sometimó
hardly recognizable. Rostrum well set off from та!
shell disc by а more prominent radial пр. The conc?

CUSPIDARIID AND VERTICORDIID BIVALVES: JUNG 43

Part of the rostrum of the left valve sometimes car-
Tying a few fine radial riblets, but corresponding area
of right valve does not. Concave part of rostrum ad-
Joining postero-dorsal margin carrying three or four
ribs. Ventral margin of main shell disc evenly rounded.
Growth lines usually more clearly developed on right
Valve, Hinge of left valve consisting of an inconspic-
"ous posterior lateral tooth behind a cavity to receive
\ е posterior cardinal tooth of right valve. This cavity
жі Situated behind the subumbonal chondrophore.

Inge of right valve consisting of a subumbonal chon-
drophore, a weakly developed anterior cardinal tooth,
and a prominent posterior cardinal tooth. Postero-dor-
sal margin of left valve somewhat thickened to fit
8'00ve of postero-dorsal margin of right valve.

. Lectotype (selected herein).—USNM 135692. This
5 the specimen figured by Dall (1903, pl. 57, fig. 16)
and Woodring (1925, pl. 11, fig. 1), a left valve.

Dimensions of lectotype.—Length 3.3 mm; height

5 mm.

: Type locality.—Bowden, Jamaica. Bowden Forma-
tion (early Pliocene).

Paralectotype.—USNM 482411. This is the speci-
Men figured by Woodring (1925, pl. 11, figs. 2, 3), a
“It valve,

Dimensions ој paralectotype USNM 482411.—

ength 3.1 mm; height 2.3 mm.

Paralectotype.—USNM 482412. This is the speci-
Men figured by Woodring (1925, pl. 11, figs. 4, 5), a
“ght valve.

Dimensions of paralectotype USNM 482412.—
ength 3.0 mm; height 2.0 mm.

Remarks, — The type lot of this species consists of
i» lectotype and the two paralectotypes. All three
қары аге glued to а piece of black paper. Origi-

УІ intended to remove the specimens from the
ack Paper in order to refigure the exterior of the lec-
p^ and to figure its interior. Various chemicals have
ро Used in an attempt to dissolve the glue but with-
Қ Success. The lectotype is therefore not refigured.

Stead several topotypes are figured (Pl. 3, fig. 5; Pl.
> figs, 1-5),

к” Dominican Neogene has yielded eight speci-
ir 8 Of this species. As hinted at in the above de-
'Plión there is some variability in the development
one radial ribs. They may be stronger or weaker.
^ *ümes they are evenly developed over the entire
lene Shell disc, sometimes they are restricted to the
tral part of the main shell disc.
is the Prisons. —As Cardiomya (Bowdenia) distira
E * only species of the subgenus known no com-

'Sons can be made.

@terial.—Eight lots with a total of 47 specimens
Sted below:

to

о

as Ji

4-
е
е
е 222% T
ee
ш? 4E
А Au
Б
5 27
c
2
1-
Т Т Т
0 1 2 3
height in mm

Text-figure 9.—Length/height diagram of Cardiomya (Bowdenia)
distira (Dall, 1903).

1. 3 spec., USNM 135692 (lectotype) and two рага-
lectotypes (USNM 482411, 482412). Bowden, Ja-
maica. Bowden Formation (early Pliocene).

2. 1 spec., NMB locality 15846: Río Gurabo, Domin-
ican Republic. Latest Miocene part of Gurabo For-
mation (Pl. 3, figs. 1-4).

3. 5 spec., TU locality 1227A (= NMB locality
18582): Arroyo Zalaya, Dominican Republic; Glo-
borotalia margaritae zone (early Pliocene).

4. 1 spec., TU locality 1352 (= NMB locality 18584):
Rio Gurabo, Dominican Republic; middle Pliocene
part of Mao Formation.

5. 1 spec., TU locality 1403 (= ММВ locality 18586):
Arroyo Babosico near Rio Yaque del Norte at La
Barranca; upper part of Globorotalia margaritae
zone (late early Pliocene).

6. 33 spec., NMB locality 10635: Bowden, Jamaica.
Bowden Formation (early Pliocene).

7. 1 spec., NMB locality 11146: Bowden, Jamaica.
Bowden Formation (early Pliocene).

8. 2 spec., NMB locality 17617: Bowden, Jamaica.
Bowden Formation (early Pliocene).

Measurements.—Plotted in Text-figure 9.

Occurrence.—Rio Gurabo section: latest Miocene
part of Gurabo Formation: NMB locality 15846 (Saun-
ders et al., 1986, text-figs. 4, 6); middle Pliocene part
of Mao Formation: TU locality 1352. Globorotalia
margaritae zone (early Pliocene) of Arroyo Zalaya
(TU locality 1227A) and Arroyo Babosico (TU local-
ity 1403).

Distribution.—Bowden Formation (early Pliocene)
of Bowden, Jamaica. Latest Miocene part of Gurabo

44 BULLETIN 351

Formation; Globorotalia margaritae zone (early Plio-
cene); middle Pliocene part of Mao Formation, north-
ern Dominican Republic.

Genus PLECTODON Carpenter, 1864
Plectodon Carpenter, 1864, pp. 611, 638.

Type species (by original designation and mono-
typy).—Plectodon scaber Carpenter, 1864, pp. 611,
638. Recent, Catalina Island, California, to Santa Inez
Bay, east coast of Baja California, Mexico (Palmer,
1958, p. 80). Throughout the Gulf of California and
south to Panama and the Galápagos Islands, Ecuador,
in 20 to 250 m (Keen, 1971, p. 302).

Diagnosis.—Shell of medium to large size (up to 24
mm in length), rostrate, moderately delicate. Antero-
ventral margin evenly rounded. Exterior surface of
shell covered by pustules. Resilium situated posterior
to the umbo. Dorsal margin of left valve twisted just
anterior to the umbo forming a small, toothlike pro-

jection. No lateral teeth in left valve. Right hinge with `

an anterior and a posterior lateral tooth.

Remarks.—The holotype of P. scaber (USNM
592441) unfortunately is broken into several fragments
(see also Palmer, 1958, p. 80). Originally it was
mounted on a piece of glass. The fragmentation prob-
ably happened when the specimen was removed from
the glass. Other material of P. scaber is figured here
(РІ. 2, figs. 5-8) for comparison with P. granulatus.

The stratigraphic range of the genus Plectodon is
given as Pliocene to Recent (Dall, 1903, p. 1507). The
single right valve from the Dominican Republic de-
scribed below has been collected from sediments of
late Miocene age, and the single valve from the middle
Miocene Shoal River Formation of Florida reported by
Gardner (1926, p. 64) as Cuspidaria (Plectodon) cf.
granulata Dall extend the range of Plectodon from
middle Miocene to Recent.

Plectodon granulatus (Dall, 1881)
Plate 2, figures 9, 10; Plate 5, figures 1-4; Plate 6,
figures 1-4; Plate 7, figures 1-4; Text-figure 10

Neaera granulata Dall, 1881, p. 111.

Leiomya (Plectodon) granulata Dall. Dall, 1886, p. 300, pl. 3, fig.
8; Dall, 1889, p. 66, pl. 3, fig. 8.

Cuspidaria (Plectodon) granulata Dall. Dall, 1903, p. 1507.

? Cuspidaria (Plectodon) cf. granulata Dall. Gardner, 1926, p. 64.

Cuspidaria (Plectodon) scabrata Olsson and Harbison, 1953, p. 67,
pL ие, 2;

Plectodon granulatus (Dall, 1881). Knudsen, 1982, p. 136.

Description.—Shell of medium size (up to 18 mm
in length), rostrate, moderately delicate. Antero-ventral
margin evenly rounded. Umbos prosogyrate. Exterior
surface of shell covered by pustules. In the umbonal
area there are fewer or no pustules; instead the growth

lines are more clearly developed. Resilium located be-
hind the umbo and somewhat toward the interior of
the shell. Dorsal margin of the left valve twisted just
in front of the umbo, forming a small, toothlike рго-
jection. No lateral teeth in left valve. Right hinge with
an anterior and a posterior lateral tooth.

Lectotype (selected herein). —USNM 63193 (РІ. 2,
figs. 9, 10).

Dimensions of lectotype.—Length 11.3 mm; height
6.7 mm.

Туре locality.—Off Sombrero Island, Leeward 15°
lands, Lesser Antilles, in 132 m (72 fathoms).

Remarks.—Lot USNM 63193 contains three spect
mens, the syntypes of P. granulatus. Тһе specimen
chosen as the lectotype is the left valve figured by Dall
(1886, pl. 3, fig. 8). One of the paralectotypes is a right
valve, the other paralectotype is also a left valve like
the lectotype, but is considerably smaller. The two
paralectotypes are USNM 887025 (ex USNM 63193).

The Dominican Neogene so far has yielded a single
specimen of this species, a right valve. It is not quite
complete: its postero-dorsal margin is somewhat dam“
aged (РІ. 5, бе. 1). Unfortunately this unique specimen
has been broken during handling for scanning electron
microscopy (Pl. 5, fig. 3). Olsson and Harbison (1953.
p. 67) state that their P. scabratus from the Pliocene
of Fort Thompson, Florida, is less “narrow” than Re
cent specimens of P. granulatus. It is not clear, how
ever, what is meant by “narrow”. It probably refers E
the ratio of height to length. In that respect there P
some variability in P. granulatus. Two Recent зресг
mens of P. granulatus are figured here for compariso"
(РІ. 6, figs. 1-4; Pl. 7, figs. 1-4).

