SAN)
6. (.-44

HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

Revision of the

balanomorph barnacles;

including a catalog

of the species

MU8. COMP. ZOOL...
r-IBRARY

NOV 221977

HARVARD
UNIVKRSiTY

William A. Newman
and Arnold Ross

MEMOIR 9

San Diego Society of Natural History

1976

Revision of the

balanomorph barnacles;

including a catalog

of the species

William A. Newman
and Arnold Ross

Scripps Institution

of Oceanography

and San Diego

Natural History Museum

MEMOIR 9

San Diego Society of Natural History

1976

SAN DIEGO SOCIETY OF NATURAL HISTORY MEMOIRS

MEMOIR 9, pages 1 - 108

Issued March 31, 1976

Frontispiece. Chionelasmus darwini (Pilsbry)*, one of the most generalized or primitive living balanomorphans, is known from
two isolated insular situations where it inhabits relatively deep water (approx. 450m|. The first specimens were taken near the
turn of the century by the U. S. Fisheries Steamer Albatross off Kauai Island, Hawaii, and then a couple of decades later by a
cable ship off Rodriguez Island, southwestern Indian Ocean. While additional specimens have been taken near the original
locaUties, there are no reports of any having been found between these two extremes. Chionelasmus therefore qualifies as a
refugial form, but not a usual one since it is both insular and in relatively deep water, well out of the mainstream of balanomorph
evolution.

♦(RA^ Te Vega Sta. 23-95, Sept. 4, 1971, S. of Molokai I., Hawaii - specimens courtesy of Dr. D. P. Abbott, Hopkins Marine
Station, Stanford University.)

PUBLISHED WITH FINANCIAL AID

FROM THE

W. W. WHITNEY PUBLICATIONS ENDOWMENT

CONTENTS

Introduction 9

How to use this work 9

Acknowledgments 10

Historical 10

Origin of the Balanomorpha 14

Monophyletic 14

Polyphyletic 15

Evolution and diversification 17

Chthamaloidea 17

Balanomorphoidea 20

Balanoidea 22

Morphology 24

Composition and definitions of suprageneric taxa 36

Order Balanomorpha 36

Superfamily Chthamaloidea 36

Family Catophragmidae 36

Family Chthamalidae 36

Superfamily Balanomorphoidea 36

Family Coronulidae 37

Family Bathylasmatidae 37

Family Tetraclitidae 37

Superfamily Balanoidea 38

Family Archaeobalanidae 38

Family Pyrgomatidae 38

Family Balanidae 39

Catalog of species 40

Superfamily Chthamaloidea 40

Family Catophragmidae 40

Family Chthamahdae 40

Subfamily Pachylasminae 40

Subfamily Euraphiinae 40

Subfamily Chthamalinae 41

Superfamily Balanomorphoidea 43

Family Coronuhdae 43

Subfamily Chelonibiinae 43

Subfamily Emersoniinae 44

Subfamily Platylepadinae 44

Subfamily Coronulinae 44

Family Bathylasmatidae 45

Subfamily Bathylasmatinae 45

Subfamily Hexelasminae 46

Family TetracUtidae 46

Subfamily Austrobalaninae 46

Subfamily TetracHtellinae 46

Subfamily TetracUtinae 47

Superfamily Balanoidea 49

Family Archaeobalanidae 49

Subfamily Archaeobalaninae 49

Subfamily Semibalaninae 55

Family PjTgomatidae 56

Subfamily Pyrgomatinae 56

Subfamily Ceratoconchinae 58

Subfamily Bosciinae 59

Family Balanidae 59

Incertae sedis 69

Literature Cited 71

Bibliographic Supplement 100

Index 103

INTRODUCTION

The Cirripedia constitutes a diverse and
abundant subclass of crustaceans, and repre-
sentatives are found in virtually all marine
environments. There are four orders, the Asco-
thoracica, Rhizocephala, Acrothoracica and
Thoracica. The Thoracica contains the true
barnacles and these are distributed between
three living suborders; the stalked barnacles or
Lepadomorpha, the asymmetrical sessile bar-
nacles or Verrucomorpha, and the sessile acorn
barnacles or Balanomorpha. These appear in the
Silurian, the middle Cretaceous, and the late
Cretaceous, respectively. The Balanomorpha en-
compasses the greatest diversity of free-living
and symbiotic forms, and as Darwin (1851a:5)
noted, the present epoch may go down in the
fossil record as the "Age of Barnacles" (Fig. 17).

The basic classification of the Thoracica
was formulated by Darwin (1854b), and his
system was expanded and somewhat revised by
Pilsbry (1907a;1916). Pilsbry's classification
formed the basis for that in the Treatise on
Invertebrate Paleontology (Newman et al, 1969).
Although the Treatise provides diagnoses of taxa
to the generic level, it does not enumerate
the species contained in each genus, nor does it
provide a guide to the hterature concerning
them. The present study fills these needs for the
Balanomorpha. It also constitutes the first major
revision of higher taxa in more than half a century.

The Balanomorpha may not be an entirely
natural assemblage, but rather a grouping of
phylogenetically parallel lineages not readily
derivable from one another nor from a common
balanomorph ancestor. The possibility of at least
a diphyletic origin was suggested by Withers
(1924:2). Thus, in preparing this revision we
were alert to the possibility that the Balano-
morpha might be separable into two or three
suborders. However, in the final analysis it be-
came clear that such a proposal was indefens-
ible or premature. Therefore, the Balanomorpha
in the broad sense has been retained. Yet three
major lineages can be recognized, and we con-
sider them to constitute superfamilies: Chthama-
loidea, Balanomorphoidea and Balanoidea. In
addition, one new family and numerous sub-
families are also proposed here, and many of
the 65 genera contained in the Balanomorpha
are redistributed within this modified systematic
framework (Fig. 1).

Much of what has been done here might be
interpreted by the casual observer as simply
"splitting" and "rank-raising." Indeed, Hyman

(1959:697) voiced concern over systematic prac-
tices in recent years: "Any acute observer can-
not fail to notice the disease prevalent in
zoological systematics today of raising rank of
groups and of assigning high ranks to groups
that differ only in minor characters." Neverthe-
less, in the present study new lines of evidence
indicate previously unrecognized affinities, and it
seems to us that the classification must be altered
and expanded to accommodate them.

Initially, classification of thoracicans de-
pended on surficial morphology of the shell,
and it is only in recent years that thin sec-
tions have revealed remarkable internal struc-
tures that have drastically altered our under-
standing of the affinities among the
Balanomorpha. Likewise, comparative studies of
trophi and chaetotaxis, or of such structures as
the base of the intromittant organ, have greatly
improved and broadened our understanding of
interrelationships between higher and lower
taxa. In addition, numerous collections by both
individuals and expeditions, from the deep sea,
from coastal waters, and especially from tropical
seas where the greatest diversity is found, have
provided new materials that have compelled us
to alter our concepts and rearrange existing
groupings in order to continue to develop a
natural system. If our system is accepted, the
practical inconvenience and annoyance will really
be quite temporary.

HOW TO USE THIS WORK

The specialist will probably have httle dif-
ficulty in using this work, but some explana-
tions seem appropriate. It is divided into three
parts: evolution, systematics and catalog of
species. We have attempted to arrange the
genera and higher taxa phylogenetically. How-
ever, for simplicity, ease, and (or) lack of
knowledge, species are listed alphabetically under
their respective genera or species groups in the
catalog. The index is the entree to species.
The first page number given after each species
leads to that species in the catalog. For genera
and higher categories, and for some species, the
index leads into the systematic and evolutionary
sections as well. Species names in the index are
given without generic indication unless they
have been used in more than one genus. In
such cases the generic names used in this work
are given.

10

Diagnoses of suprageneric taxa, and for a
single new genus (Notobalanus), are provided
in the systematic section. Diagnoses for estab-
lished genera can be found in the Treatise on
Invertebrate Paleontology (1969); original sources
for subsequently described genera are cited
herein.

The general arrangement of the catalog fol-
lows that of the preceding evolutionary and
systematic sections. The original author, date
and page are cited for each species and, where
appropriate, a citation of the most compre-
hensive synonomy, which may not necessarily
be the most recent. This is followed by a
relatively complete list of references through
1973, but including many through 1975; many
of the non-systematic papers are briefly anno-
tated. Finally, general distributional and some-
times bathymetric and stratigraphic data are
included, but needless to say, distribution of the
majority of the species is very poorly known.
Following the body of the catalog there is a list
of species incertae sedis.

ACKNOWLEDGMENTS

The primary data base for this work was,
quite naturally, the hterature, and we have
cited virtually all that was available to us.
Much of the contemporary literature was made
available as reprints by authors and others,
and we thank them for their generosity. A
large portion, however, came from various
university and museum libraries, over many
years, through direct borrowing and interhbrary
loans. Librarians involved are too numerous to
mention individually, but we thank them, known
and unknown to us, for their services.

As with many data bases, sources extend
well beyond pubhshed works, and we are much
indebted to numerous cirripedologists for vol-
uminous oral and written communications. There
have been so many we hesitate to mention
them by name, for fear of not including all.
But we must acknowledge Huzio Utinomi of the
Seto Marine Biological Laboratory; Alan J.
Southward of The Laboratory, Plymouth; EUza-
beth C. Pope of the Australian Museum; and
Victor A. ZuUo, University of North Carolina
at Wilmington.

Data were also extracted from the vast
collections of the Scripps Institution of Oceanog-
raphy and the San Diego Natural History
Museum, and from materials made available on
loan by curators of collections in other insti-
tutions. In particular, we would Uke to thank
Thomas E. Bowman of the National Museum

of Natural History; Torben Wolff of the Zoologi-
cal Museum, Copenhagen; Jan Stock of the
Zoological Museum, Amsterdam; L. B. Holthuis
of the Rijkmuseum, Leiden; J. P. Harding of
the British Museum (Natural History); WiUiam
K. Emerson of the American Museum of Natural
History; and J. Wyatt Durham of the Paleon-
tology Department, University of California,
Berkeley. We have also garnered knowledge and
experience from innumerable specimens sent to
our laboratories for identification by ecologists
from all over the world.

Development of the catalog has passed
through the hands of several assistants. It be-
gan many years ago as a compilation of
references to species of immediate interest in
contemporary literature, and was subsequently
expanded to include all primary hterature on
all species by Mrs. Carol Platt-Kourtz, who
carried it forward for five years as a sideline
to her regular work. Mrs. Cecelia Ross spent
nearly a year of intensive work on it, and
finally Ms. Gayle Kidder aided substantially in
bringing it to its present state. We thank
these young ladies for their concerted efforts
and ask their forgiveness for the moments
when attention to detail became excessively
tedious.

This revision is for the most part a by-
product of our work on the systematics of the
Cirripedia. Support, in part, was provided by
several grants from the National Science Foun-
dation (to W.A.N. : GB-4973X through BMS575-
17149), and these are gratefully acknowledged.

HISTORICAL

Classification of the thoracican Cirripedia,
beginning in good part with the work of Leach
(1817, 1818, 1825) and Gray (1825), was placed
on a firm foundation by Darwin (1851-1854).
Darwin's three basic divisions, the Lepadidae,
Verrucidae and Balanidae, are the principal
ones recognized today (Pilsbry, 1907a, 1916;
Kriiger, 1940; Withers, 1953; Newman et al,
1969). Progress in the classification of the
Thoracica, from Leach (1817-1825) to that being
proposed, is given in Figure 1. Gruvel's (1903b)
classification is omitted. Suprageneric taxa are
indicated only under the Balanomorpha.

The Lepadomorpha (= Lepadidae sensu
Darwin) contains the most primitive Thoracica,
members of which are inferred to have arisen
from a free-hving stem near the Ascothoracica
(see Newman et al, 1969; Newman, 1974:437).
While the unity of the Lepadomorpha has never
been questioned, the relationships of the scalpeUi-

11

Leach 1825

Ord Campylosomata^
Ord Acamptosomala

Fam Clisiadae

Fam. Balamdae ■^^-- —
Fam, Coronuladae

Gray 1825

-Fam Analifidae

"^Fam PolUcipedidae

- Fam Balamdae

- Fam Pyrgomatidae

- Fam Coronubdae

Darwin 1854

Ord Thoracica y

Fam Lepadidae''^^
Fam Verrucidae ^
Fam Balamdae -^

Subfam Chlhamalm.