Comparisons.—The only other species of Plectodo"
is the Recent Eastern Pacific P. scaber Carpente"
(1864, pp. 611, 638) (for figures see Schenck, 194
pl. 67, figs. 1-4; Palmer, 1958, pl. 6, figs. 6-8; Keen
1971, p. 302, fig. 786; and Pl. 2, figs. 5-8). The mai"
differences between the two species is size. P. scabe!
is considerably larger; it actually may be twice as larg
as P. granulatus.

Material.—Twelve lots with a total of 22 specimé
as listed below:

19

1. 1 spec., ММВ locality 16857: Río Cana, Domi
ican Republic; Cercado Formation (late Міюселе”

2. 1 spec., USNM 63193: lectotype. Recent; so
brero Island, Leeward Islands, Lesser Antilles»
fms.; Blake Coll. t

3. 2 spec., USNM 887025: paralectotypes. Кесин
Sombrero Island, Leeward Islands, Lesser And
les, 72 fms.; Blake Coll. 100

4. 1 spec., USNM 63194: Recent; Barbados,
fms.

CUSPIDARIID AND VERTICORDIID BIVALVES: JUNG 45

5. 6 spec., USNM 94214: Recent; Station 2648: off
Cape Florida, 84 fms., sand.

без врес., USNM 667843: Recent; Station 1306:
Campeche Bank off Yucatan, Mexico (22°10'N,
91°40'W), 42 fms., sand.

kel spec., USNM 667668: Recent; Station 470:
Campeche Bank off Yucatan, Mexico (22°30'N,
90°15'W), 46 fms., sand.

б] spec., USNM 157815: Recent; Station 2404: be-
tween Mississippi Delta and Cedar Keys, Gulf of
Mexico; 60 fms., sand.

9. 1 spec., USNM 64003: Recent; Station 2646: off
Cape Florida; 85 fms.

10. 3 spec., USNM 97157: Recent; Station 2646: 5
miles off Cape Florida, Gulf of Mexico; 85 fms.,
sand.

B. 1 spec., USNM 157986: Recent; Station 2646: Re-
cent; off Cape Florida, Gulf of Mexico; 85 fms.,
sand.

12. 1 spec., USNM 667737: Recent; Station 1241:
Campeche Bank off Yucatan, Mexico (20?15'N,
92°10'W), 32 fms., sand.

Measurements.—Plotted in Text-figure 10.

Depth range.—From 37 to 274 m (Knudsen, 1982,
7137).

Occurrence.—Cercado Formation (late Miocene) of
Rio Cana section: NMB locality 16857 (Saunders er
al, 1986, text-figs. 15, 16).

Distribution.—Shoal River Formation (middle Mio-
cene), Florida? Cercado Formation (late Miocene),
tern Dominican Republic. Pliocene, Florida. Re-
“nt, southern Florida and Gulf of Mexico throughout
the West Indies,

Family VERTICORDIIDAE Stoliczka, 1871
Genus HALIRIS Dall, 1886
Нау» Dall, 1886, p. 287.

Type species (by original designation).—Verticor-
44 fischeriana Dall, 1881. Recent, Gulf of Mexico.
orth Carolina to Gulf of Mexico to Barbados (Ab-
tt, 1974. р. 563).
lagnosis.—Shell small (up to 7 mm in length),
8 ОБове, Umbos strongly prosogyrate. Lunule some-
at depressed. Ventral margin evenly rounded or
Mewhat angulated near its middle. Entire surface
кіріге by numerous radial ribs. Мо secondary ra-
its ribs. Surface granulated. Interior surface nacreous,
Ventral margin fluted. Hinge of left valve consisting
а hardly recognizable, subumbonal cardinal tooth
За (only in fully adult shells) an inconspicuous pos-
‚ Or lateral tooth. Hinge of right valve with a prom-

In А :
а Subumbonal сагата! tooth and а posterior lateral

50
Se

an

14-

13:5

12 4 +

11”

length in mm
е

10 4 ага

Т Т

0 5 6 7 8
height in mm

Text-figure 10.—Length/height diagram of Plectodon granulatus
(Dall, 1881).

Remarks.—Numerous Recent specimens of the type
species, H. fischeriana, are available. Some of them
are figured (Pl. 8, figs. 1-6) for comparison with H.

Jamaicensis. The stratigraphic range of the genus is

Eocene to Recent.

Haliris jamaicensis (Dall, 1903)
Plate 9, figures 1—6; Plate 10, figures 1—4

Verticordia (Haliris) jamaicensis Dall, 1903, p. 1511; Woodring,
1925, p. 93, pl. 11, figs. 9-11.

Description.—Shell small (up to less than 5 mm in
length), globose. Umbos strongly prosogyrate. Lunule
depressed. Ventral margin slightly angulated near its
middle. Surface sculptured by 23 to 26 radial ribs; in-
terspaces narrower on anterior part of shell. Whole sur-
face granulated. Inner surface nacreous, its ventral
margin fluted. Hinge of left valve consisting of a hard-
ly recognizable, subumbonal cardinal tooth and (only
in fully adult shells) an inconspicuous, posterior lateral

46 BULLETIN 351

tooth. Hinge of right valve with a prominent, subum-
bonal cardinal tooth and a posterior lateral tooth.

Lectotype (selected herein). —USNM 135686. This
is the specimen figured by Woodring (1925, pl. 11, fig.
10), a right valve.

Dimensions of lectotype.—Length 4.5 mm; height
4.7 mm.

Type locality.—Bowden, Jamaica. Bowden Forma-
tion (early Pliocene).

Paralectotype.—USNM 482413. This is Ше speci-
men figured by Woodring (1925, pl. 11, fig. 9), a right
valve.

Dimensions of paralectotype USNM 482413.—
Length 3.8 mm; height 3.8 mm.

Paralectotype—USNM 482414. This is the speci-
men figured by Woodring (1925, pl. 11, fig. 11), a left
valve.

Dimensions of paralectotype USNM 482414. —
Length 3.3 mm; height 3.1 mm.

Remarks.—The lectotype of И. jamaicensis is the
largest of the 10 available specimens. Its height is
greater than its length. In all the other specimens the
length is greater (or the same as) than the height.

The type material of this species consists of the lec-
totype and the two paralectotypes mentioned above.
All three specimens are glued to a piece of black paper.
As explained under Cardiomya (Bowdenia) distira it
has not been possible to remove the specimens from
the black paper. Therefore the lectotype is not refigu-
red Bere.

Woodring (1925, p. 93) mentioned a fragment from
the early Pliocene Bowden Formation of Jamaica that
is almost three times as large as the specimens listed
below under “Material”. More specimens from Bow-
den would be needed in order to be able to identify
the fragment mentioned above.

Comparisons.—H. jamaicensis is obviously closely
related to the living H. fischeriana (Dall) (1881, p.
106), the type species of the genus. H. fischeriana is
larger than H. jamaicensis and has more radial ribs. In
addition the ventral margin is evenly rounded in H.
fischeriana but somewhat angulated in H. jamaicensis.

Material.—Four lots with a total of only ten speci-
mens as listed below:

1. 2 spec., NMB locality 15832: Río Gurabo, Domin-
ican Republic; middle Pliocene part of Mao For-
mation.

2. 1 spec., TU locality 1227A (= NMB locality
18582): Arroyo Zalaya, Dominican Republic; Glo-
borotalia margaritae zone (early Pliocene).

3. 4 spec., NMB locality 10635: Bowden, Jamaica;
Bowden Formation (early Pliocene).

4. 3 spec., USNM 135686 (lectotype) and two para-

•
..
а
•
•
e
өө
слиза е
=
Е
~
+
P 24
Ф
14
Ew
T Si T T
0 1 2 3 4
height in mm
Text-figure 11.—Length/height diagram of Haliris jamaicensis
(Dall, 1903).

lectotypes (USNM 482413, 482414). Bowden, Ja-
maica; Bowden Formation (early Pliocene).

Measurements.—Plotted in Text-figure 11.

Occurrence.—Globorotalia margaritae zone (early
Pliocene) of Arroyo Zalaya: TU locality 1227A and
middle Pliocene part of Mao Formation of Río Gur abo
section: ММВ locality 15832 (Saunders et al., 1986,
text-figs. 4, 6).

Distribution.—Bowden Formation (early Pliocene)
of Bowden, Jamaica. Early Pliocene Globorotalid
margaritae zone and middle Pliocene part of Mao For
mation, Dominican Republic.

Genus TRIGONULINA d'Orbigny, 1842
Trigonulina d'Orbigny, 1842 (see also d’Orbigny, 18457, p. 327).

Type species (by monotypy).—Trigonulina ornata
d’Orbigny, 1842 (see also d’Orbigny, 18457, p. 327).
Recent, Massachusetts to Florida and Ше West Indies:
Bermuda, Brasil (Abbott, 1974, p. 563). |

Diagnosis.—Shell small (up to almost 6 тт m
length), oval. Umbos low, strongly prosogyrate. Ви
пше deeply depressed. Sculpture consisting of fom
widely spaced, high, and narrow radial ribs projecting
beyond ventral margin. On the posterior slope there |
а large area without radial ribs. Interior surface пасте“
ous, its ventral margin fluted. Ligament internal.
teeth in left valve. Right valve with a strong, projec”
ing, subumbonal, cardinal tooth and a groove alone
postero-dorsal margin to receive postero-dorsal marg!
of left valve.

CUSPIDARIID AND VERTICORDIID BIVALVES: JUNG 47

Remarks.—The Spanish edition of Ramón de la Sa-
gras Historia física, política y natural de la Isla de
Cuba, volume 5 (molluscs) of the second part (natural
history), is dated 1845. On page 327, where Trigon-
ulina and its type species, T. ornata, are described, the
date is given as 1846. According to Aguayo (1943, p.
38) publication of this edition appears to have started
ІП 1844 already and was probably completed only in
1853, Aguayo is quoting the Spanish edition as of
18452, to which the present author is adding [1844-
18537] in the "References Cited". Dall (1889, p. 18)
did not have access to the Spanish edition.

Both Dall (1889, p. 18) and Aguayo (1943, p. 38)
Commented on the dates of publication of the French
edition. Both authors state that the atlas was published
ІП 1842, The figures of T. ornata given in this atlas
(Pl. 27, figs. 30-33) are an indication as defined in
Article 12b(7) of the International Code of Zoological

omenclature (third edition, 1985).

Trigonulina is here used as a full genus, whereas
Abbott (1974, p. 563) and Woodring (1925, p. 92)
treated it as a subgenus of Verticordia J. Sowerby
(1812-1846, p. 68, pl. 639, 1844 [for date of publi-
Cation of plate 639 see Renevier, 1855, and Sykes,
1906]. On the other hand Keen (1971, p. 302) consid-
ered Trigonulina as a synonym of the subgenus Ver-
"cordia s.s. The type species of Verticordia is V. car-
diiformis y. Sowerby (1812-1846, p. 68, pl. 639, 1844)
rom the Pliocene of England. The original figure
Shows that the 13 radial ribs are evenly distributed
dl the entire shell disc. In Trigonulina, however,

еѓе is a space without radial ribs on the postero-
dorsal slope.

T. ornata d’Orbigny, the type species of Trigonuli-
а 18 not only reported from Western Atlantic waters,

ut is also said to occur in the Eastern Pacific (Keen,
En р. 302). Having looked at a number of lots from
Oth Oceans I come to the conclusion that they are
n and that the species from the Eastern Pacific
| tote needs a name. For this reason these two liv-
8 Species are briefly discussed and compared below.

he Stratigraphic range of Trigonulina is Eocene to
Recent.

p Trigonulina ornata d'Orbigny, 1842
late 11, figures 1-4; Plate 12, figures 1-4; Text-
figures 12, 13

Tri "
Eos ornata d'Orbigny, 1842, pl. 27, figs. 30-33; 18457, p.

Уер.
шш caelata Verrill, 1882, р. 566; 1884, р. 278, pl. 30, figs.
» да,

Verton,
міз (Trigonulina) ornata D’Orbigny. Dall, 1886, р. 290
E Dall and Simpson, 1901, p. 498 (part). (For further citations
“е Dall, 1886.)

Verticordia (Trigonulina) ornata (Orbigny, 1842). Abbott, 1974, p.
563, fig. 6158.

Verticordia ornata (Orbigny, 1846). Knudsen, 1982, p. 128 (part).
For additional citations see this publication.

Verticordia ornata (Orbigny, 1842). Rios, 1985, p. 282, pl. 99, fig.
1390 (part).

Description.—Shell small (up to 5 mm in length),
oval. Umbos low, strongly prosogyrate. Lunule deeply
impressed in both valves, but more so in left valve.
Sculpture consisting of eight to twelve high, narrow,
radial ribs anterior to the unsculptured posterior slope
projecting beyond the ventral margin. Surface of per-
fectly preserved valves covered by minute pustules
forming rows parallel to the ribs. Interior surface na-
creous, its ventral margin fluted. No teeth in left valve.
Right valve with a strong, subumbonal, cardinal tooth
and a groove along postero-dorsal margin to receive
postero-dorsal margin of left valve.

Holotype.—BMNH Cat. no. 493; Reg. no.
1854.10.4.557, a left valve.

Dimensions ој holotype.—Length 2.6 mm; height
2.3 mm.

Туре locality.—''Jamaica" (from sand). This is the
only information given with the original description.

Remarks.—The holotype of 7. ornata 15 at hand. Its
ventral margin is somewhat damaged. It is a left valve.
It is the specimen figured in an idealized way and in
mirror-image by d’Orbigny (1842, pl. 27, figs. 30, 31).
His Figure 31 shows the interior of the valve with a
hinge without teeth, a hinge typical for left valves. The
holotype is one of the rare cases of a specimen having
only eight radial ribs in front of the posterior slope.
Due to its imperfect preservation it is not refigured
here.

As listed under “Material”, six lots with 1049 spec-
imens have been used for the description given above.
Out of these 1049 specimens 190 valves have been
measured (Text-fig. 12) and their ribs in front of the
unsculptured posterior slope counted (Text-fig. 13). As
indicated in the description the range of the number
of ribs is eight to twelve. However, the extremes are
rare (Text-fig. 13); there are only three valves with
eight ribs, eight valves with nine ribs, and twelve
valves with twelve ribs. In other words one should
really describe 7. ornata as having ten or eleven ribs.

The original description of Verticordia caelata Ver-
rill was based on a single right valve with eleven ribs.

Comparisons.—Comparative remarks are given be-
low under 7. pacifica and T. bowdenensis.

Material.—Six lots with a total of 1049 specimens
as listed below (quoted from specimen labels):

1. 439 spec., USNM 444664: Eolis Station 368: off
Ajax Reef, Florida; 80-100 fms.

48 BULLETIN 351

length in mm
ЕЛ

Т Т T

0 1 2 3 4.
height іп mm

Text-figure 12.—Length/height diagram of Trigonulina ornata
d’Orbigny, 1842.

2. 54 spec., USNM 444479: Eolis Station 178: off
Fowey Light, Florida; 68 fms.

3. 3 spec., USNM 63214: off Hatteras; 15-124 fms.

4. 172 spec., USNM 444514: Eolis Station 311: off
Govt. cut, Miami, Florida; 75 fms.

5. 337 зрес., USNM 444665: Eolis Station 370: off
Ajax Reef, Florida; 70-90 fms.

6. 44 spec., USNM 444653: Eolis Station 363: off
Fowey Light, Florida; 85 fms.

Measurements.—Plotted in Text-figure 12.

Depth range.—From 5 to 850 m (Knudsen, 1982,
p. 128) and 15 to 1256 m (Hertlein and Grant, 1972,
p. 344).

Distribution.—Massachusetts to Florida and the
West Indies, Bermuda, Brazil (Abbott, 1974, p. 563)
or from about 42%N to about 30%42'S (Knudsen, 1982,
p. 128). So far T. ornata has not been reported as a
fossil.

Trigonulina pacifica, new species
Plate 2, figures 11, 12; Plate 13, figures 1-4;
Plate 14, figures 1-4; Text-figures 14, 15

Verticordia ornata (d’Orbigny). Grant and Gale, 1931, р. 266, pl.
13, fig. 4 (part).

Verticordia (Verticordia) ornata (Orbigny, 1846). Keen, 1971, p.
302, fig. 789 (part).

Verticordia (Trigonulina) ornata d’Orbigny. Hertlein and Grant,
1972, p. 344, pl. 43, figs. 23, 26, 27, 31.

80 -
o
5 604
Е
о
Ф
e.
o
5
5 40-
Q
Е
-
c
20-
--

1
8 9 10 14 12
number of ribs

Text-figure 13.—Histogram showing rib number distribution о!
Trigonulina ornata d'Orbigny, 1842.

Description.—Shell small (up to almost 6 mm іп
length), oval. Umbos low, strongly prosogyrate. Lus
nule deeply impressed in both valves, but more 50 10
left valve. Sculpture consisting of six to 12 high, nar
row radial ribs anterior to the unsculptured posterior
slope projecting beyond the ventral margin. Surface 5
perfectly preserved valves covered by minute, no
closely spaced pustules forming rows parallel to the
ribs. Interior surface nacreous, its ventral margin flut-
ed. No teeth in left valve. Right valve with a strong»
subumbonal, cardinal tooth and a groove along pO%
tero-dorsal margin to receive the postero-dorsal margi”
of left valve.

Holotype. АСМ 2718 (РІ. 2, figs. 11, 12).

Dimensions of holotype.—Length 4.8 mm; heig
4.2 mm.

Type locality.—(quoted from specimen label
LACM 65-6.22: 0.4-0.7 miles 110 to 132 degrees
from Ship Rock, Santa Catalina Island, California
Channel Islands, California (33°27'N, 118°30'W”
Depth: 82 m.