Subfam Balanmaf

Present

Subor Lepadomorpha
Subor Brachy lepadomorpha
rSubor Verrucomorpha

///

Pilsbry 1907-16 /X/sybor Balannmorpha

Subor Lepadomorpha //
Subor Verrucomorpha y'v"^
Subor Batanomorpha
Fam Chthamahdae
Balamdae
bfam Chelonibiinae
Subfam Coronu
Subfam Balanmae

/Fam I
Fam

Fam Chthamahdae
Subfam Catophragminae
Subfam. Pachylasmmae
Subfam Chthamahnae

Fam Bathylasmatid

Fam Tetrachtidae

Fam Balamdae
Subfam Chelonibi
Subfam CoronuUn
Subfam Fmersoniinae-
Subfam Balaninai
Subfam- f'yrgomatinae

Proposed for Balanomorpha

Subor Balanomorpha Pilsbry 1916
Superfam Chthamaloidea Darwin 1854
Fam CaLopfiragmidae Utinomi 1968
Fam Chthamahdae s s

Subfam Pachylasmtnae Utinomi 1968

Subfam Euraptumae nov.

Subfam. Chthamahnae s s

Superfam Balanomorphoidea nov

Fam Coronubdae Leach 1817

Subfam Chelombunae Pilsbry 1916

Subfam Emersonimae Ross 1967

Subfam Platylepadinae nov.

Subfam Coronuhnae s s
Fam Bathylasmatidae Newman & Ross 1971

Subfam Bathylasmatmae s.s

Subfam Hexelasminae nov.
F"am Tetrachtidae Gruvel 1903

Subfam Austrobalamriae nov

Subfam TetracUtethnae nov

Subfam Tetracbtmae s s.

Superfam Balanoidea Leach 1817

Fam Archaeobalanidae nov

.Subfam Archaeobalaninae s s

Subfam Semibalaninae nov
Fam Pyrgomatidae Gray 1825

Subfam Pyrgomatinae s.s.

Subfam Ceratoconchinae nov.

Subfam Bosciinae nov.
Fam. Balamdae s.s.

Figure 1. History of the classification leading to that proposed for the Balanomorpha.

form and lepadiform groups remain obscure.
However, problems that arise in this regard
have no direct bearing when considering the
origins of the Balanomorpha, because it is
generally agreed that one looks to the scalpelli-
form or pollicipoid barnacles for the antecedents
of the sessile barnacles (Darwin, 1854b; Withers,
1953; Broch, 1924; Newman et al, 1969; Newman
and Ross, 1971).

Darwin's (1854b) classification of the
Thoracica reflects the view that the sessile
barnacles (Verrucidae and Balanidae) evolved
from the Lepadomorpha as independent lineages.
The Verrucomorpha was recognized by Darwin
(1854b:495) as sharing a number of character-
istics with the Lepadomorpha and the Chtha-
mahdae among the Balanomorpha. However, the
sum of the characters he enumerated favor a
lepadomorph rather than a balanomorph an-
cestry for them. Withers (1914:945) considered
Prouerruca from the upper Cretaceous to "con-
stitute, in fact, the 'missing link' between the
pedunculate Cirripedes of the family Polhcipedi-
dae [= Scalpellidae] and the sessile asymmetri-
cal Cirripedes of the family Verrucidae." The
two lateral plates of one side seen in Prouerruca,
and Eouerruca but missing in Verruca, are
homologous with those of the presumed an-
cestor of the Balanomorpha as well as the
Verrucomorpha. Thus, what these early verrucids
indicate is that the lepadomorph ancestors of
both suborders were comparable (Fig. 2).

The next sessile suborder, the extinct
Brachylepadomorpha, was unknown to Darwin.
It was instituted by Withers (1923:37) to ac-

commodate Brachylepas, which Woodward (1901:
150) previously considered a pedunculate bar-
nacle. Withers (1953) subsequently discovered
that Pycnolepas Withers (1914) was not only
a sessile barnacle, but also that it was inter-
mediate in structure between stalked barnacles
and Brachylepas (Fig. 2). He stated that,
while the Brachylepadomorpha "includes the
commonest and most widespread of the Cre-
taceous symmetrical sessile cirripedes. . . ."
they "do not appear to be in the direct line
of descent of the Balanomorpha, as already
pointed out by Pilsbry. They apparently repre-
sent an independently developed sessile type,
which, except for the reduced number of capitu-
lar valves, probably resembled the ancestor of
the Recent primitive Balanomorpha (Catophrag-
mus)" (Withers, 1953:344; see Fig. 2).

Gruvel's (1903b) classification of the Balano-
morpha departed radically from Darwin's scheme,
but it was rejected by Pilsbry (1907a, 1916)
and subsequent workers as in good part un-
natural. Pilsbry (1907a, 1916) elevated Darwin's
famihes to suborders, primarily to allow for an
expanded classification at subfamihal levels.
Darwin's Balanidae thus became the Balano-
morpha, containing two families, the Chthamah-
dae and Balanidae. He further divided the
Balanidae into the Balaninae, Chelonibiinae, and
Coronuhnae, all primarily on the basis of shell
characters.

Numerous subgenera, in good part based
on characters Darwin (1854b) used in formu-
lating sections, have been proposed, particularly
by Pilsbry (1916) and Hoek (1907), especially

12

Ir i Ic

PoUicipoid Lepadomorpha

Figure 2. Monophyletic origin of the Balanomorpha and inferred relationships: The principal divisions (superfamilies) of the
Balanomorpha are directly related to and stem from a pedunculate stock allied but distinct from that which gave rise to the
Verrucomorpha and Brachylepadomorpha. Radiations and relationships of the superfamilies are illustrated in figures 4, 5 and 6.
(see text for discussion.)

13

in the Balaninae. Yet, in the years since Pilsbry
(1916), few alterations have been made in the
basic classification of the Balanomorpha. Nilsson-
Cantell (1921) resurrected GruveFs (1903b) Tetra-
clitinae (in part) as a subfamily of the Balani-
dae, and fostered the Stellatus- and HembeU-
groups of Chthamalus, suggested by Pilsbry
(1916). Ross (1968) subsequently elevated the
Tetraclitinae to familial level; Utinomi (1968)
divided the Chthamalidae into three subfamilies;
Ross (in Ross and Newman, 1967) created the
subfamily Emersoniinae for an extinct form allied
to the turtle barnacles; Newman and Ross (1971)
proposed the family Bathylasmatidae for a group
of relatively primitive deep water balanoids;
and Ross and Newman (1973) resurrected Gray's
(1825) Pyrgomatidae (in part), a name available
for a group of coral barnacles designated
Creusiinae by Baluk and Radwanski (1967b:
468). Despite these advances the broad aspects
of the classification have remained the same. As
it stands, it fails to portray many actual or
inferential relationships and this has necessitated
the present revision.

Although appUcation of the biological species
concept spread in systematic studies of other
groups, the Darwinian tradition of numerous
varieties (subspecies) in the cirripeds has con-
tinued to prevail, especially in the Stellatus-
group of Chthamalus, Tetraclita s.s., the Balanus
amphitrite group, the subgenus Megabalanus,
and the coral barnacles.

Students of the balanomorphs will find some
unfamiliar features in what we propose and
these may be quite disconcerting without back-
ground information. Darwin's work on the Cirri-
pedia had a profound two-fold effect. On one
hand, he established the basic classification and
brought order to a chaotic and wide spread
literature. On the other hand, he arranged the
higher taxa in such a manner as to bias
virtually all subsequent phylogenetic studies.
While Hoek (1913) and Pilsbry (1916) expanded

upon the basic framework, they retained the
Darwinian order in their monographs in which
the morphologically primitive forms, the chtha-
malids and coronuhnids, appeared at the end
and the more highly evolved forms such as
Megabalanus, appeared at the beginning. The
first break in tradition came with Gruvel
(1905a), and in a more acceptable manner, with
the work of Nilsson-Cantell (1921), Kriiger (1940),
and Withers (1953), where the various groups
were, with the exception of the turtle and whale
barnacles, placed in a more or less acceptable
phylogenetic sequence.

The present study is a further attempt to
order the balanomorphs as naturally as possible,
down to and including the subgeneric level. In
doing so, some marked departures from pre-
vious classifications have been made. This may
not prove upsetting to new students of the
Balanomorpha, but the "old Hne" may find it
difficult to accept the turtle and whale barnacles
as groups having relatively primitive origins,
and to find the tetraclitids closer to them than
to the balanids.

It may also prove disconcerting to find
that most free-living acorn barnacles cannot be
readily assigned to either Chthamalus, Balanus
or Tetraclita. But it must hkewise have been
upsetting to earUer students of the group when
certain workers decided that most barnacles
were not Lepas as Linneaus had established.
The changes proposed herein reflect a sharpen-
ing of resolving power over the past decade or
so, a sharpening made possible through the
efforts of many students of this remarkable
and fascinating group of animals.

Beginning on page 25 we illustrate various
features and relationships of the shells and ap-
pendages of the balanomorphs. These were orig-
inally prepared to aid us in our understanding
of the diversity of morphologies involved, and it
is hoped that they will be useful to the reader.

14

ORIGIN OF THE BALANOMORPHA

MONOPHYLETIC

Until recently the Balanomorpha consisted
of the Balanidae and Chthamalidae. Darwin
(1854a:152, 176) and subsequent authors, con-
sidered the Chthamahdae the more primitive
and directly derivable from scalpelliform bar-
nacles. The criteria for this judgment cover both
homologies of hard parts and morphology of
appendages, especially in the most primitive or
generalized chthamalid, Catophragmus (Catomer-
us). The fossil record supports this interpreta-
tion, because Catophragmus appears in the late
Cretaceous. Representatives of the Balanidae do
not appear until the early Eocene.

In the chthamalid Pachylasma, while the
body and appendages are wholly chthamaloid,
the shell wall and to some extend the opercu-
lum are in certain respects balanoid in appear-
ance. Darwin (1854b:475) stated that when he
first examined the shell of Pachylasma he "did
not doubt that it was . . . Balanus." But when
he examined the animal's body, he found the
characteristics preeminently chthamaloid, and
concluded that (1854b:477) "Pachylasma would
be the point of contact [of the Chthamalidae]
with the Balaninae, . . . [for] when the shell
alone ... is examined, it is hardly possible to
separate this genus [Pachylasma] from Balanus. "
Unfortunately, the fossil record does not lend
support to this view because Pachylasma first
appears in the Miocene, well after the appear-
ance of Balanus. Nonetheless, the implication
exists; the Balanidae may have come from the
Chthamahdae via Pachylasma.

Subsequent work on the origin of the Balani-
dae seemed to make a chthamahd ancestry
more plausible. Hoek (1883, 1913) described a
number of deep-water species that appeared by
shell characters to belong to Balanus, but the
nature of the soft parts, particularly the structure
of the labrum and the third cirrus, was atypical,
and while he considered them balanids, he pro-
posed the genus Hexelasma for them. Pilsbry
(1916) reviewed the status of this genus and
concluded that the species in Hexelasma be-
longed instead to the Chthamalidae, close to
Pachylasma, and this assignment was followed
by Kruger (1940), Withers (1953), Zullo (1963a),
and Utinomi (1968). Zullo (1963c:190) oversimpli-
fied this picture with his sweeping statement
that "the Balanidae . . . differ materially from
the Pachylasma group [including Hexelasma]
only in the structure of the labrum . . . and

that they were derived from the Pachylasma
group stock." Despite this oversimphfication it
would seem at this point that there would be
relatively little difficulty in deriving balanids
from chthamahds, for the shell of Pachylasma
and soft parts of Hexelasma would appear,
superficially, to bridge the gap.

Taking this simplistic view at face value, a
model for a monophyletic diversification of the
Balanomorpha would be as follows (Fig. 2). The
Chthamalidae, containing the most primitive
members of the suborder, gave rise to the re-
mainder of the Balanomorpha. Fundamentally,
chthamahd hard parts consist of deeply articu-
lated opercular valves, a wall of eight solid
plates (three pairs of laterals overlapping the
unpaired carina and rostrum), and several whorls
of small imbricate plates, comparable to the
peduncular plates of certain scalpeUids, surround
the region where the wall contacts the sub-
stratum. The basis is membranous. A large
bullate, lepadomorphan-Uke labrum surrounding
the mouth parts has mandibular palps situated
on its lateral margins. The scalpellid-Uke man-
dible, composed of several incisor-hke teeth and a
spinous rather than molariform inferior angle,
is simple. The first and second cirri are modified
to assist in the transfer of food captured by
the posterior four pairs to the mouth parts; that
is they have been modified to serve as maxiUi-
peds. The cirri, armed with simple setae and lack-
ing speciaUzed spines are hke those of the Le-
padomorpha. The penis, originating between the
last pair of cirri below the anus and flanked by
a pair of multiarticulate caudal rami or appen-
dages (the furca), lacks a basidorsal point. All
these features are found in the most generaUzed
members of the extant Chthamaloidea, Catophrag-
mus sensu lato, fossil forms of which are the old-
est balanomorphs known (late Cretaceous).