Remarks.—The basis for the above description СО
sists of the 23 lots with 1840 specimens listed UN
“Material”. A total of 320 specimens have been me?
sured (Text-fig. 14) and their ribs in front of the rible?

ht

)

CUSPIDARIID AND VERTICORDIID BIVALVES: JUNG 49
6 -
160 y
е
е
ее
5+ АНН
к e
“ 140-
4- е
е
Е 2
Е :
с На
5 34 өөө 120-
е 2200009
Ф HIT
е е.
ее
МА
2- e
57
100-
E g
Ф
Е
Ф
= 80-
o
"mou T T T T 5
0 2 3 4 5 5
о
height in mm Е
7 | с 604
9xt-figure 14.—Length/height diagram of Trigonulina pacifica,
New species.
ле slope counted (Text-fig. 15). The range of the
Umber of ribs as given in the description is six to 12. 407
ut again—as in Т. ornata—the extremes are rare:
Ко are only 16 valves with six ribs, 17 valves with
» ribs, six valves with 11 ribs, and а single valve
Ith 12 ribs, It is therefore appropriate to say that the 20-
Steat majority of the valves has seven to nine ribs.
he ribs of 7. pacifica as a rule are high and narrow.
б. Sometimes there are exceptions with lower and
OMewhat broader ribs. |
‚ Comparisons. —T. pacifica reaches larger dimen-

Моп than the Recent West Indian T. ornata d'Orbigny
A has fewer radial ribs. In addition the general out-
and n the shell is more rounded or oval іп 7. ornata,
за T area without ribs on the posterior slope of T.
миң 15 wider than that of 7. pacifica. Т. pacifica 15
а d larger than T. bowdenensis Dall from the
à ^ liocene Bowden Formation of Jamaica but has
ЧЕ the same number of radial ribs.

D microsculpture of T. ornata and T. pacifica are
das In both species the microsculpture
tows sts of rounded pustules, which are aligned in

parallel to the ribs; but in T. ornata they are

E more closely spaced (Pl. 12, Figs. 2, 4; Pl. 14,
85. 2, 4). In T. bowdenensis on the other hand the

о У
~

8 9 1952741 12

number of ribs

Text-figure 15.—Histogram showing rib number distribution of
Trigonulina pacifica, new species.

pustules are more closely spaced than in 7. pacifica
but not as closely as in 7. ornata (Pl. 18, Figs. 2, 4).
However, the pustules of 7. bowdenensis have a dif-
ferent shape; they are not rounded but pointed (Pl. 16,
Figs. 3, 5).

The diagnostic features of the three species of Tri-
gonulina discussed above are tabulated in Table 3.

Although there are clear differences in the micro-

BULLETIN 351

Table 3.—Diagnostic features of the three species of Trigonulina dealt with herein. Numbers of measured specimens: T. ornata: 190; Т.
pacifica: 320; Т. bowdenensis: 79.

теап ribless most microsculpture
ratio area on range of frequent depth
maximum maximum length/ posterior number number spacing of form of range
length height height slope of ribs of ribs pustules pustules (in m)
T. ornata 5.0 4.9 1:13 wide 8-12 11 close rounded 5-1250
T. pacifica 5.8 22 1.09 narrower 6-12 Y wide rounded 18-168
T. bowdenensis 4.0 S9 1.16 wide 7-10 10 not wide pointed --

sculpture of these species, considerably more numer-
ous, well preserved specimens should be looked at in
the scanning electron microscope in order to determine
the range of variability of the microsculpture.

sand and gravel, W of Isla Smith, Bahía de 105
Angeles, Gulf of California, Mexico (29%03.7'N,
113°31.0’W). Leg. Gale Sphon, О.К. Mulliner
10-16 May 1976.

Material. —23 lots with a total of 1840 specimens 9. 25 spec., paratypes. LACM 71-158.38: 31-37 m»
as listed below (lots arranged from north to south) shelly sand, Kellett Channel, S of Isla Cedros, Pa-
(quoted from specimen labels): cific Coast, Baja California, Mexico (27°57.0'N,

1. 3 spec., paratypes. LACM 63-50.12: 60 m, Hump- Петя Dep DEI ME ME

back Rock, off Hopkins Marine Station, Pacific R/V Searcher, 20 October 1971.
Grove, Monterey Bay, California (36°38'N, 10. 24 spec., paratypes. LACM 78-120.18: 43-99 г
121°54’W). Leg. J.H. McLean, R/V Tage, 26 No- sandy, off Isla Danzante, Bahía Escondido, Gu
veraber 1063. of California, Baja California Sur, Mexico
2. 3 spec., paratypes. LACM 41-80.19: 99-102 m, (25°46’N, 111*15'W). Leg. D. Mulliner, G. Spho™
mud, sand and shell, 1.5 mi NW of Cavern Pt., 6 November 1978. Е 2
Santa Cruz Id., California Channel Ids., California 1. 384 ЖҮ” paratypes. USNM 211469: off La Ра?
(34°04'N, 119°34.4’W). Leg. R/V Velero Ш (АНЕ BAS SAP MENO: Че КОЕ, „ои
1300-41), 12 April 1941. Ex AHE 12. 623 spec., paratypes. USNM 211458: off La Paz,
3. 2 spec., paratypes. LACM 40-164.20: 27-91 m, Baja California, Mexico; 26% fms. 2
sand and gravel, Anacapa Passage, W of Anacapa 13. 591 spec., paratypes. USNM 151959: near La Pa
Id., California Channel Ids., California off Baja California, Mexico; 9%-10 fms.
(33°59.0'N, 119°32.1’W). Leg. R/V Velero III 14. 13 spec., paratypes. LACM 66-23.22: 27-31 T
(AHF 1190-40), 30 October 1940. Ex АНЕ sand, off Punta Arena de la Ventana, Gulf of СЕ
4. 1 зрес., paratype. LACM 41-74.19: 62-75 m, sand ifornia, Baja California Sur, Mexico (24704 |:
and shell, 0.5 mi S of Gull Id., Santa Cruz Id., 109°49’W). Leg. J.H. McLean, P. M. Oringeb 7
California Channel Ids., California (33°56.5'N, Marincovich, 8 April 1966.
119°49.6’W). Leg. R/V Velero Ш (АНЕ 1294-41), 15: 6 spec., paratypes. LACM 66-22.40: 18-55 T^
11 April 1941. Ex АНЕ sand and shell, directly off anchorage at Bahía de
5. 8 spec., holotype and 7 paratypes. LACM 65-6.22: los Muertos, Gulf of California, Baja California
82 m, 0.4-0.7 mi 110 to 132 degrees T from Ship Sur, Mexico (23%58'N, 109*46'W). Leg. J.H. МС
Rock, Santa Catalina Id., California Channel Ids., Lean et al., 8 April 1966.
California (33°27'N, 118°30'W). Leg. В. Reimer 16. 6 spec., paratypes. LACM 66-17.62: 18-37 "h
et al., R/V Velero IV, 13 February 1965. Ex АНЕ sand, between Rancho El Tule and Rancho Pal
6. 4 spec., paratypes. LACM 41-25.17: 75 m, shell, milla, Gulf of California, Baja California 55"
mud and gray sand, 4 mi N of Islas Todos Santos, Mexico (22°58'N, 109745" W). Leg. J.H. McLean
Pacific Coast, Baja California, Mexico (31%53.3'N, P.M. Oringer, 5 April 1966.
116°48.3'W). Leg. R/V Velero Ш (АНЕ 1245-41), 17. 10 spec., paratypes. LACM 38-5.9: 37-73 M, n
24 February 1941. Ex AHE Һа Banderas, Jalisco, Mexico (20°40 ^"
7. 12 spec., paratypes. LACM 75-93.17: 27 m, grav- 105°25'W). Leg. С. Willett, 14 February 1938.
el and shell, W of Isla Smith, Bahía de los An- 18. 31 spec. paratypes. LACM 34-2.20: 26-33 M
geles, Gulf of California, Mexico (29?04'N, sand, nullipores, Bahía Braithwaite, Isla Socorro
113°33'W). Leg. Gale Sphon, D.K. Mulliner, 10 Islas Revilla Gigedo, Mexico (18%42-5 №
October 1975. 110%56.22'W). Leg. R/V Velero Ш (AHF 129-34)
8. 21 spec., paratypes. LACM 76-2.21: 18-22 m, 3 January 1934. Ex AHE

оп ат па

CUSPIDARIID AND VERTICORDIID BIVALVES: JUNG 51

19. 8 5рес., paratypes. LACM 38-9.11: 73-128 m, Ва-
hía Guatulco, Oaxaca, Mexico. Leg. G. Willett, 7
March 1938.

20. 11 spec., paratypes. LACM 72-13.26: 37 m, mud,
0.5 to 1.5 mi W Roca Vagares, Bahía Juanillo,
Guanacaste Prov., Costa Rica (10%57.47'N,
85%45.3"W). Leg. D. Cadien, PI. LaFollette, R/V
Searcher (Searcher 393), 14 February 1972.

21. 27 spec., paratypes. LACM 72-54.45: 37 m, off
Bahía Herradura, Puntarenas Prov., Costa Rica
(9*38.8'N, 84°40.8'W). Leg. LH. McLean, W.
Bussing, R/V Searcher (Searcher 451, 457), 10
March 1972.

22. 15 врес., paratypes. LACM 72-53.27: 21 m, sand,
anchorage in Bahía Herradura, Puntarenas Prov.,
Costa Rica (9°37.97'N, 84°40.5’W). Leg. J.H. Mc-
Lean, R/V Searcher, 9 March 1972.

23. 12 врес., paratypes. LACM 72-57.33: 21 т, sand,
anchorage inside small islet 1.5 Кіп 5 Punta Que-
pos, Puntarenas Prov., Costa Rica (9°22.72'N,
84°09.68'W). Leg. Г.Н. McLean, R/V Searcher, 11
March 1972.