Diversification of the chthamahds included
the appearance of a number of Uneages in all
of which the whorls of imbricate plates were
lost, the number of wall plates was reduced from
eight to six, and in some cases four, and the
caudal appendages inevitably disappeared
(Fig. 4). The reduction of wall plates from eight
to six was accomphshed in two different ways —
most commonly the carinolaterals drop out,
thereby retaining the arrangement where the
rostrum as well as the carina remain over-
lapped; and less commonly, the rostrolaterals
fuse with the rostrum forming a compound plate
that overlaps the adjacent laterals. The latter

15

arrangement is the same as that seen in higher
non-chthamalid Balanomorpha and is presumed
to herald them. The chthamalid bullate labrum,
inherited from the Lepadomorpha, gave way to
the thick but non-bullate condition, with concomi-
tant changes in the nature of the mandibles
to the more advanced balanid type. The third
cirri became intermediate in structure between
the second and fourth rather than more closely
resembling the fourth, and the opercular valves
became complexly but not deeply articulated;
all features seen in Hexelasma and related genera
(Bathylasmatidae).

Further advances included a flattened labrum
that became cleft, aiding in the removal of food
from the cirri. Concomitant with this, the third
pair of cirri completed the transformation to
maxiUipeds. Apparently at this point the sohd-
waUed Balanidae and Tetraclitidae appeared and
diverged from the ancestral Bathylasmatidae
(Figs. 2, 4 and 5). Both went on to develop
distinctly different complex wall tj^es, variously
armed cirri, and in the balanids, a penis with
a basidorsal point.

POLYPHYLETIC

As palatable as the monophyletic scheme
may be, Zullo (1963c:190) noted that there were
conflicting views regarding affinities within the
Balanomorpha and that it is possible that the
balanomorphs are polyphyletic. Withers (1924:2)
stated that he was "not at all convinced that
the Chthamalidae and Balanidae . . . are so
nearly related as is supposed," but he did not
pursue the subject in subsequent writings
(through 1953). Recently, Utinomi (1968:33),
expressed a similar view, suggesting that the two
families were independently derived from lepado-
morph ancestors, but unfortunately he did not
elaborate further on the matter. We became
involved in the problem of the unity of the
Balanomorpha when working on a revision of
Hexelasma (Newman and Ross, 1971). In this re-
gard, Bage (1938:10) had already pointed out
that, "from the examination of the soft parts
of the animal it is apparent that the reference
of the genus [Hexelasma] to the Balanidae or

Sessile Cirripedia
Pedunculate Cirripedia

Figure 3. Polyphyletic origin of sessile cirripeds: It is weU documented that extinct Brachylepadomorpha and the Verruco-
morpha (Jurassic to middle Miocene, and middle Cretaceous to Holocene, respectively) evolved from pedunculate ancestors be-
fore the appearance of the Balanomorpha (Cretaceous to Holocene). The Balanomorpha also descended from pedunculates rather
than from earlier sessile groups (see fig. 2 for fundamental structural differences). Thus, sessiUty evolved three times. However,
there have been suggestions in the literature that two or more of the principal divisions of the Balanomorpha also may have had
separate pedunculate ancestors, as illustrated here. If this were so, sessihty within the Thoracic would have evolved four or five
times (see text for discussion).

16

Chthamalidae, discussed by Hoek (1913) and
Pilsbry (1916), remains unsettled." In our study
(Newman and Ross, 1971:143) it became clear
that Hexelasma and its allies stand distinctly
apart from the chthamaloids and balanoids and
that it was impossible to supply facts supporting
the long standing view that it is from the
chthamaloids, through Pachylasma and Hexe-
lasma, that the balanoid barnacles have descend-
ed. Therefore, we instituted the Bathylasmatidae,
to accommodate several genera, including
Hexelasma, because the group could not be
properly assigned to either the Chthamalidae or
the Balanidae, and suggested that antecedents of
the bathylasmatids may well be found among the
scalpellid Lepadomorpha rather than the
Balanomorpha.

However, in the aforegoing, the balanoid
rather than chthamaloid affinities of the Bathy-
lasmatidae were emphasized, and while the Tetra-
clitidae was recognized as a distinct family,
it was placed rather closer to the Balanidae
than the Chthamalidae, with the "elminoids"
standing in a somewhat intermediate and ques-
tionable position (Newman and Ross, 1971:143).
Ross (1970:9) provided the solution to the last
problem by demonstrating that at least two
species referred to Elminius by previous authors
were in fact tetraclitids rather than balanids.
Hence the difficulties posed by the elminoids

evaporated and the four famiUes of the Balano-
morpha became more sharply defined.

The gulf between chthamaloids and balanoids
is particularly great. The tetrachtids and bathy-
lasmatids are envisaged here as more closely
related to one another than to either the
chthamaloids or the balanoids. Although the
tetraclitids and bathylasmatids are separable,
there is presently no reason to believe that the
former have not evolved from the latter, and it
is to this complex rather than the balanoids
that we infer the coronulines and chelonibiines
are most closely related. Thus, it remains pos-
sible that the Balanomorpha is triphyletic; an
artificial assemblage of three independently
evolved sessile types (Fig. 3). If this were the
case, sessility was achieved five times in the
Thoracica: once in the Verrucomorpha, once in
the Brachylepadomorpha, and three times in the
Balanomorpha — all from comparable, but lepado-
morph ancestors. Nonetheless, for the aforemen-
tioned reasons, the Balanomorpha is retained
here, even though it may not be a natural
grouping. As a partial solution to this problem
we propose that the families recognized here be
distributed between three superfamiUes, the
Chthamaloidea, the Balanomorphoidea, and the
Balanoidea. Should polyphyly be documented in
the future, one or more of these would of neces-
sity become suborders.

17

EVOLUTION AND DIVERSIFICATION

CHTHAMALOIDEA

Darwin (1854b:486) commented that
"... Catophragmus forms, in a very remark-
able manner, the transitional link [between the
Chthamalidae and the Lepadidae], for it is
impossible not to be struck with the resemblance
of its shell with the capitulum of Pollicipes.
In Pollicipes, at least in certain species, the
scuta and terga are articulated together — the
carina, rostrum, and three pairs of latera, mak-
ing altogether eight inner valves, are consider-
ably larger than those in the outer whorls —
the arrangement of the latter, their manner of
growth and union, — all are as in Catophrag-
mus. If we, in imagination, unite some of the
characters found in the different species of
Pollicipes, and then make the peduncle so short
(and it sometimes is very short in P. mitella)
that the valves of the capitulum should touch
the surface of attachment, it would be impos-
sible to point out a single external character
by which the two genera . . . could be dis-
tinguished." As Withers (1928b) noted, Pilsbry
(1916) suggested an even nearer hkeness with
the more speciaUzed Sciliaelepas, and this model
was adopted by Newman et al (1969:R269, fig. 90).

Darwin further noted that the trophi of
chthamaloids are similar to those of lepado-
morphs. The labrum is thick and bullate, and
this is basically a lepadomorphan character.
The tridentoid mandibles and the multiarticulate
caudal appendages of the primitive chthamaloids
are typical of the pollicipoid lepadomorphans,
and it is the sum of these arthropodal struc-
tures that separate the chthamaloids from other
Balanomorpha. The first and second cirri are
modified for cleaning the posterior net-forming
pairs of particulate matter and transferring it
to the mouth. In lepadomorphs, generally but
one pair of cirri is so modified, but pollicipoids
have modifications of the second and even the
third pairs (Darwin, 1851b:313). Finally, the base
of the penis is simple, without basidorsal point,
as in all thoracicans except the Balanoidea. In
general, the trophi and chaetotaxis are most con-
servative throughout the lower chthamaloids and
readily distinguish the entire stock from the re-
mainder of the Balanomorpha. The facies simi-
larity with pollicipoids is indeed most striking.

Virtually nothing more could be asked for in
a generalized ancestor for the higher Chthamaloi-
dea than Catophragmus and Catomerus — all
the essential parts are there. All that needs

to be done is to modify the form, by loss or
fusion of shell parts and loss or specialization
of appendages and trophic structures, in or-
der to produce the diversity of taxa presently
observed within the superfamily (Fig. 4).

Relationships between genera have been noted
by various authors. Pilsbry (1916:291) took a
somewhat Gruvellian approach, and arrayed
them in phylogenetic order, primarily according
to number of wall plates. He also divided the
most species-rich genus, Chthamalus, into two
groups based on the nature of the mandible.
These were refined and named informally by
Nilsson-Cantell (1921) as the quadridentoid
Stellatus-group and the tridentoid Hembeh-group.

Zullo (1963c) observed that the more gen-
eralized tridentoid mandible of the Hembeli-
group was the type common to more primitive
chthamaloids and, coupled with differences in
mode of shell reduction, proposed that the
Chthamalidae be divided into three groups:
the quadridentoid Chthamalus, Chamaesipho,
and Octomeris, and tridentoid Catophragmus,
Chionelasmus, and Euraphia, and the tridentoid
Pachylasma. The Pachylasma-group included
Hexelasma (Zullo, 1963a). This is essentially
Pilsbry's (1916:291) classification. Pope (1965),
in a most scholarly review, pointed out some
problems with the tri- and quadridentoid aspects
of the division and this will be returned to
shortly.

Although Utinomi (1968:36) avoided dealing
with the problems that arise when using the
mandibles as a key taxonomic character, he
formally designated subfamilial divisions for
what were essentially the Pilsbry-ZuUo group-
ings: the Catophragminae (Catophragmus, Cato-
merus, and Chionelasmus), the Chthamahnae
(Chthamalus, Chamaesipho, and Octomeris), and
the Pachylasminae (Pachylasma, Hexelasma,
and Tessarelasma). To the Catophragminae one
must add the late Cretaceous Pachydiadema
Withers (1935); to the Chthamahnae, the Recent
Tetrachthamalus Newman (1967) and Jehlius
Ross (1971); and from the Pachylasminae, or
rather from the Chthamaloidea in general, re-
move Hexelasma and Tessarelasma (see Newman
and Ross, 1971:142). We are otherwise in ac-
cord with Utinomi 's groupings, but not as
coordinate taxa (Fig. 4).

The Catophragmidae comprises an ancient
and generalized stock; there is a significant
gap between it and the remaining subfamihes.
The differences, aside from the supplementary

18

Catophragmidae
CHTHAMALOIDEA

Figure 4. Radiation of the Chthamaloidea: It seems unlikely that Chionelasmus and Pachytasma (the only deep-sea members of
the superfamily) evolved from intertidal catophragmids {Catophragmus and Catomeris) whose trophi and anterior cirri are much
more specialized. By default then, the extinct Pachydiadema becomes a more hkely candidate. It also seems unlikely, for the
same reasons, that Octomeris gave rise to Pachytasma, or vice versa since the opercular valves of Pachylasma are already
advanced. Finally, while it seems unlikely that Chionelasmus gave rise to six-plated euraphiines, the possibility cannot presently
be ruled out.

If higher balanomorphans arose from chthamaloids, workers since and including Darwin (1854b) consider that it would have
been via a pachylasmine ancestor (see text for further discussion).

19

whorls of plates in the Catophragmidae, are
the extremely primitive nature of the primary
wall plates and the opercular valves. Even in
Chionelasmus. where the supplementary plates
have been reduced to but a single whorl,
and the primary wall plates from eight to six,
the primitive pollicipoid facies is retained.

The jump from Catophragmidae to Chtha-
malidae is wholly in "modernization" of the
wall, the arthropodal structures remain essen-
tially the same in the primitive Euraphiinae
and Pachylasminae, as do the number of pri-
mary wall plates. Whether catophragmines gave
rise to these two subfamilies independently,
or the apparently more generalized shallow water
euraphian Octomeris gave rise to the deep-water
Pachylasminae, cannot be resolved at this time.