Measurements.—Plotted in Text-figure 14.
p range.—From 18 to 168 m (Keen, 1971, p.

).

Distribution. —Pliocene of San Diego, California
(Hertlein and Grant, 1972, р. 344); Pleistocene of Cal-
Нога (Grant and Gale, 1931, р. 266). Recent from
Catalina Island, California, through the Gulf of Cali-
?rnia, south to Peru and the Galápagos Islands (Keen,
1971, p. 302) or from about 34°N to 12°S (Knudsen,
1982, p. 128). The record from Monterey Bay (lot 1)
extends the distribution to the north to almost 37°N.

Trigonulina bowdenensis (Dall, 1903)
Plate 15, figures 1-4; Plate 16, figures 1-5; Plate 17,
figures 1-4; Plate 18, figures 1-4;
Text-figures 16, 17

Vert; я
еИсогфа (Trigonulina) bowdenensis Dall, 1903, р. 1512; Wood-
"Ing, 1925. р. 92, pl. 11, figs. 6-8.

Я Description.—Shell small (up to 4 mm in length),
Val to rotund. Umbos low, strongly prosogyrate. Lu-
(е more deeply impressed in left valve. Sculpture
Dsisting of seven (о 10 high, narrow, radial ribs an-
“rior to unsculptured posterior slope, which project
а ventral margin. Surface of perfectly preserved
E" covered by minute, pointed pustules forming
5 parallel to ribs. Along postero-dorsal margin
ere are two closely spaced, narrow ribs. Interior sur-
ace Nacreous, its ventral margin fluted. No teeth in
eft Valve, Right valve with a strong, subumbonal, car-
al tooth and a groove along postero-dorsal margin
Тесеіуе postero-dorsal margin of left valve.

30+
[72]
| =]
Ф
Е
о
® 20-
Ф%
5
Ф
ке!
Е
2 104

Т T T T

T 8 9 10
number of ribs

Text-figure 16.—Histogram showing rib number distribution of
Trigonulina bowdenensis (Dall, 1903).

Lectotype (selected herein).—USNM 135689. This
is the specimen figured by Woodring (1925, pl. 11, fig.
6), a left valve.

Dimensions of lectotype.—Length 3.1 mm; height
2.8 mm.

Type locality.—Bowden, Jamaica. Bowden Forma-
tion (early Pliocene).

Paralectotype.—USNM 482415. This is the speci-
men figured by Woodring (1925, pl. 11, fig. 7), a left
valve.

Dimensions of paralectotype USNM 482415.—
Length 2.7 mm; height 2.4 mm.

Paralectotype.—USNM 482416. This is the speci-
men figured by Woodring (1925, pl. 11, fig. 8), a right
valve.

Dimensions of paralectotype USNM 482416.—
Length 3.0 mm; height 2.5 mm.

Remarks.—The type material of this species consists
of the lectotype and the two paralectotypes mentioned
above. All three specimens are glued to a piece of
black paper. As explained under Cardiomya (Bowden-
ia) distira, it has not been possible to remove the spec-
imens from the black paper. The lectotype is therefore
not refigured here.

As hinted at in the above description, there is some
variability as to the number of radial ribs. The large
majority of the specimens at hand has eight to 10 ra-
dial ribs. Only two valves out of the 90 available spec-
imens have only seven radial ribs (Text-fig. 16). One
is from Jamaica, the other from the Dominican Re-
public.

Comparisons.—The Recent Caribbean 7. ornata
d’Orbigny is larger than 7. bowdenensis and has more
radial ribs. The Recent Eastern Pacific 7. pacifica has
about the same number of radial ribs as 7. bowdenen-
sis but reaches even larger dimensions than 7. ornata
(see also “Comparisons” under T. pacifica).

Material. —32 lots with a total of 112 specimens as
listed below:

1. 1 spec., ММВ locality 16802: Río Mao; Cercado
Formation (late Miocene).

2. 2 spec., TU locality 1293 (= NMB locality
18583): Rio Mao, Bluff 1 of Maury; late Miocene.

3. 1 spec., NMB locality 15804: Rio Gurabo; Gurabo
Formation (early Pliocene).

4. 1 spec., NMB locality 15823: Rio Gurabo; Mao
Formation (early Pliocene).

5. 2 spec., NMB locality 15828: Rio Gurabo; Mao
Formation (early to middle Pliocene).

6. 5 spec., NMB locality 15829: Río Gurabo; Mao
Formation (middle Pliocene).

7. 12 spec., NMB locality 15846: Río Gurabo; Gur-
abo Formation (late Miocene).

BULLETIN 351

4 + е
eco”
е”
2 Ke
ә
3- e
Е 299
Е "ag
£ 2222”
2.9 33333
D ө
© е
Ф е
14
T T T
0 1 2 3
height in mm

Text-figure 17.—Length/height diagram of Trigonulina bowde-
nensis (Dall, 1903).

. | spec., NMB locality 15849:

Formation (late Miocene).

. 6spec., NMB locality 15863:

Formation (late Miocene).

. 1 spec., ММВ locality 15864:

Formation (late Miocene).

. 2 spec., NMB locality 15865:

Formation (late Miocene).

. 1 spec., NMB locality 15869:

Formation (late Miocene).

Río Gurabo; Gurabo 22.
Río Gurabo; Gurabo 23.
Río Gurabo; Gurabo 24.
Río Gurabo; Gurabo 25.
Río Gurabo; Gurabo 26.

Río Gurabo; Gurabo 2T.

3 spec., NMB locality 16818:

Formation (early Pliocene).

4 spec., NMB locality 16824:

Formation (early Pliocene).

1 spec., NMB locality 16828:

Formation (late Miocene).

1 зрес., ММВ locality 16832:

Formation (late Miocene).

2 spec., NMB locality 16833:

Formation (late Miocene).

1 spec., ММВ locality 16961:

Formation (early Pliocene).

1 spec., NMB locality 17026:

Formation (late Miocene).
10 spec., TU locality 1354

Río Cana; Gurabo
Río Cana; Gurabo
Río Cana; Gurabo
Río Cana; Gurabo
Río Cana; Gurab?
Río Cana; Сигабо
Río Cana; Cercado

(= NMB locality

18585): Río Cana, Cañada de Zamba; Gurabo For

mation (early Pliocene).
5 spec, TU locality 1250

(= NMB local у

18558): Río Verde; Gurabo Formation (late Mio"

cene or early Pliocene).

. 29 spec., NMB locality 10635: Bowden, Jamaica,

Bowden Formation (early Pliocene).

13. 1 spec., NMB locality 15937:
Formation (early Pliocene).

14. 1 spec., NMB locality 15945: Rio Gurabo; Gurabo 28.
Formation (late Miocene).

15. 1 spec., NMB locality 15962: Rio Gurabo; Gurabo 29.
Formation (early Pliocene).

16. 1 spec., NMB locality 16031: Río Gurabo; Мао
Formation (early Pliocene). 30.

17. 2 spec., NMB locality 16034: Río Gurabo; Mao
Formation (early Pliocene).

18. 1 spec, TU locality 1210 (= NMB locality ^ ^!
18579): Río Gurabo; Gurabo Formation (early
Pliocene).

19. 2 spec, TU locality 1211 (= NMB locality
18580): Río Gurabo; latest Miocene part of Gur-
abo Formation.

20. 2 spec., TU locality 1215 (= NMB locality

18581): Rio Gurabo; Gurabo Formation (late Mio-
cene).

21. 6 spec., NMB locality 16817: Río Cana; Gurabo
Formation (early Pliocene).

32. 3 spec., USNM 135689 (lectotype) and two Рза
lectotypes (USNM 482415, 482416). Bowden, +2
maica; Bowden Formation (early Pliocene).

Measurements.—Plotted in Text-figure 17.

Occurrence.—This species is recorded from the
lowing areas: B

Río Mao: Cercado Formation (late Miocene): NM

fol-

locality 16802, TU locality 1293 (Saunders е! al.
1986, text-fig. 29).

CUSPIDARIID AND VERTICORDIID BIVALVES: JUNG 58

Río Gurabo: late Miocene part of Gurabo Forma-
tion: NMB localities 15846, 15849, 15863, 15864,
15865, 15869, 15945 and TU localities 1211, 1215.
Early Pliocene part of Gurabo Formation: ММВ lo-
Calities 15804, 15937, 15962, and TU locality 1210.

ao Formation (early Pliocene and early to middle
Pliocene): NMB localities 15823, 15828, 15829,
16031, 16034. For location see Saunders ef al., 1986,
lext-figs. 4-6).

Río Cana: Cercado Formation (late Miocene): NMB
locality 17026. Late Miocene part of Gurabo Forma-

tion: NMB localities 16828, 16832, 16833. Early Plio-
cene part of Gurabo Formation: NMB localities 16817,
16818, 16824, 16961, and TU locality 1354 (Saunders
et al., 1986, text-figs. 15, 16).

Río Verde: Gurabo Formation: TU locality 1250
(Saunders et al., 1986, text-fig. 38).

Distribution.—Bowden Formation (early Pliocene)
of Bowden, Jamaica. Cercado Formation (late Mio-
cene), late Miocene and early Pliocene parts of Gurabo
Formation, and Mao Formation (early Pliocene and
early to middle Pliocene), Dominican Republic.

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New England Region, during the past ten years. Tran
actions of the Connecticut Academy of Arts and Science?
vol. 5, art. 6, pp. 447—587, pls. 42-44, 57, 58.