Although the arthropodal structures seem to
be similar, it is evident by shell characteristics
that the euraphiines and pachylasmines are not
closely related. Zullo (1963c:190) emphasized
that further advances in shell arrangement differ
in the two groups. Consequently, it is clear that
the pachylasmines attained a six-plated condi-
tion by development of a compound rostrum,
and the euraphiines by loss of the carino-
laterals.

The transition from Euraphiinae to Chthama-
linae is clearly by way of Euraphia, and con-
sists primarily of the first and only significant
change in the trophic apparatus. This change,
the development of the so-called quadridentoid
mandible of Pilsbry and others, is probably an
adaptation, along with the specialized setae
(grapples or cards) on the anterior cirri, to life
in the high intertidal, as suggested by Pope
(1965:965).

The important feature of the quadridentoid
mandible is probably not so much that there
are four teeth, or that the second and third
teeth are commonly bifid, but that the inferior
portion rather than forming an angle support-
ing a group or tuft of spines, is drawn out
into a relatively long straight comb. Neverthe-
less, Pope (1965:27) questioned the taxonomic
value of this character. For example, she stated
that the finding of "large individuals of certain
AustraUan species (Chthamalus antennatus: 49)
in which normally 4-toothed species have devel-
oped only 3 teeth, or conversely, of 3-toothed
species [Euraphia withersi: 43) with 4 teeth,
is going to make the drawing of distinctions
between Zullo's generic groups somewhat
difficult."

While there are difficulties in placing a few
species in one or the other of these two groups,
they are minor, and Pope herself explains most
of them away. There is some variation in the

number of teeth in E. withersi, and Pope
(1965:43) pointed out that the majority of
specimens will be found to have three teeth,
and under any circumstance, the inferior angle is
pectinated, not combed. Thus there would appear
to be no real difficulty here. And with regard
to C. attennatus, Pope (1965:49) stated, "Some-
times mandibles of the right and left sides
may vary and while the left one may have a
s^e//af US-pattern for its lower tip, the right
may have a "hembeli" one. However, in indi-
viduals with somewhat hembeli-\\\ie jaws, the
small, fourth double tooth can be seen, thus
enabhng the real affinities of C. antennatus
with Chthamalus to be recognized."

Pope (1965:58) goes on and provides further
evidence that alleviates her own objection to
the recognition of Euraphia as separate from
Chthamalus. In C. malayensis "juveniles, or
during regeneration in certain individuals, the
lower tip of the mandible is reminiscent of the
Hembeli pattern." She then (1965:59) notes that
mandibles regenerating after having been dam-
aged take on a euraphian form, and further-
more, that it seems as though juveniles and
regenerating C. antennatus have to pass through
a euraphian stage during the development of
their much toothed and highly complex man-
dibles. The juvenile situation is clearly onto-
genetic; it is indeed an ancestral euraphian
reminiscence, as suggested by Pope, and that
a regenerating Umb would have to repeat the
process is not surprising. Therefore, the dis-
tinction between mandibles in the two groups
recognized by Pilsbry and used informally by
Nilsson-Cantell seem not only to be useful
taxonomically, but they aid in elucidating the
evolution of higher chthamaHds. A combed
stellatoid mandible is seen elsewhere only in
some Tetraclitidae, which also develop specialized
cirral setae and occur high in the intertidal
(Ross, 1970).

Despite the great age of the Chthamaloidea,
the group has been relatively conservative,
undergoing little diversification with regard to
both structure and habitat. None (with the pos-
sible exception of certain Pachylasma on crinoids)
has formed an obligate symbiotic association.
The catophragmoid facies, first appearing in rocks
of late Cretaceous age, was apparently an adapta-
tion to high energy conditions along the shore
and must have been abundant and widely distrib-
uted in the past. Extant species have restricted
distributions in the austral region and the tropi-
cal Americas.

The most advanced catophragmid, Chionelas-
mus, and the relatively generalized chthamalid
Pachylasma are presently the only deep-water

20

members of the entire superfamily — all others
are intertidal. Where Euraphia and Chthamalus
occur together the former and more generalized
occupies the higher reaches of the intertidal,
the highest of all balanomorphs (Pope, 1965;
Southward, 1964b). Littoral and shallow water
habitats that would otherwise appear suitable
for chthamaloids are occupied, presumably
through competitive exclusion and other bio-
logical interactions, by higher balanomorphoids
and by balanoids.

BALANOMORPHOIDEA

The Balanomorphoidea, proposed here, en-
compasses the Coronulidae, Bathylasmatidae and
Tetraclitidae (Figs. 1 and 5). Taken together
one finds a suite of fundamentally primitive
or generalized characters, including 8 wall plates,
membranous basis, generalized opercular plates,
no basidorsal point on the penis, and a labrum
and cirrus III of intermediate form.

Until recently the Tetraclitidae occupied an
uncomfortable position as a subfamily of the
Balanidae (Ross, 1968, 1970), whereas certain
species of the Bathylasmatidae had been placed
at one time in the Balanidae and at another
in the Chthamalidae before being recognized as
constituting a distinct family (Newman and Ross,
1971). The suite of characters that unites the
Tetraclitidae and Bathylasmatidae under the
Balanomorphoidea is the same as that which
prevents their being satisfactorily assigned
to either the Chthamalidae or Balanidae. The
same holds true for the coronulid Chelonibia.
But in addition it has 8 wall plates, a con-
dition that previously complicated understanding
the evolution of Balanidae. Our proposed re-
assignment of the coronulids to this super-
family not only removes this difficulty, but
also allows for further insights into the funda-
mental organization and evolution of the
balanomorphoids.

The intermediate position of Hexelasma s.l.
and related genera between Chthamalidae and
Balanidae, appeared ideal in arguments for
derivation of the latter (ZuUo, 1963c: 190). How-
ever, it was shown (Bage, 1938; Newman and
Ross, 1971:148) that the nature of the soft
parts are not altogether intermediate, but rather
possess many unique characteristics. Also, argu-
ments requiring bathylasmatids as intermediate
between chthamaloids and balanoids neglected
the apparent eight rather than six-plated origin
of the latter. Such arguments side-stepped what
was considered a living representative of an
early balanid, Chelonibia. At the same time.

the bathylasmatids could not be considered
directly derivable from chthamalids, and be-
cause of Chelonibia they did not appear to be
appropriate ancestors for the balanids. The
obvious conclusion was that they must have had
a separate origin, and probably then from a
comparable pollicipoid lepadomorphan stock
(Newman and Ross, 1971).

The preparation of this revision afforded
us the opportunity to take a fresh look at the
matter. We found that the apparent, obstacle
raised by Chelonibia was actually not a problem
at all. As stated previously, and as will be
given diagnostic documentation in the system-
atic account to follow, Chelonibia and its aUies
have hitherto been incorrectly placed among the
Balanidae, and this has stifled our thinking
on the matter. Once freed of this constraint
the whole picture becomes simplified and emi-
nently clearer. Chelonibia and other coronuhds
appropriately fall in the Balanomorphoidea.

Because of their extreme specialization as
obligatory commensals of marine reptiles and
mammals (Ross and Newman, 1967), what is
known of the Coronuhdae, beyond Chelonibia,
tells us nothing about the evolution of the
higher Balanomorphoidea. It is the Bathylas-
matidae that provides us with the data base
from which further inferences can be drawn.

The Bathylasmatidae form a natural group
and we propose that it be divided into the
subfamilies Bathylasmatinae and Hexelasminae
(Fig. 5). Opercular valves are generalized in
the former, and form a vertically oriented cone.
In the latter, the opercular valves are more
balanoid, and the plane of the scuta lies almost
horizontal, across the orifice of the shell. Within
the family, Hexelasma stands in an intermediate
position between Bathylasma and Aaptolasma.
However, it is more closely related to the latter
and together they form the Hexelasminae.

Many features in Aaptolasma herald the
Tetraclitidae. The comparable form of the man-
dible and labrum, the tendency for the third
cirri to be antenniform, comparable opercular
valves, and the peculiarity of the wall plates in
being permeated by longitudinal chitin-filled
tubes, are all characteristics that draw them
together.

In the original diagnosis of Aaptolasma,
only a small number of differences could be
assembled to distinguish the genus from Tetra-
clita s. 1., but at that time no six-plated
tetraclitids were known. However, it subse-
quently became clear that Balanus (Austro-
balanus) imperator Darwin was not just closer to
Tetraclita than to Balanus, as Darwin (1854b:290)
had recognized, but that it was a tetraclitid

21

BALANOMORPHOIDEA

Figure 5. Radiation of the Balanomorphoidea: The CoronuUdae are specialized obligate commensals of large crustaceans, some
fish, sea turtles and snakes, and marine mammals. The Tetraclitidae are, on the other hand, specialized for an intertidal
existence. These two families apparently did not give rise to higher forms. If the Balanoidea arose from the Balanomorphoidea,
as proposed here, it probably would have been via the hexelasmines, species of which are presently confined to the shelf (see
text for further discussion).

22

(Ross, 1971:266). Thus a distinction based on
the number of wall plates, between Hexelas-
minae and Tetraclitidae, fails. What remains
is that the tetraclitids have radii (at least
fundamentally), cirri II and III commonly are
armed with bipectinate and other complex setae,
and the labrum is wholly non-bullate; all ad-
vances above the more generalized bathylasma-
tid plan.

In Aaptolasma, the solid wall is permeated
by strips of chitin in much the same manner as
in certain tetraclitids (e.g. Epopella). All other
tetraclitids have tubiferous walls whose charac-
teristics provide the distinguishing features of
the subfamilies.

There is a marked correlation between ad-
vances in specialization of appendages, shell
wall, and bathymetry. The most generalized
forms in the Bathylasmatidae occur at the great-
est depths; in fact Tetrachaelasma, a close
relative of Bathylasma, is the deepest known
balanomorphan (2,300 m). Members of the
Hexelasminae occur on the shelf between ap-
proximately 100 and 1,000 m. All members
of the Tetrachtidae on the other hand, hke
most Chthamaloidea, are intertidal or restricted
to very shallow water. The hiatus between the
low intertidal and 100 m or so is exploited
by the Balanoidea. The Balanomorphoidea (ex-
cept Tesseropora sp. on Heliopora, and the
coronulids), hke the Chthamaloidea, do not form
obligate commensal relationships as do many
members or groups of the Balanoidea.

BALANOIDEA

It has been tacitly assumed that the Bal-
anidae s. I. had an eight-plated ancestry, as
did the chthamalids (cf. Newman et al, 1969).
Darwin (1854b) pointed out the tripartite ros-
trum of the chthamahd Pachylasma, and of the
presumed balanid Chelonibia. Runnstrom (1925)
reported that the rostrum in Balanus balanoides
formed ontogenetically by fusion of the rostro-
laterals, and this has been interpreted as a re-
duction in the tripartite origin of the balanid
rostrum. However, subsequent workers have
failed to confirm this finding in this or any
other balanid, much less a balanomorph.

Direct evidence of a tripartite rostrum is
found in Pachylasma and in Chelonibia, but
as already discussed, these genera fall near
the stem of the Chthamaloidea and Balano-
morphoidea, respectively, and are not directly
involved in the origin of the Balanoidea. It
follows then, that there is no evidence for a
tripartite rostrum (eight-platedness) in the stem

of the Balanoidea. Nonetheless, it is appropriate
that we review arguments to the contrary.

Zullo (1963c: 190), following Darwin and Pils-
bry suggested that the balanoids stemmed from
the "Pachylasma-group." While not specifying
which genera the group contained, he included
Bathylasma ( = Hexelasma, in part), and pos-
sibly Bathybalanus, as did Pilsbry (1916:291,
328). At the time, inclusion of these two genera
previously unknown to Darwin, made acceptance
of the group as the stem from which the
balanoids could have arisen more palatable,
for not only did they have the proper type
of rostrum but also the appendages were con-
sidered to range from somewhat chthamaloid
in Bathylasma to somewhat more balanoid in
Bathybalanus. The labrum in Bathylasma, while
not bullate as in chthamaloids, is relatively
thick and lacks a deep median incision. Also, the
third cirri are somewhat intermediate between
the second and fourth pairs. The situation in
Bathybalanus was thought to be comparable,
although more balanoid. However, we have
shown that Bathybalanus is in all respects
a true balanid and that Bathylasma, while
not a balanoid, is not a chthamaloid either
(Newman and Ross, 1971:142). Furthermore, the
Pachylasma-Bathylasma-Bathybalanus-Balanus
s.l. transition from the chthamaloids to the
balanoids by-passed Chelonibia, previously con-
sidered the only eight-plated balanoid. The
problem of Chelonibia was removed in the pre-
ceding section of this paper, where it was
shown that Chelonibia and its alhes were
primitive balanomorphoids rather than balanoids.
We are left then with the prospect that the
principal balanoid groups descended from balano-
morphoidans rather than chthamaloids.