1884. Second Catalogue of Mollusca recently added to the
na of the New England Coast and the adjacent parts Е
the Atlantic, consisting mostly of Deep-Sea species, wit
notes on others previously recorded. Transactions 0 |
Connecticut Academy of Arts and Sciences, vol. 6, ай.”
pp. 139-294, pls. 28-32.

Vokes, H. E.

1939. Molluscan faunas of the Domengine and Arroyo
Formations of the California Eocene. Annals of the ye
York Academy of Sciences, vol. 38, pp. 1-246, pls.
2%

Faw

Hond?

Woodring, W. P.
1925. Miocene mollusks from Bowden, Jamaica. Pelecy
and scaphopods. Carnegie Institution of Washington
lication No. 366, pp. 1-222, pls. 1-28.

род

PLATES

Figure

1-6. Cardiomya (Cardiomya) islahispaniolae (Maury, 1917)
x

EXPLANATION OF PLATE 1

NMB G 14145. NMB locality 15907: Rio Gurabo, Dominican Republic; uppermost part of Cercado Formation (late Miocene).
Exterior of right valve. Length 9.3 mm; height 5.4 mm.

. NMB G 14144. NMB locality 16923: Rio Mao, Arroyo Bajón, Dominican Republic; Cercado Formation (late Miocene).

Right valve. 2. Enlargement of hinge, X30. 3. Interior. Length 8.4 mm; height 5.2 mm.

. NMB G 14146. NMB locality 15903: Rio Gurabo, Dominican Republic; upper part of Cercado Formation (late Miocene).

Left valve. 4. Interior. 5. Enlargement of hinge, X30. Length 8.7 mm; height 5.8 mm.

. NMB G 14147. NMB locality 16929: Rio Mao, Dominican Republic; Cercado Formation (late Miocene). Exterior of left

valve. Length 7.6 mm; height 4.8 mm.

95

ISE NILATIN Y

PLATE |

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 110 PLATE 2

EXPLANATION OF PLATE 2

Figure
ЕТ ОНИЕ (Cadena) islahispaniolac (May, 10017) 22-22. x uM а EU еи
1, 2. Holotype of Сизрдапа ornatior Pilsbry and Johnson, 1917. ANSP 2790. “Santo Domingo”. Left valve. 1. Exterior; 2.
Interior. Length 5.0 mm; height 2.9 mm.
3, 4. Holotype of Cuspidaria gabbi Pilsbry and Johnson, 1917. ANSP 2791. “Santo Domingo”. Left valve. 3. Exterior; 4.
Interior. Length 8.0 mm; height 5.3 mm.
5-8. Pleciodon caber еккен Кати Еа ET Dee ој gehe a, БИ ro иш Bee
5, 6. USNM 63943. Catalina Island, California; 16 fms., mud. Left valve. 5. Exterior; 6. Interior. Length 17.6 mm; height 9.9
mm.
7, 8. USNM 63943. Catalina Island, California; 16 fms., mud. Right valve. 7. Exterior; 8. Interior. Length 17.4 mm; height 9.2
mm.
9, 10. Plectodon NR ER O A И O a s erill
Lectotype. USNM 63193. Off Sombrero Island, Leeward Islands, Lesser Antilles; 72 fms. Left valve. 9. Exterior; 10.
Interior. Length 11.3 тт; height 6.7 mm.
М; 12. Tresore ee 2-2... 1 I AL occu IC a у...
Holotype. LACM 2718. Locality LACM 65-6.22: 0.4—0.7 miles 110 to 132 degrees T From Ship Rock, Santa Catalina
Island, California Channel Islands, California (33°27’N, 118°30’W). Depth: 82 m. Right valve. 11. Exterior; 12. Interior.
Length 4.8 mm; height 4.2 mm.

45

45

50

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CA
SI

EXPLANATION OF PLATE 3

Figure
1=5. Cardiomya (Bowdenia) distira (Dall, 1903)
1-4. NMB G 14139. NMB locality 15846 Río Gurabo, Dominican Republic; latest Miocene part of Gurabo Formation. Incomplete
right valve. 1. Exterior; 2. Enlargement of sculpture, X120; 3. Interior; 4. Enlargement of hinge, X60. Length 2.7 mm; height
2.2 mm.

- NMB С 14140. NMB locality 10635: Bowden, Jamaica; type locality of Bowden Formation (early Pliocene). Interior of right
valve. Length 3.2 mm; height 2.0 mm.

86

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PLATE 3

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 110 PLATE 4

EXPLANATION OF PLATE 4

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Figure
1-5. Cardiomya (Bowdenia) distira (Dall, 1903)
All specimens from NMB locality 10635: Bowden, Jamaica; type locality of Bowden Formation (early Pliocene).
1, 2. NMB G 14141. Right valve. 1. Exterior; 2. Enlargement of rostrum, X90. Length 3.0 mm; height 2.0 mm.
3. NMB G 14142. Interior of left valve. Length 2.9 mm; height 2.3 mm.
4, 5. NMB С 14143. Left valve. 4. Exterior; 5. Enlargement of rostrum, X65. Length 3.3 mm; height 2.3 mm.

09

EXPLANATION OF PLATE 5
Figure Page
1-4. Plectodon granulatus (Dall, 1881)

NMB G 14138. NMB locality 16857: Río Cana, Dominican Republic; Cercado Formation (late Miocene). Right valve. 1. Interior;

2. Enlargement of hinge, Х20; 3. Exterior; valve broken during handling for SEM photography; 4. Enlargement of sculpture, X60.
Length 10.6 mm; height 6.4 mm.

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PLATE 5

PLATE 6

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 110

EXPLANATION OF PLATE 6

Figure
КОА NA аа E Еа ya ur es ee ea 46
USNM 94214. Recent. Station 2648: off Cape Florida, 84 fms., sand.
1, 2. Left valve. 1. Exterior; 2. Interior. Length 8.8 тт; height 5.6 mm.
3, 4. Right valve. 3. Exterior; 4. Interior. Length 11.5 mm; height 6.5 mm.

ONAF ¿SANIVAIG GIIQHOOLL?IHA аму апмуашазпо

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EXPLANATION OF PLATE 7

Figure Page
i4 Pleciodo SERIAS ЭБ ASI аан везан en a а A A 46
USNM 94214. Recent. Station 2648: off Cape Florida, 84 fms., sand.

1. Enlargement of left hinge (X30) of specimen shown on Plate 6, figure 2.
2. Enlargement of right hinge (X20) of specimen shown on Plate 6, figure 4.

3, 4. Enlargement of sculpture of right valve shown on Plate 6, figure 3. 3. Near center of shell disc, X60; 4. Towards rostrum,
х50.

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PLATE 8

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 110

EXPLANATION OF PLATE 8

Figure
1-6. Наших euer УЗИ ERBE A A We PUR eia og Jav мент кү әре 47
USNM 444416. Recent. Eolis Station 370: off Ajax Reef, Florida, 70-90 fms.
1. Interior of right valve. Length 6.0 mm; height 5.6 mm.
. Exterior of right valve. Length 6.1 mm; height 5.7 mm. 3. Enlargement of sculpture, x50.
. Interior of left valve. Length 6.1 mm; height 5.6 mm.
- Exterior of left valve. Length 6.4 mm; height 5.8 mm. 6. Enlargement of sculpture, х 50.

ON > ш
ONAF SAATVA АПАЯООМИЯЯД аму ANIVAISNI

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EXPLANATION OF PLATE 9
Figure Page
16 ТОУ panacea (РЕШЕНИ ае Киа IS IO AR На A ЗС РАДЕ АА 47
1-3. NMB G 14148. NMB locality: 15832: Río Gurabo, Dominican Republic; middle Pliocene part of Mao Formation. Right valve
1. Interior; 2. Exterior; 3. Enlargement of sculpture, X60. Length 4.3 mm; height 4.1 mm.
4-6. NMB G 14149. NMB locality 10635: Bowden, Jamaica; Bowden Formation (early Pliocene). Right valve. 4. Exterior; 5
Interior; 6. Enlargement of hinge, X40. Length 4.3 mm; height 3.9 mm.

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PLATE 9

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 110

PLATE 10

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EXPLANATION OF PLATE 10

Figure

ОСКЕ cc (ЕКШЕ ENT СТ... a атта ла лы уу 47
NMB G 14158. NMB locality 10635: Bowden, Jamaica; Bowden Formation (early Pliocene). Left valve. 1. Exterior; 2. Enlargement
of sculpture, х 100; 3. Interior; 4. Enlargement of hinge, X50. Length 3.2 mm; height 2.6 mm.

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EXPLANATION OF PLATE 11

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USNM 444479. Recent. Eolis Station 178: off Fowey Light, Florida, 68 fms. d

1, 2. Interior of right valve. Length 4.2 mm; height 3.8 mm. 2. Enlargement of hinge, X45. сл

3, 4. Interior of left valve. Length 3.4 тт; height 3.2 mm. 4. Enlargement of hinge, Х55. "и

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 110 PLATE 11

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 110 PLATE 12

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EXPLANATION OF PLATE 12

Figure
ee 49
USNM 444479. Recent. Eolis Station 178: off Fowey Light, Florida, 68 fms.
1, 2. Exterior of right valve. Length 3.7 mm; height 3.3 mm. 2. Enlargement of sculpture, Х50.
3, 4. Exterior of left valve. Length 4.0 mm; height 3.6 mm. 4. Enlargement of sculpture, X50.