Early balanoids had a solid wall, as borne
out by both fossil and ontogenetic evidence.
The evolution of higher balanoids has in good
part centered around the development of a com-
plex wall, an evolutionary advance not achieved
to any comparable degree in the chthamaloids
(Darwin, 1854b), but paralleled in many re-
spects in the higher balanomorphoids. At this
point it is not difficult to envisage the Bal-
anoidea as having descended from hexelasmine-
Uke balanomorphoid ancestors, since the trends
are already beginning there: comparably con-
structed wall of six plates, labrum thin and
broadly notched, third cirri somewhat modified
as maxillipeds, and balanoid opercular parts.

We include three families in the Balanoidea.
The solid-walled forms, those included in
Semibalanus, and those having irregular wall
tubes of the non-balaninae type, such as found
in Archaeobalanus, differ so markedly in wall

23

Archaeobalanidae
BALANOIDEA

Figure 6. Radiation of the Balanoidea: A few of the more generalized balanoids, such as Bathybaianus. are found in deep water.
But most free-living forms occur in relatively shallow water, and in the intertidal (Semibalaninae and Balanidae) where upper
limits tend to set the lower limits of the chthamahd zone. Archaeobalanines, on the other hand, are usually subtidal and many
have formed obligate commensual relationships (i.e. Conopea on gorgonians, Acasta on sponges, Hexacreusia, etc, on scleractiniansl.
It is from the archaeobalanines that the Pyrgomatidae, occurring on scleractinians (one exception, on sponges), are inferred to
have been derived, Hkely polyphyleticaUy (see text for discussion).

24

structure from the Balanidae s.s., that we
relegated them to two separate families —
the Archaeobalanidae (including the Archaeo-
balaninae and Semibalaninae) and the Pyrgo-
matidae (including the Pyrgomatinae s.s.,
Bosciinae and Ceratoconchinae). Thus arranged,
the archaeobalanids, which first appear in the
Eocene, are envisaged as having stemmed from
six-plated hexelasmine bathylasmatids or bal-
anomorphoids.

The Archaeobalanidae fall into two subfam-
ilies, the more diverse Archaeobalaninae and the
strictly intertidal Semibalaninae. The Archaeo-
balanus-like forms with tubiferous walls have
undergone the most marked diversification of
any of the balanomorphs. They are well repre-
sented in the intertidal even though the higher
reaches have been left to the tetraclitid balano-
morphoids and most chthamaloids. The semi-
balanines which may have an Actinobalanus
ancestry, apparently did not give rise to any
higher taxa.

Two families stem from the Archaeobala-
ninae. They are fundamentally the solid-walled
Pyrgomatidae and the tubiferous-walled Balani-
dae. The Pyrgomatidae encompasses the coral
barnacles and, while the monophyletic origin of
the group is in question (Withers, 1929a; Ross
and Newman, 1973; Newman and Ladd, 1974),
the consensus is that some, particularly those
contained in the principal subfamily, the Pyrgo-
matinae, and possibly the Bosciinae, have
descended from Armatobalanus (Zullo, 1969b,
1967:127; Ross and Newman, 1973). The Cerato-
conchinae apparently had a different and ap-
parently non-armatobalanid origin (Newman and
Ladd, 1974).

The Balanidae, as envisaged here, may have
stemmed from an irregularly tubiferous-walled
ancestor having a calcareous basis such as
Archaeobalanus. In the Balanidae the principal
evolutionary advance was the establishment of a
regular tubiferous wall in conjunction with an in-
tricate dovetailing between the marginal portion
of the internal ribs of the wall and the margins

of the basis, thereby enabling an individual to
continue to grow diametrically while maintaining
a strong mechanical interlock with the sub-
stratum (Newman et al, 1967). Perfection of this
system, including delayed apphcation of an
inner lamina to the internal ribs of the wall,
produced the unique tubiferous structure dis-
tinguishing balanids from all other Balano-
morpha. Tubiferous walls occur in other Bal-
anomorpha (Pyrgomatidae, Semibalaninae, and
Archaeobalaninae among the Balanoidea, and
the Coronulidae and Tetraclitidae among the
Balanomorphoidea (Darwin, 1854b; Newman et
al, 1967; Ross and Newman, 1967; Ross and
Newman, 1973), but differences in ontogeny
and the nature of the resultant structures
indicate separate origins.

The Balanidae is the most diverse family,
and Pilsbry (1916:78 et seq.) began to group
species of Balanus s.s., informally, into eight
"series." However, he did not follow through
with the matter in his monograph, and today
only the "Series of B. amphitrite" is commonly
referred to in the literature. We have at-
tempted to follow Pilsbry s lead, and have ar-
ranged the species of Balanus in six more or
less natural groups. While some of these are
readily recognizable by a number of characters,
others have been assigned on the basis of un-
defined facies similarities. Thus, while some of
the groups may eventually become genera or
subgenera, such a proposal at the present would
be premature. Those species that we could not
readily assign to one group or another are
placed incertae sedis, at the end of the catalogue.

A considerable part of the diversification
of the Balanoidea has come about through
establishment of obhgate symbiotic relationships
verging on, but in only one case becoming
wholly parasitic (Hoekia on the coral Hydno-
phora, Ross and Newman, 1969). If the present
epoch goes down in the fossil record as
the "Age of Barnacles," as suggested by Darwin
(1851a:5), it will in good part be due to the
remains of symbiotic as well as free-living forms.

MORPHOLOGY

The figures appearing on the following ten pages illustrate various features of the shell and
appendages of balanomorphans. The figures are arranged sequentially, beginning with the shell and
ending with the appendages. In those figures where comparisons are made, the arrangement is
essentially phylogenetic.

1

25

Thoracic
limbs

Tergum

Lateral

Carinolateral

Anterior cirri

Posterior cirri

Carina

Penis

Anus

Tergal depressor muscles

Branchia (right)

Mantle cavity-

Oral cone

Scutum

Lateral

Scutoral adductor muscle

Rostral sinus

Rostrum

Prosoma

Aperture to maxillary gland

Female genital aperture

Oviduct

Rostral depressor muscles

Ovaries / Lateral depressor muscles \ Ovaries

Mantle cavity lining Membranous basis

A.

B.

C.

Posterior net-forming cirri
Anterior cirri (maxillipeds;

Tergum

Scutum

Tergoscatal flaps

Opercular aperture

Orifice of shell wall

Carina Carinolateral Lateral

Figure 7. Model of Semibalanus balanoides (L.): A and B, viewed from left side; C, viewed from rostral end (frontal aspect).
In A, the left carinolateral and lateral wall plates as well as the left tergum and scutum have been removed revealing the in-
terior of the mantle cavity containing the body of the animal as it resides when withdrawn. B, as A, but with the missing
parts replaced and the cirri extended. The posterior three pairs of cirri (in Balanoidea and some Balanomorphoidea) form the
cirral net while the anterior three pairs act primarily as maxillipeds. In C, it can be observed how the cirral net is formed
and how the anterior cirri are positioned to aid in transferring food from it to the oral cone. Photographs courtesy of the
American Museum of Natural History.

26

max'

Figure 8. Principal anatomical relationships:

A. A balanomorph montage*, viewed from the right side, with right cirri extended from aperture formed between the
occludent margins of the opercular valves, primarily the scuta. The six cirri are always biramous. The posterior three form
the right half of the plankton-capturing cirral net, while the anterior three are reduced and otherwise modified, primarily
as "maxiUipeds," for removal of food from the cirral net (cf. Figs. 14-161. Cirri are extended by circulatory hydrostatic pres-
sure and withdrawn by retractor muscles.

B. The tentorial operculum, composed of paried terga and scuta, attaches along its basal margin to the lower margin of
the sheath and is operated by three principal pairs of longitudinal depressor muscles, a transverse adductor between the
scuta, and circulatory hydrostatic pressure.

C. and D. Exploded operculum illustrating scuta and terga respectively, (am, insertion of adductor muscle; rd, insertion of
rostral depressor; Id, of lateral depressor; td, of tergal lateral depressor).

E. Body torn free from its attachment in the operculum and surrounding mantle (carapace), exposing the adductor muscle
(a), the oral cone or labrum surrounding mouthparts (6), the pedicles and proximal portions of the right cirri, the penis (c)
originating between the sixth cirri and resting for the most part between the pedicles of the adjacent pairs, and the right
caudal appendage or ramus of the caudal furca (d).

F. Oral cone enlarged, illustrating arrangement of trophi, from left to right: Labrum (/) with mandibular palps (p) attached
to each side, followed by mandibles (m) and first and second maxillae (max', max").

G. Basal region of a balanid penis illustrating gross form of the pedicel (a) and basidorsal point or horn (6).

*A fully bullate labrum and caudal appendages are characteristics of lower chthamalids. while much reduced third cirri and a penis with a
basidorsal point are characteristics of balanitis (see subsequent illustrations). A pair of outgrowths of the interior mantle lining extends into
the mantle cavity in which the body of the barnacle resides. These, termed branchiae, are variable in structure between taxa, but their
taxonomic value is yet to be determined. They have been little used in systematic studies and consideration of them has not been included here.

27

Figure 9. The balanomorph wall; modifications of basic plan and nature of the basis:

A. Wall of six solid plates; basis on left (not indicated) membranous, on right calcareous. In the latter, the basal margin
of the wall may form some minor denticles, the older portions of which may appear as riblets on the interior of the wall, but
the denticles form neither complex interdigitation with the basis nor anything other than very simple interlaminate figures
in cross-sections of the wall (cf. Fig. 12A, B).

B. As above, but with a "false basis" which may not, as in Euraphia intertexta, include the central portion of the
membranous basis.

C. Longitudinal sections of wall. Left, false basis, formed by successive layers of secondary calcification, in which fusion
to the wall precludes further growth. Right, true basis indicating suture where marginal growth increments can occur. Var-
iously thickened apical portions of wall (sheath) support the opercular valves. The sheath is ordinarily solid and its basal
margin may become dependent (right). When a dependent sheath contacts ribs on the interior of the wall a type of tubiferous
wall if formed, but the sheath does not constitute a true inner lamina (some coronulids and pyrgomatids).

D. Tubiferous wall, in this case accompanied by a tubiferous basis. Well developed, uniformly deployed basal denticles form
complex interdigitation with the basis which in turn is firmly cemented to the substratum. When, with growth, the inner por-
tions of the denticles become secondarily fused, forming an inner lamina, the type of tubiferous wall seen here appears
(Balanidae). If the denticles are simple, the interlaminate figures will be simple; if they have subsidiary lateral cusps, the
interlaminate figures will be complex (cf. Fig. 12H, I).

E. Longitudinal sections of wall. Left, a form with solid wall and basis where growth has been precluded by secondary
calcification (false basis, as in some species of Euraphia). Right, a situation where transverse septa have developed in the
tubiferous wall and basis, and where the cavity formed by the dependent sheath has become secondarily filled and/or cancellated.
Aforegoing structural developments can occur in various combinations.

28

•^^^

Figure 10. The balanomorph wall; modifications of the basic plan. Fundamentally balanomorphs grow diametrically, increasing
in height and basal width by depositing new shell around the basal parietal margins, and the size of the aperture is increased
by additions to the lateral parietal margins to varying degrees (Darwin. 1854b). When lateral increments are negligible or ab-
sent, the aperture is often enlarged by corrosion, as in Tetraclita. Other variations in the fundamental plan are illustrated
here.
A-C. Alterations in growth in gregarious species due to crowding.

A. Pattern in forms such as Semibatanus balanoides and Balanus glandula (without or with carcareous bases respectively)
where normally conical forms become columnar through elongation of the parietes.

B. Alternative response to crowding, as seen in species of Megabalanus and members of the concavus group of Balanus.
where elongation is primarily accompHshed by the formation of a cup-shaped basis. The basis may be permeated by one or
numerous rows of tubes.

C. Situation in some species producing a cup-shaped basis, such as some species of the concavus group and in Balanus
laevis (illustrated), where the extensive cavity formed by elongation of the basis becomes secondarily transversely septate
(cancellate or cystose).

D. Growth in symbiotic forms, such as Acasta imbedded in sponges and Conopea (illustrated) occurring on gorgonians,
in which a cup-shaped basis is formed. In the former it apparently assists in maintaining the apertural end of the barnacle at
the surface of the growing sponge and (or) in enlarging the body chamber without forcing the wall above the surface of the
sponge. In the latter, it elevates the barnacle well above the general surface of the gorgonian, the "keel" of its boat-shaped
basis attaching firmly to the gorgonian axial skeleton.

E-G. Coral barnacles keep pace with the surface of the coral, generally by elongation of the basis.

E. As seen in species of Armatobalanus and Boscia, where elongation of the basis is not extensive and growth of the wall
elevates the barnacle above the surface of the coral.

F and G, where elongation is extensive and the wall plates grow so as to remain more or less flush with the coral surface,
as in Eoceratoconcha and most members of Ceratoconchinae and Pyrgomatinae. In Eoceratoconcha and an early species of
Ceratoconcha, the chamber formed by the basis is canceUate (F). while in most members of the Pyrgomatidae it is open (G).

29

radius

Figure 11. The balanomorph wall; parts and relationships of plates (schematic):

A. Exploded eight-plated wall viewed from the right side. The plates are named, from left to right, the rostrum (R),
paired rostrolaterals, laterals and carinolaterals (RL, L and CL respectively! and the carina (C). The central triangular portion
of each plate is termed the paries (pi. parietes). The basal margin (blackened portion) contacts the substratum. Contacting
surfaces between adjacent plates are stippled, and the portion seen from the exterior is termed the ala (pi. alae). The parietal
margin overlapping an ala may develop a lateral portion which fills the space between it and the adjacent paries. This
structure is termed a radius (pi. radii). Parietes can be sohd or permeated by longitudinal tubes; radii (except in Megabalanus
and some Tetraclitidae) and alae are always solid. It can be observed that the rostrum and carina have alae, the rostro-
laterals have radii on both margins while the laterals and carinolaterals have alae on their rostral and radii on their carinal
margins.

B. An articulated eight-plated wall (as in lower Catophragmidae and Chthamalidae).

C. Eight-plated wall in which the rostrolaterals have become inseparably but discernibly fused to the rostrum forming a
"compound rostrum" (as in some species of Pachylasma and in Chelonibia). A true rostrum has alae and is overlapped by
adjacent plates, while a compound rostrum has radii and overlaps adjacent plates.

D. Compound rostrum where fusion demarcations are no longer discernible. May consist of fused rostrolaterals and
rostrum, or fused rostrolaterals alone (see text for discussion of divergent views).

30

I \ ^-'^ t, yl^

Conopea galeata

B

Solidobalanus hesperius laevidomus

Semibalanus cariosus

Balanus glandula

Megabalanus californicus

Megabalanus californicus
Figure 12. Wall structure in some Balanoidea. See caption on page 35.

31

D Octomeris sulcata

A Catomerus polymerus B Chionelasmus danvini C Pachylasma scutistriatum

E Euraphia hembeli F Euraphia aestuarii

G Chthamalus fragilis H Bathylasma corolliforme I Aaptolasma americanum

J Austrobalanus imperator K Tetraclitella divisa L Tetraclita rufotincta

M Armatobalanus nefrens

N Balanus niveus

O Megabalanus psittacus P Nobia grandis

Figure 13. Opercular plates of the Balanomorpha. See caption on page 35.

32

A Chionelasmus darwini

'"'X^.^J^

B Euraphia intertexta

C Tetrachthamalus oblitteratus

D Chionelasmus darwini

E Euraphia intertexta

F Tetrachthamalus oblitteratus

Figure 14. Trophi and cirri of the Chthamaloidea. See caption on page 35.

33

A Chelonibia patula

B Bathylasma corolliforme

C Aaptolasma leptoderma

D Epopella breviscutum

E Chelonibia patula

\y ^11 iiix-'v VI

F Bathylasma corolliforme /

G Aaptolasma leptoderma

H Epopella breviscutum

III

I Tesseropora pacifica

Figure 15. Trophi and cirri of Balanomorphoidea. See caption on page 35.

34

A Solidobalanus pentacrini

B Balanus amphitrite

C Balanus laevis

D Hoekia monticulariae

G Megabalanus
psittacus

Armatobalanus terebratus / J Armatobalanus allium

Figure 16. Trophi and cirri of Balanoidea. See caption on page 35.

K Balanus
nubilus

35

Figure 12. Wall structure in some Balanoidea. AH, transverse thin sections: I, photograph of basal margin. Increments
resulting in vertical growth of the shell are due to deposition of new sheU along the basal margin of the plates. In primitive
forms lacking a calcareous basis (most Chthamaloidea and the lower Balanomorphoidea and Balanoidea) transverse sections
reveal little complexity in gross structure. With the advent of a calcareous basis there is an opportunity for a complex suture
to form, interlocking the wall to the substratum through the basis. In solid wall forms the interlocking is generally accomplished
by a regular array of simple denticles, the development of which is that of planar mineraUzed entities along centers of calcifi-
cation, and these are visible as simple interlaminate figures in the older portion of the shell (A and B). If subsidiary
denticles are produced perpendicular to the main denticle (as in I), the interlaminate figures wiU be arborescent (in solid walled
forms as in C: in tubiferous walled forms as in E, G and HI. Some species without calcareous bases have tubiferous walls, and
transverse sections commonly appear as in D. Some basically tubiferous walled species with calcareous bases have given up
denticle formation and have all but completely filled irregularly formed tubes (F).

Figure 13. Opercular plates of the Balanomorpha: A-G, Chthamaloidea; H-L, Balanomorphoidea; M-P, Balanoidea (all right
terga and scuta, viewed from within). In the chthamaloids, terga tend to be triangular in outline (without spur, except in
higher forms hke Chthamalus fragilis |G| where a rudimentary spur is developed), scuta never develop a strong adductor
ridge, and the terga and scuta of each side tend to be deeply articulated especially in shallow-water forms (A and D-G).

The situation in the balanomorphoids is somewhat intermediate between that seen in the chthamaloids and lower balanoids;
a tergal spur is variously developed, an adductor ridge becomes prominent in higher forms (J-L) and the plates of shallow water
forms are less deeply articulated. (In the first two superfamilies the tergum is never beaked and a spur furrow, where developed,
is always open.)

Lower balanoids tend to resemble higher balanomorphoids except that the adductor ridge is not particularly strong and
the insertions of the scutal depressor muscles are simple. Closiu-e of the spur furrow (a result of the shaft of the spur becoming
virtually internal), occasionally accompanied by production of a beak (O). apparently develops independently in various lines
(within Semibalaninae, members of the Balanus concavus group, and Megabatanus). Partial or complete fusion of terga and
scuta of each side occurs in all three superfamilies, but marked alterations in general form occur primarily in the coral sym-
bionts (Pyrgomatinae, P). The whale-turtle symbionts (coronuUds) have reduced the opercular plates, and in Xenobalanus
they have been lost completely.

Figure 14. Trophi and cirri of the Chthamaloidea: A and D, Chionelasmus darwini; B and E, Euraphia intertexta: C and F,
Tetrachthamalus oblitteratus. Trophi of lower chthamaloids are similar to those polUcipoid scalpellids — labra are buUate with
crests variously concave but without a median notch (A and B); the mandibular teeth may have spinous superior margins (B);
and the cutting edges of the first maxillae are usually stepwise or notched. In Chthamalinae (C) the mandibular teeth are
never spinose, the second and third teeth are frequently bifid, a fourth bifid tooth is developed and the inferior portion is
drawn out into a straight comb with the inferior angle supporting but a few sptnules, and the cutting edge of the first maxillae
tends to be straight and slightly notched.

The third cirri in general tend to resemble the fourth more than the second (E and F) but in Chionelasmus (D) even the
second are more similar to the posterior ones. SpeciaHzed setae, ranging from bipectinate (E) to pinnate (F) are generally
found on the second cirri. Posterior rami of the third cirri may be antenniform (F). apparently seasonally in intertidal forms.
The caudal appendages or furca, a pair of uniramous appendages attached near the bases of the sixth cirri, and commonly
found in lepadomorphans, are known in a few chthamaloids (D).

Figure 15. Trophi and cirri of Balanomorphoidea; A and E, Chelonibia patula; B and F, Bathylasma corolliforme: C and G,
Aaptotasma leptoderma: D and H, Epopella breviscutum; I, Tesseropora pacifica. Labra of balanomorphoids are similar to
those of chthamaloids in being thick (although not bullate) and with variously concave crests (although there is a tendency to
form a shallow notch). In Chelonibia (A) the labrum is distinctly notched and multidenticulate, in a manner reminiscent of
Balanus amphitrite (Fig. 16 B). Mandibles of lower forms (A-C), in generally having four major teeth and pectinate inferior
angles, are similar to lower balanoids (Fig. 16, A|. In higher tetrachtines (D) the inferior portion becomes combUke, much as
in higher chthamaloids (Fig. 14, C), apparently an adaptation to life in the high intertidal. The first maxillae are essentially
balanoid. The third cirri resemble the fourth more than the second (E-G), as in chthamaloids, or one or both rami are antenniform
or bear specialized serrate setae (F, H and I).

Figure 16. Trophi and cirri of Balanoidea. A and E, Solidobalanus (Bathybalanus) pentacrini; B, Balanus amphitrite amphi-
trite; C and F, Balanus laevis; D, Hoekia monticulariae; G, Megabalanus psittacus; H, Balanus amphitrite inexpectatus;
I, Armatobalanus (Armatobalanus) terebratus. Acasta conica and Acasta nitida respectively; J, Armatobakmus allium, K, Balanus
nubilus. Labra of balanoids are generally thin and deeply incised; mandibles tend to have molariform rather than pectinated
or combed inferior portions; first maxillae are undistinguished (B and C). In primitive species (A), the labrum has but a shallow
notch and the mandible, in having an incisiform inferior portion, resembles that of the lower balanomorphoids. Marked de-
partures from this facies are seen in commensal forms such as the wholly parasitic coral barnacle, Hoekia (D).

The third cirrus always more closely resembles the second than the fourth, even in primitive species (E), and its rami
are never antenniform. Rather, it is the anterior ramus of the first cirrus that takes on antenniform characteristics in
some higher forms (F and K). Anterior cirri become variously thickened and in species without markedly speciahzed setae or
spines, the anterior margin of the articles may become markedly protuberant (G). Complex setae, as seen in many chthamaloids
and higher balanomorphoids, are not found in balanoids. Conversely, the so-called furcate (H) and multifurcate types
found in certain species of the group of Balanus amphitrite (Henry, 1973) have not been observed in the first two
superfamilies. On the other hand, compUcated arrays of spines commonly develop in balanoids (I and J), in free living forms,
but especially in commensals of sponges and corals where they may be used to clear the aperture and prevent overgrowth by
the host.

36

COMPOSITION AND DEFINITIONS OF SUPRAGENERIC TAXA

An abbreviated definition is given for all
suprageneric taxa. Where appropriate, the type
genus and related genera are indicated. For each
genus, the author, date and page, and the number
of fossil and extant species included are given.

BALANOMORPHA Pilsbry (1916: 14)

Thoracic cirripeds lacking peduncle; bilater-
ally symmetrical shell composed of carina,
rostrum, and one to three pairs of lateral com-
partmental plates that may be variously fused
or totally concrescent; opercular valves paired
when present, with members of each pair
separate, articulated or concrescent; hermaphro-
ditic (a few species of Archaeobalaninae have
complemental males).

CHTHAMALOIDEA Darwin (1854b: 446)
n. status

Wall composed of rostrum and one to three
pairs of laterals; rarely supplemented with one
or more whorls of imbricating plates around
basal margin; rostrum rarely compound; parietes
solid; radii solid; internally wall lacks uniform
ribs; articulation of opercular valves generally
deep, articulating pairs occasionally secondarily
cemented or calcified together; basis commonly
membranous, when calcareous, solid, and not
forming complex interdigitations with wall;
labrum bullate; crest nearly straight or shallowly
concave, but without medial incision; mandible
tri- or quadridentoid, with teeth usually simple;
inferior angle finely pectinate or coarsely serrate;
cirrus III resembling IV more than II; cirrus II
frequently with specialized terminal setae; cirri
lacking specialized hooks and spines; anterior
ramus of cirrus III occasionally antenniform;
penis without basidorsal point; caudal
appendages when present multiarticulate.

CHTHAMALIDAE Darwin (1854b: 446)

Wall of 8, 6, or 4 plates; lacking basal whorl
of supplementary plates; mandible tridentoid or
quadridentoid.