DNAS 25ЧАЛУЛІҢ GIKTHOOLLSHHA аму GIPIVGIdSD;)

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89

EXPLANATION OF PLATE 13

1, 2. Interior of right valve. Length 4.8 mm; height 4.7 mm. 2. Enlargement of hinge, Х35.
3, 4. Interior of left valve. Length 4.7 mm; height 4.3 mm. 4. Enlargement of hinge, Х40.

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LM Та HOLTON. ИЕ SEORKSS 2. аа ee ias Lid qul анат 02; 50 Е
USNM 211469. Recent, off La Paz, Baja California, Mexico; 95-10 fms. Оз

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BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 110 PLATE 14

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EXPLANATION OF PLATE 14

Figure

1, 2. Exterior of right valve. Length 5.1 mm; height 4.9 mm. 2. Enlargement of sculpture, X50.
3, 4. Exterior of left valve. Length 4.5 mm; height 4.0 mm. 4. Enlargement of sculpture, х 50.

DNAS :SHA'IVAIQ апанозпачд аму AMAVAIdsao

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EXPLANATION OF PLATE 15

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1.4 Батаны DO РЕНЕ 1903)... 3 aud lu мало 22---222,2424.2-42.2..... a3 Е
NMB locality 15846: Río Gurabo; Gurabo Formation (late Miocene). do
1, 2. ММВ С 14151. Interior of right valve. Length 3.5 mm; height 3.1 mm. 2. Enlargement of hinge, Х40. EA

3, 4. NMB G 14153. Interior of left valve. Length 3.3 mm; height 3.0 mm. 4. Enlargement of hinge, X50.

PLATE 15

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 110 PLATE 16

EXPLANATION OF PLATE 16

Figure
A рел ds РИ О ua зу ыл E UE E UM 53
NMB locality 15846: Río Gurabo; Gurabo Formation (late Miocene).

1-3. NMB G 14150. Exterior of right valve. Length 3.8 mm; height 3.2 mm. 2. Enlargement of sculpture, X75; 3. Further
enlargement of sculpture, х 150.

4, 5. NMB G 14152. Exterior of left valve. Length 4.0 mm; height 3.3 mm. 5. Enlargement of sculpture, х 150.

ONAF :5ЯАЛУЛІҢ MIAJODILAYA аму AIRIVAIdSAD

SI
—

EXPLANATION OF PLATE 17

Figure Page
1-4. Prigonulina bowdenensis Gall, 1903)... A 05522529220 53
NMB locality 10635: Bowden, Jamaica; Bowden Formation (early Pliocene).

1, 2. NMB С 14154. Interior of right valve. Length 2.8 mm; height 2.5 mm. 2. Enlargement of hinge, X60.
3. 4. NMB С 14156. Interior of left valve. Length 2.1 mm; height 1.8 mm. 4. Enlargement of hinge, X90.

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BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 110 PLATE 17

PLATE 18

BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 110

ч

EXPLANATION OF PLATE 18

Figure

1, 2. NMB G 14155. Exterior of right valve. Length 2.3 mm; height 1.9 mm. 2. Enlargement of sculpture, х 100.
3, 4. NMB G 14157. Exterior of left valve. Length 2.8 mm; height 2.6 mm. 4. Enlargement of sculpture, х 100.

ОМОГ :5ЧАЛУЛІҢ GIKTHOOLLWHA аму AMNMAVAIdsao

N
о

74 BULLETIN 351

INDEX

Note: Page numbers are in light face; plate numbers are in bold face; page numbers on which principal discussions occur are in italics.

Abbott, 1974 cin env A anne on wre ruhen 45-48
АЛОЕ АС 1804 A ken euren are tee O a 40
Aguayo, 1943... eese cernere rne nennen натака свита 47
Ajaz Reef ..,‚.......›› A reas 47, 48
ает, NOTET esa cab vice кези кбй уккан КККК Вт ЕЕК DY PEN 40
Anacapa Island. ....s eee eren nean nhe enn rhe aree reri rne na 50
Arroyo Babosico ........................ еее еее еее ж... 36, 43
Агоуо Вајбӣ 4, eere eir eren nh nnn enn arra hne no 39, 41, 42
Arroyo Zaloya ....ocoooocronccccccnacancconon carro nano non 36, 43, 46
A A O a 50
Bahía Baritlwalte |... esee eee rn error rr rmt rre hn reni 50
Bahía de los Angeles ............................ rn ееееееее nnns 50
Bahia de los Muros ..iisrcer rhe rrr enr ren en tho eee 50
ЫНШЫ iS 27020 ү.-...72 50
Bahia СкішісӘ-;22.5220;55 O ката бей 50
Ваша Нела оен ета еерее тоате E
Баша ЈАТО пи | | || 21
Baja California ...................... HH 50
Baja California Sur... 50
pr yn exkestekixsexo vade == en 44, 45
таи [о ПН АНИ nn nenne Er Y РАК FEY 46, 48
bowdenensis, Trigonulina ............ 15-18, 35-37, 39-41, 49-51
Bowden Formation»... ee э. уукка лек 37-43, 46, 49-53
Ра a E = VE UE TAY XR VENAE E ваа 42
БИШІ 27, reso Fee KE н UE РТ RES БЕК ERE AT CER 46, 48
caelata, Verticordia о.о toan nnn 47
СНА uere Колер ке ee жеті Pe Fd кет, 48, 50, 51
California Channel Islands ...................................... 48, 50
Campeche Bank ....... menn 45
Cape Florida ............................ 4 04 еее. 45
cardiiformis, Verticordia ......................0.ы ы. 47
Cardiomya А. Adams, 1864 .......................1ҮЛ.:.2.... 40
craspedonia 2.2.2.2... 19665 еее себе hne nenne ne tonne 39, 41
Ia dd MEER VE кета кат 36, 37, 39, 40
islahispaniolae гй. nnn 35-37, 39-41
Cardiomya (Bowdenia) distira ............. 3, 4, 40-42, 43, 46, 51
Cardiomya (САРА ОУ) есенва uen rrr nre 40
craspedonlg, пени ВЕЌЕ ea vie ко ee eu Ee qua ee en nee enne non 41
ISIARISDaNIOlGE өзекктекекеті rriren rintt R ee rre 1, 2, 38-40
Carpenter, 1864 ......›.... eerte en en eher yn фа hehehe nenne naa 44
Catan ШИШ o а ТТТ” Si
Cedar Коув 2577 ка eerte везна Т 45
Cercado PortitatiotE оеро irre 40-42, 44, 45, 52, 53
СИН ВИЕ o ooo D Re NN 51
craspedonia, Cárdiomya ....... eene 39, 41
стагредота, Cardiomya (Сатйїотуа)............................. 41
Cuspidaria
A 200000 07... 77 40, 41
islalispantolae обе eee енне ео мҺəмМәМцөцəц0Мцөц0Һ0ШДШ 40
OPI И a AA 40, 41
Cuspidaria (Bowdenia) йїзїїўа_..................................... 42
Cuspidaria (Plectodon)
Paaa AN 44
lo A O A AE 44
A Т 222277777277 35, 39-.41
Du Iso еркек кН oe PERDE R E 44

De 1680 от 40, 44, 45, 47
DE ет MN T LEE 47
DS 003 ти 39, 41-45, a
Dall and Simpson, 1901 ..............................4.4.0.2 еее
distira
CN ТАТЫ MT ATE 36, 37, 39, 40
Cardiomya (Bowdenia) unn 3, 4, 40-42, 43, 46, 51
Cuspidaria (Bowdenia) ........ een 4
Durham, 1944. „оно erre errorem mee erri 40
ОЛА е dE РЕКИ ӨТ hrs HEP КЕ СИЛНИ ИТ 44
БЕТТЕН ыы uH dear HM 39, 40
fischeriana
Hals een een ЕСЕН EVE AE СКОК ЕНЕ КЗД АА 46
Verticordla сый, уута беке _______________ | 45
Wile ос правени ну а O xus 37, 44-48
шан 01000000 47, 48
QUE. INA NN. 40
BABB, Cuspidaria «iiie eee eee entere n e ener 40, 41
Galápagos Islands |... eer rennen ete nen tnn nter 44, 51
Gardner 1926 arco dada nn 44
ВИЕ DR nina А
gouldiana, Меавға „лон наа нан ri viii" 40
ПИ кан ТИ оно A бија 40, 48,5!
granulata
Cuspidaria (Plectodon) ..........111. еее еее еккен ене 44
Leiomya (Plectodon) ..... линии осон 4
МАПЕ ТК еее тектік TO CER секске т еке секта CUR cee
ат етте о 2, 5-7, 36, 37, 40, 4
Guanacaste Province .....................554886ө еск ка ККЕ
SAA 22 50, |
GE ФЕ MARIDO, A terim кита e Ке ree Epor v een кк 4
¡AU ee ak Cre e er ro lL 40
Guppy, 1876. sau nennen era eese d Fev ov СЕ 4
Gurabo Formation ve eo oo YE Eur ro rrr rmn 41-43, 52 >
ЛКЕН. 0.2... у ^
fischeriaa nia кина int eoe ho rb o EN YR Y ek da вен ка re nr АЁ 6
ДП ПИЙ ea 9, 10, 36, 37, 39—41, 45, E
Harris 19 an ТЕН CO XT кал Ма e CPV b ba rv vec OC 24
FIO A Анка | а КРЕНА НЕ
Hertlein and Grant, 1972... erre e кити 5
НИШ 1949 rin Cre Y V GE FI enr a 50
Hopkins Marine Station .................... конна бий РР”
Institut Francais du Pétrole |... creen |
ТИЙ СЕДОВ | coi ce rec yr peso V кы ana КЕТТЫ 50
IMD С, ағас етекке 18 tip v pa EC e ER err XEM
islahispaniolae "
Cardlomy eio nt ex tin com EUER OX eer e 35-31, ША”
Cardiomya (Cardiomya) ..... eee enn 1,2, 38 40
CRspldanla „СЛЕВА СЕТУ rola osa ез 50
o A 50
ШЗ lao RA A re 50
Islas Revilla Gigedo |... eren e Ц 50
Tela Todos ВАШЕ: 2.2222... еск exo eri E
50