PACHYLASMINAE Utinomi (1968a: 36)

Wall of 8, 6, or 4 plates; wall sutures finely
denticulate; rostrum compound or with weakly
developed alae; scutum higher than wide: basis
commonly calcareous; mandible tridentoid;
commonly with caudal appendages.

Genus: Pachylasma Darwin (1854b: 475), type

genus, 9 spp.

EURAPHIINAE n. subfam.

(Group of C. hembeli.
Nilsson-Cantell, 1921: 275)

Wall of 8 or 6 plates; sutures often coarsely

serrate; rostrum with well developed alae;

scutum higher than wide; basis commonly

calcareous; mandible tridentoid; generally lacking

caudal appendages.

Genera: Euraphia Conrad (1837: 261), type

genus, 10 spp.; Octomeris Sowerby (1825: 326),

3 spp.

CHTHAMALINAE Darwin (1854b: 446)

(Group of C. stellatus,

Nilsson-Cantell. 1921: 275)

Wall of 6 or 4 plates; sutures usually finely

denticulate; rostrum with well developed alae or

rarely compound; scutum wider than high; basis

membranous; mandible quadridentoid; teeth two

through four commonly with subsidiary cusps;

generally lacking caudal appendages.

Genera: Chthamalus Ranzani (1817: 276), type

genus, 24 spp.; Jehlius Ross (1971b: 269),

1 sp.; Tetrachthamalus Newman (1967a: 425),

1 sp.; Chamaesipho Darwin (1854b: 470), 3 spp.

CATOPHRAGMIDAE Utinomi (1968a: 36)
n. status

Wall of 8 or 6 plates; having one or more
basal whorls of supplementary plates; mandible
tridentoid.
Genera: Catophragmus Sowerby (1826: 328),
type genus, 1 sp.; Catomerus Pilsbry (1916:
335), 1 sp.; Pachydiadema Withers (1935: 389),
1 sp.; Chionelasmus Pilsbry (1911: 82), 1 sp.

BALANOMORPHOIDEA n. superfam.

Wall composed of rostrum, carina, and one
to two pairs of laterals; rostrum compound;
parietes solid or tubiferous; when tubiferous
often secondarily filled with chitinous and(or)
calcareous material; radii solid or tubiferous;
internal surface of compartments generally with-
out uniform ribs; articulations between pairs of
opercular valves generally shallow, valves never
calcified together secondarily; basis commonly

37

membranous, when calcareous solid and not
forming complex interdigitations with wall;
labrum thick, weakly bullate; crest nearly
straight or shallowly concave, frequently with
median depression, rarely with medial incision;
mandible quadridentoid; teeth simple or teeth
two through four with subsidiary cusps; inferior
angle finely pectinate or coarsely serrate; cirrus
III resembling II more than IV or more or less
intermediate between II and IV; cirri without
speciahzed spines or hooks, but cirri II and III
may be armed with specialized setae; rami of
cirrus III normal, or inner, outer or both rami
antenniform; penis lacking basidorsal point;
caudal appendages lacking.

CORONULIDAE Leach (1817: 68)

Wall of 8 (rostrum discernibly tripartite) or
6 plates; plates of six-plated forms with or with-
out a median longitudinal sulcus; parietes tubifer-
ous; tubes formed between inner and outer lam-
ina, between internal buttresses, or between ex-
ternal ribs; interlaminate figures simple, dendritic
or anastamosing; radii solid; basis membranous;
opercular plates when present, reduced, not
articulated and not occluding aperture.

CHELONIBIINAE Pilsbry (1916: 262)

Wall of 8 or 6 plates, each lacking a median
longitudinal sulcus; opercular plates weakly
articulated; terga well developed; borders of
mantle not forming a hood over the cirri; one
row of confluent wall tubes formed between irmer
and outer lamina.

Genera: Chelonibia Leach (1817: 68), type

genus, 12 spp.

EMERSONIINAE Ross (1967: 7)

Wall presumably of 6 plates, each lacking a
median longitudinal sulcus; several rows of
vertically discontinuous wall tubes between
inner and outer lamina.

Genus: Emersonius Ross (in Ross and Newman,

1967: 7), type genus, 1 sp.

CORONULINAE Leach (1817: 68)

Wall of 6 plates, each lacking a median
longitudinal sulcus; terga vestigial; opercular
plates lacking in Xenobalanus; borders of mantle
forming a hood over the cirri; single row of wall

tubes formed by infoldings of outer lamina

against the sheath.
Genera: Coronula Lamarck (1802: 464), type
genus, 8 spp.; Cetopirus Ranzani (1817: 276),
1 sp.; Cetolepas Zullo (1969a: 17), 1 sp.;
Cryptolepas Dail (1872: 300), 2 sp.; Tubicinella
Lamarck (1802: 461), 1 sp.; Xenobalanus
Steenstrup (1851: pi. 3), 1 sp.

BATHYLASMATIDAE Newman and Ross
(1971: 138)

Wall of 6 or 4 plates; parietes solid and lack-
ing regular internal ribs, or with chitin-filled
longitudinal tubes arranged in a single row;
plates lacking radii; inferior margin of mandible
commonly pointed, bearing a few small spines;
all cirri lacking specialized setae; one or both
rami of cirrus III and occasionally cirrus II may
be antenniform.

BATHYLASMATINAE n. status

Wall of 6 or 4 plates; wall not permeated by
tubes; basis membranous, but inner shelf may
form by secondary calcification; scuta oriented
essentially perpendicular to basis; tergum lack-
ing distinct spur; cirrus II resembhng III more
than I.
Genera: Bathylasma Newman and Ross (1971:
143), type genus, 3 spp.; Tessarelasma Withers
(1936: 591), 1 sp.; Tetrachaelasma Newman
and Ross (1971: 152), 1 sp.

HEXELASMINAE n. subfam.

Wall of 6 plates; permeated by chitin-filled

tubes; basis calcareous; scuta oriented essentially

parallel to basis; tergum with distinct spur;

cirrus II resembling I more than III.

Genera: Hexelasma Hoek (1913: 224), type

genus, 3 spp.; Aaptolasma Newman and Ross

(1971: 158), 5 spp.

TETRACLITIDAE Gruvel (1903b: 160)

Wall of 6 or 4 plates; parietes solid, or per-
meated by chitin, or having one or more rows of
tubes containing living tissue or secondarily
filled with calcareous and chitinous material;
radii well developed or obsolete, basis commonly
membranous; inferior margin of mandible pecti-
nate or coarsely serrate; cirrus II and III com-
monly armed with specialized setae; inner or
outer or both rami of cirrus III either normal or
antenniform.

38

AUSTROBALANINAE n. subfam.

Wall solid, or permeated by chitinous rods
or lamellae; radii solid, narrow or obsolete.
Genera: Austrobalanus Pilsbry (1916: 218 in
part, ref. to B. imperator only),' type genus,
1 sp.; Epopella Ross (1970: 3), 3 spp.

TETRACLITELLINAE n. subfam.

Wall tubiferous; tubes never filled: radii
tubiferous or solid, broad, well developed.
Genera: Tetraclitella Hiro (1939e: 273), type
genus, 10 spp.; Newmanella Ross (1969: 242),
1 sp.

TETRACLITINAE Gruvel (1903: 160)

Wall tubiferous; tubes commonly partly filled

with chitinous and calcareous material; radii

solid, narrow or obsolete:

Genera: Tetraclita Schumacher (1817: 91),

type genus, 18 spp.; Tesseropora Pilsbry (1916:

259), 5 spp.; Tesseroplax Ross (1969: 241), 1 sp.

BALANOIDEA Leach (1817: 68) n. status

Wall composed of rostrum, carina, and one
to two pairs of lateral compartments, or wholly
concrescent; parietes solid or tubiferous. when
tubiferous rarely secondarily filled; radii solid or
tubiferous; when basis calcareous internal sur-
faces of compartments commonly with uniform
ribs; basis commonly calcareous, solid or per-
meated by tubes, rarely membranous; when
calcareous commonly forming complex inter-
digitations with wall; opercular valves occlude
aperture; articulations between pairs generally
shallow, or fused; labrum thin, never bullate;
crest with pronounced medial incision; mandible
quadri- or quinquidentate; second and following
teeth with one or more subsidiary cusps; fifth
tooth often vestigial; inferior angle commonly
molariform; cirrus III resembUng II more than

'Darwin (1854b: 290) noted, on the basis of several shell
characters and the nature of the third cirrus, that Balanus
imperator was closer to Tetraclita than to Balanus. but he
nonetheless assigned it to Balanus. Pilsbry (1916: 2181 pro-
posed the subgenus Austrobalanus, with Balanus imperator
as the type species. However. Ross (1971: 266) noted that
imperator was not a Balanus. but a six-plated tetracUtid,
and subsequent studies on arthropodal structures confirms
this affinity; Austrobalanus imperator is assigned to the
Tetraclitidae herein. This change necessitates erecting a
new genus for the remaining three taxa originally assigned
to Austrobalanus by Pilsbry. We propose Notobalfinus Ross,
herein (Gr. notos. southern, and Balanus). with Balanus flos-
culus Darwin, 1854b, as the type species, and assign this
genus to the Archaeobalanidae herein. The species assigned
to Notobalanus may be characterized as follows: shell small,
non-tubiferous: inner basal surface bears irregular ridges;
radii narrow; basis calcareous, and non-tubiferous; scutum
with crests for insertion of lateral depressor muscle.

IV; cirri usually without specialized setae, but not
infrequently armed with specialized hooks and
spines; rami of cirrus II or III never antenni-
form; rami of cirrus I subequal or grossly
unequal; lacking caudal apendages; penis with
basi-dorsal point (rudiment thereof in Semibala-
ninae).

ARCHAEOBALANIDAE n. fam.

Wall of 6 or 4 plates; parietes solid, rarely
tubiferous; tubes uniformly or irregularly ar-
ranged and formed between inner and outer
laminae; when regularly arranged interlaminate
fingers simple, hnear; radii solid; basis commonly
calcareous, rarely tubiferous.

ARCHAEOBALANINAE n. subfam.

Wall of 6 or 4 plates; parietes solid or
tubiferous; when tubiferous, tubes uniformly
arranged in single row; interlaminate figures
simple; basis calcareous or membranous, when
membranous wall sohd.
Genera: Archaeobalanus Menesini (1971: 9)
type genus, 1 sp.; Actinobalanus Moroni (1967:
923), 7 spp.; Kathpalmeria Ross (1965a: 61),
2 spp.; Armatobalanus s. s. Hoek (1913: 159),
15 spp.; Armatobalanus (Hexecreusia) Zullo
(1961b: 72), 2 spp.; Chirona s. s. Gray (1835:
37), 6 spp.; Chirona (Striatobalanus) Hoek
(1913: 159), 8 spp.; Solidobalanus s. s. Hoek
(1913: 159), 15 spp.; Solidobalanus (Hesperi-
balanus) Pilsbry (1916: 192), 15 spp.; Solido-
balanus (Bathybalanus) Hoek (1913: 230), 1 sp.;
Notobalanus n. gen., 3 spp.'; Elminius Leach
(1825: 210), 3 spp.; Membranobalanus Hoek
(1913: 159), 7 spp.: Acasta Leach (1817: 69),
54 spp.; Conopea Say (1822: 323), 16 spp.;
Pseudoacasta Nilsson-Cantell (1930b: 11),
1 sp.; Eoceratoconcha Newman and Ladd
(1974: 387), 2 spp.

SEMIBALANINAE n. subfam.

Wall of 6 plates; parietes tubiferous; basally
tubes irregularly spaced, not in discrete rows;
interlaminate figures lacking; basis membranous.

Genus: Semibalanus Pilsbry (1916: 182), type

genus, 5 spp.

PYRGOMATIDAE Gray (1825: 104)

Wall of 4 plates or wholly concrescent;
parietes solid or tubiferous; when tubiferous
tubes occur between outer lamina and sheath, or
between external ribs of wall; interlaminate
figures complex, essentially arborescent; radii
solid; basis calcareous, rarely tubiferous, mem-
branous in Pyrgopsella.

39

PYRGOMATINAE Gray (1825: 102)

Wall of 4 plates or wholly concrescent; oper-
cular valves normal or modified; when normal,
tergum with weakly developed lateral depressor
muscle crests, or crests lacking; when shell con-
crescent, sheath lacking paired sulci.