CUSPIDARIID AND VERTICORDIID BIVALVES: JUNG 75

Jamaica Ты осы 37, 39-43, 46, 47, 49, 51-53
Jamaicensis
la 9, 10, 36, 37, 39-41, 45, 46
OA TAS) a A a 45
IR a MEHR re ИНЕ Оя 35, 36
ШЕШ AR ИР ОООО 40
Honn, „ОЦЕНА УНИ ОС NR 36
BEEN ББ 30 одо л юл. 36
И ле eod 44, 47, 48, 51
EE ons aus st T 50
EE OE IL LLL O CDE 44, 45, 47, 48, 51
La Қана ава ERE e Ed 36, 43
A ОО ОАК х 50
ВЕНИ oo o a A 44
€lomya ТОСОО) ТИИИШ re енеке CO ED n 44
ОШ Ce EE БАТЫ, 44
ШЕ оо око a 43, 46, 52, 53
SISAL Tbh I ee 46, 48
ООО
EE 00.1 45, 50
acu nd НЫНАН 2. 40
ні БЛ RN EE пина 48
БАНА e M c a E 45
өлше ДА ТТК ORE 30, $1
Neaera
ов јада ТАРИ лушы си узнала 40
РН нул АА нс ыы 40
кани НЕОН ППТ M АЦА ЈО Б 44
ИИ КСС т 40
"hi (Chat лк a 45
ЕВЕ Со ОС... 50
A A 44
БЫ. 46,47
p» any, ee 46
ata
Б... 11, 12, 40, 46, 47, 48-52
сине НЫН 4... 47, 48
Verticordia (ТОЛАДЫ ae 47, 48
m erticordia (УИШН) nn molar 48
B ИО ри pM ү 40, 41
И ОА A ез ише 40
а И ПСВ, Trigonulina .......... eene 2, 13, 14, 35, 40, 48-52
EE Az ov о АИ 50
o o у о ш... ER ра 44
"аа Se ei ыо. 51
"M u 40, 41
| КЕШ ШОШ, ТОЙ 2. 40, 41
A A eie rete trente 44
вени вен Ты Т 2, 5-7, 36, 37, 40, 44

“aber 44

Punta Arena de la VENTANA vie rro ns 50
Е ОИЕ A A NE DEI UD 51
доо ис Са пе вади ђе ео ае варана ра СЕ АЦО 51
TOO ЕЛІН east 50
ИИО PERLE er re 50
ZEE IE A RE Т ИО 47
PRO Gonna Sarin Ud OH EUCH ОКИ ланци 38, 41, 42, 44, 52, 53
ARO a e МИШ ach tic awed tices nae vas sus dane EA XN 36, 45
генге ліне ТЕН” 41-43, 46, 52
IO DUO ие она азн на CHA IHE eret ert 36, 43, 46
IIO NIBUS И 0-22 ST POR 38-42, 52
ONG ОВО URL, оо ее OH M E S 36
NIU UNS S Os А МУНА КАНОН SQUE ween EC wand SE LESER TUA 47
RIO даны, БЕТЕР Те ыры CIERRE UR X ERR CIO е 26; 32, 94
RIO BUS US БОЙ. eios спане ает 36, 43
ОСЕ VERTUS. eec eee Wa Gn сњачња ав ее ee 51
НИ Ро cU АНИ ари? Сер ae dns 24
Sante ети ISTUD Oo И баула n Ie жа UE ead 48, 50
sena Mea USING У ео ное 50
Saunders, Jung, and BijusDuval, 1986 „сено cantara rte ee
Re rr rer а ГАЛ 35, 37-39, 42, 43, 45, 46, 52, 53
Saunders, Jung, Geister, and Biju-Duval, 1982 ................... 35
БОДИНА PISCIORDN очи ну ver ex vien Don i ex lut А 44
scabrata, Cuspidaria tPIectodon). nina rr a hes 44
ЗОНА REA ameter es Ener bore rr кон. dn LH A REIR 44
Shoal River Formation ..................................... 37, 44, 45
SOMO ИШ CU oen COUR GEAR CU VAN A RODA а сеа re 44
ОТУ: QUIM cars клеток ODIO AA OCC HO GE EIN 47
ТОО DET as аана еа оен Dt ee se е 45
Swiss National Science: Foundation ¿ciar ek een err 35
A карала евала ува иа hehe ran 47
МЕ ене а ek стена o MAINE
y оо слава A 15-18, 35-37, 39-41, 49-51
BOND ооа ГТ есе тан 11, 12, 40, 46, 47-52
ПРОБИ A A tUe соби Ск 2, 13, 14, 35, 40, 48-52
ДОО LOS. asin уйны кө ea каскада ride 40
A ИРА РТА O 47
VERA DENS E O E A O И 47
MA ТТТ qut T 47
O eher 47
DUPLO O A S А 47
A за E n 45
GIU d ОТОО ORE Lus e eer UNE TA S SERT EXEn (QA Vx pen a 47
ornata ОРОТО Шай... nase. 47
Vertieormia ТИЧ) JOPIGICENSIS a aet ert er tea exa eR 45
Verticordia (ПРОПИСЕ) OMIA „оо erra ee exer eh Rn 47, 48
УНС t VEFRODPEIRS) OPHBIM aera eeiam mE HEU COPIE 48
hi eremia i vn seen 35, 39-41, 45
VOR; EIER seen 40
WOOGIE, 10 serien hen Erie au rena 41, 43, 45-47, 51
Үлбі rennen асаа ені OX DUE CER re ЕНА 45

PREPARATION OF MANUSCRIPTS

Bulletins of American Paleontology usually comprises two or more sep-
arate papers in two volumes each year. The series is a publication outlet for
significant, longer paleontological monographs (i.e., more than approximately 50
printed pages), for which high quality photographic illustrations and the large
quarto format are required.

Submissions are welcome from any author, regardless of institutional or or-
ganizational affiliation. Authors must, however, be members of the Paleontological
Research Institution at time of publication; annual membership is currently
US$25.00. Publication costs of the Bulletins are heavily subsidized by the Insti-
tution, but authors are currently required to pay illustration charges at a rate of
$120.00 per plate and $35.00 per text-figure.

Important references for style and format are 1) Bulletins of American Pa-
leontology “Instructions for Authors’ (volume 108, number 347, pages 149-153);
2) Chicago Manual of Style (fourteenth edition) 1993. Recent issues of the Bul-
letins provide useful guides but note changes with the “Instructions for Authors"
mentioned above.

Manuscripts must be typewritten, and double-spaced throughout (including
direct quotations, tables and references). All manuscripts should contain a table
of contents, lists of text-figures and/or tables, and a short, informative abstract
that includes names of all new taxa. Format should follow that of recent numbers
in the series. All measurements must be given in the metric system, alone or in
addition to their English system equivalents, The maximum dimensions for pho-
tographic plates are 178 mm X 229 mm (7 inches X 9 inches; outlined on this
page). Single-page text-figures should be drafted for reproduction as single col-
umn (82 mm; 3% inches) or full page (178 mm; 7 inches) width, but arrangements
can be made to publish text-figures that must be larger.

Authors must provide three (3) printed copies of the text and accompanying
illustrative material. Authors are strongly encouraged to prepare their manuscripts
on word processors, as this considerably expedites publication. On initial sub-
mission, however, only printed copies of the manuscript should be sent; the disk
copy should be retained so that revisions can be made after the review process.
The text and line-drawings may be reproduced xerographically, but glossy prints
at publication size must be supplied for all half-tone illustrations and photographic
plates. These prints should be identified clearly on the back.

Referenced publication titles must be spelled out in their entirety. Particular
care should be given to reference format. Consult the “Instructions for Authors,”
referred to above, as several changes have been introduced with that issue. Ci-
tations of illustrations within the text bear initial capitals (e. g., Plate, Text-figure),
but citations of illustrations in other works appear in lower-case letters (e.g., plate,
text-figure).

Format of systematic descriptions should follow that in any recent number
of the Bulletins.

Original plate photomounts should have oversize cardboard backing and
strong tracing paper overlays. These photomounts should be retained by the author
until the manuscript has been formally accepted for publication. The approximate
position of each text-figure in the text should be indicated. Explanations of plates
and text-figures should follow the References Cited.

Gilbert Dennison Harris
(1864 - 1952)

Founder of the Bulletins of American Paleontology (1895)

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ISBN 0.877 10-44 |