Genera: Pyrgoma Leach (1817: 68), type genus,

1 sp.; Cantellius Ross and Newman (1973:

150), 17 spp.; Creusia Leach (1817: 68), 3 spp.;

Hiroa Ross and Newman (1973: 153), 1 sp.;

Hoekia Ross and Newman (1973: 161), 1 sp.;

Nobia Sowerby (1823: no pagination), 6 spp.;

Savignium Leach (1825: 210), 4 spp.; Pyrgop-

sella Zullo (1967a: 123), 2 spp.

CERATOCONCHINAE n. subfam.

Wall of 4 plates; opercular valves normal;
tergum with a single large crest for lateral
depressor muscle.

Genus: Ceratoconcha Kramberger-Gorjanovic

(1889: 50), type genus, 21 spp.

BOSCIINAE n. subfam.

Wall wholly concrescent; opercular valves
normal; tergum with feebly developed lateral
depressor muscle crests, or crests lacking; sheath
with paired sulci.

Genus: Boscia Ferussac (1822: 145), type

genus, 4 spp.

BALANIDAE Leach (1817: 68)

Wall of 6 or 4 plates; parietes tubiferous;
tubes basically in single uniform row formed
between inner and outer laminate although
supplementary tubes may form basally; inter-
laminate figures complex, arborescent; radii
either solid or tubiferous; basis calcareous,
commonly tubiferous.

Genera: Balanus DaCosta (1778: 248), type
genus, 131 spp.; Megabalanus Hoek (1913:
158), 49 spp.; Tetrabalanus Cornwall (1941:
227), 1 sp.

40

CATALOG OF SPECIES

Superfamily Chthamaloidea Darwin, 1854, n. status
Family Catophragmidae Utinomi, 1968

Genus Catophragmus Sowerby. 1826

Catophragmus imbricatus Sowerby, 1827, figs. 1-6

Synonymy DiAC.NOsis: Henry. 1958: 217.

References: Broch, 1922:298 (as Catophragmus pilsbryi
n. sp.); Darwin, 1854b:490: Gruvel, 1905a:196; Pilsbry,
1916:335, VerriU, 1901:22; Weltner. 1897:274.

Distribution: Atlantic: Antigua, Bermuda; Pacific:
Panama, Costa Rica.

Genus Pachydiadema Withers, 1935

Pachydiadema cretaceum Withers, 1935:390
Distribution: Upper Senonian (Cretaceous), Ifo, Sweden
(Withers 1953:103).

Genus Catomerus Pilsbry, 1916

Catomerus polymerus (Darwin), 1854b:487

Synonymy diagnosis: Pope. 1965:16.

References; Barnes & Klepal, 1971:79 (pedicel of penis);
Broch, 1922:299, 301; 1927a:.506; Bennett & Pope, 1953:
105; 1960:182; Dakin et al, 1948:176; Endean et al,
1956:88; Gruvel, 1903b:lll; 1905a:195; Guiler, 1952:20;
NUsson-CanteU, 1926:8; Pilsbry. 1916:336; Pope, 1945:356;
Weltner, 1897:274; Wisely & BUck, 1964:162 (first stage
naupUi); Womersley & Edmonds, 1958:217.

Distribution: Southeast Australia.

Genus Chionelasmus Pilsbry, 1911

Chionelasmus darwini (Pilsbry), 1907c:188.
Synonymydiagnosis: Nilsson-Cantell, 1928b:446.
Reference.s: Gordon, 1970:105; Nilsson-Cantell, 1938b:14;

Pilsbry, 1911:82; 1916:335; Pope, 1965:10.
Distribution: Hawau; Rodriguez Is., Western Indian

Ocean; 450-460m.

Family Chthamalidae Darwin, 1854
Subfamily Pachylasminae Utinomi, 1968

Genus Pachylasma Darwin, 1854

Pachylasma aurantiacum Darwin. 1854b:480

Synonymy diagnosis: Darwin, 1854b:480.

References: Gruvel, 1905a:199; Weltner, 1897:273.

Distribution: New South Wales, Australia.
Pachylasma chinense Pilsbry, 1912:293

Synonymydiagnosis: Pilsbry, 1912:293.

Reference: Pilsbry, 1916:329.

Distribution: East China Sea; 400m.
Pachylasma crinoidophilum Pilsbry, 1911:81

Synonymy: Utinomi, 1968a:24.

Diaunosls: Pilsbry, 1911:81; Utinomi, 1968a;24.

References: Kruger, 1911b:460; Nilsson-Cantell, 1932a;
14; Utinomi, 1958a:307.

Distribution: Tokyo Bay to Kyusyu, Japan; 300-400m.
Pachylasma daru'inianum Pilsbry, 1912:293

Reference: Pilsbry, 1916:329."

Distribution: Sulu Arch.; 150m.
Pachylasma ecaudatum Hiro, 1939b:52

Synonymydiagnosis: Utinomi, 1968a:31 (as Hexelasma
ecaudatum).

Distribution: Ogaswara I.; 200m.
Pachylasma giganteum (Philippi), 1836:250

Synony.my DiAGNOSLS: Darwin, 1854b:477.

References: Gruvel, 1905a:198; Kolosvary, 1942c;143;

1943a:77; 1951c:412; Pilsbry, 1916:329; Rehni, 1969:169;

Stubbings, 1967:263; Weltner. 1897:273; Withers, 1953:

60,61.

Di.stribution: Mediterranean (Sicily); West coast of
Africa. Tertiary: Messina, Sicily.
Pachylasma integrirostrum Broch, 1931:50

Distribution: Kei Is.; 140m.
Pachylasma japonicum Hiro, 1933:65

Synonymy diagnosis: Utinomi, 1958a:22.

Reference: Hiro, 1937c:430.

Distribution: Southwest coast of Japan; 55-364m.
Pachylasma scutistriata Broch, 1922:301

Synonymy diagnosis: Utinomi, 1968a:26.

Reference: Nilsson-Cantell, 1927a:781.

DisTKiBurioN: Southern Japan, South China Sea to S.
Australia; 132-2050m.

Subfamily Euraphiinae n. subfam.

Genus Octomeris Sowerby, 1825

Octomeris angulosa Sowerby, 1825:244

Synonymy: Barnard, 1924:98.

DiAONOSl.s: Darwin, 1854b:483.

References: Barnes & Barnes, 1965a:391 (variation in
egg size); Barnes & Klepal, 1971:79 (pedicel of penis);
Gray, 1825:104 (as O. stuchburii n. sp); Gruvel, 1903b:
109; 1905a:197; Hiro, 1932b:478; Nilsson-CanteU, 1938b;
12; Pilsbry, 1916:334; Ritz & Foster, 1968:545 (tempera-
ture responses); Sandison, 1954:69 (nauplii); Stebbings,
1910:575, Weltner, 1897:274.

Distribution: South Africa.
Octomeris brunnea Darwin, 1854b:484

Synonymy/diagnosis: Pope, 1965:20.

References: Barnes & Klepal, 1971:79 (pedicel of penis);
Gruvel, 1903b:110; 1905a;197; Hiro, 1932b:471; 1939e:
252 (includes discussion of O. intermedia): Nilsson-
Cantell, 1921:303 (as Octomeris intermedia n. sp.); 1925:
1; 1930b:10; 1931a:108; 1932a:13; 1938b:33 (as O. inter-
media): Utinomi, 1949a:25; 1954:22; 1958a:307; Weltner,
1897:274; Withers, 1932:123 (as O. crassa n. sp.).

Distribiition: Southern Japan: Philippines; Indonesia;
New Hebrides; Australia; Mergui Arch.
Octomeris sulcata Nilsson-Cantell, 1932a:8

Synonymy: Utinomi, 1970:345.

Diagnosis: Hiro, 1939e:254.

References: Hiro, 1932b:471; 1939d:242; 1939f:207;
Ooishi, 1964:195; Rosell, 1973b:75; Utinomi, 1949a:21;
1970:345; Utinomi & Kikuchi, 1966:5.

Distribution: Southern Japan to Formosa.

Genus Euraphia Conrad, 1837

Euraphia aestuarii (Stubbings), 1963b:7

Synonymy: Stubbings, 1967:257.

Diagnosis: Stubbings, 1963b:7.

References: Gauld, 1957:10 (as Chthamalus stellatus
depressus): Kolosvary, 1941b:70 (as Chthamalus cirratus):
1943a:75 (as Chthamalus cirratus): Longhurst, 1958:32,
59 (as Chthamalus rhizophorae and C. withersi): Nilsson-
Cantell, 1938a:177 (as C s. depressus): Sandison, 1967:
166 (naupliar stages); Utinomi, 1968b: 169.

Distribution: West Africa.
Euraphia apelloefi (NUsson-CanteU), 1921:292

Synonymy diagnosis: Nilsson-Cantell, 1921:292.

References: Hiro, 1936d:229; Kolosvary, 1941b:70; Nilsson-

41

CanteU, 1926:1.

DisTHiiunioN: Java.
Euraphia calcareobasis (Henry), 1957:30

Rkkkrenik: Newman. 1961:148.

Distribution: Tuamoto Is.
Euraphia caudata (Pilsbry), 1916:315

Sv.NONV.MV DIAGNOSIS Pilsbry. 1916:315: Pope, 1965:35.

Refekencks: Endean et al, "l956:88: Foster, 1974:42; Hire.
1937b:51; Kolosvary, 1941b:70; NUsson-CanteU, 1921:278.
296: 1930b:8: 1932d:3; Rosell, 1972:184; Stephenson et al,
1958:268; Zevina & Tarasov, 1963:84.

Distribution: Australia; Philippines; Palau Is.; Indonesia.
Euraphia depressa (Poli). 1791:27

Synonymy: Southward. 1964b:241.

Diagnosis: Utinomi, 1959a:392.

References: Barnes, 1956c:309 (biometry); Barnes &
Barnes, 1964a:19 (exposure to air); 1964b:3 (distribution
and ecology); 1968a: 146 (variations in egg production);
Barnes & Klepal, 1971:77 (pedicel of penis); Carli, 1966a:
277 (mandible deformities); 1966b:115 (morphology and
ecology); Darwin, 1854b:456; Gauld, 1957:10; Gruvel,
1905a:"211; Hammen, 1972:435 (lactate oxidation); Have &
Have. 1954:330 (zonation); Kolosvary. 1939c:169; 1941b:
68; 1943:74; Monterosso. 1933:17 (morphology and biol-
ogy); Nilsson-CanteU. 1938a:177; Pilsbry, 1916:17 (mor-
phology and biology); Nilsson-Cantell. 1938a:177; Pilsbry,
1916:304; Ranzani, 1818:83 (as Chthamalus glaber n. sp.);
ReUni, 1964:402; 1969:170; Riedl, 1963:2.56; Stubbings,
1963a:7; TenerelU, 1952:92 (biology); Utinomi. 1959a:382
(as Chthamalus stellatus maxima): Weltner. 1897:273.

Distribution: Mediterranean: Gibraltar to Israel, Adriatic
and Black Seas.
Euraphia hembeli Conrad. 1837:261

Synonymy: Henry. 1957:29.

Diagnosis: Pilsbry. 1916:324; Newman, 1961:145.

References: Darwin, 1854b:465; Gruvel. 1905a:205
Gordon. 1970:107; Kolosvary, 1941b:70; Kruger, 1911a:4
1911b:460; Nilsson-Cantell, 1921:278, 290; Pilsbry, 1928
310; Weltner. 1897:272.

Distribution: Hawaiian. Caroline, and Sunda Is.; Ceylon.
Euraphia intertexta (Darwin) 1854b:467

Synonymy/diagnosis: Pope, 1965:29.

References: Foster, 1974:39; Gordon. 1970:110; Gruvel,
1905a:206; 1912a:349; Hiro. 1936d:227; 1939e:251; Hoek.
1913:269; Kolosvarv. 1941b:70; Newman. 1961:143;
Nilsson-CanteU. 1921:278; Pilsbry. 1916:324; 1928:310;
Tokioka. 1953:123; Utinomi. 1949:21; 1954:22: 1968:169.

Distribution: Indonesia north to Ryukyu and Tokara Is.,
eastward to Hawaii and Fitcairn I.
Euraphia pilsbryi (Hiro), 1936d:227

Synonymy/diagnosis: Hiro, 1936d:227.

References: Hiro, 1937c:429; 1938c:1687 (resistance to
exposure); Kolosvary, 1941b:70, 76 (forma typica and
neuseelandicus): 1943a:77; Ooishi. 1964:195; Utinomi,
1949a:21; 1954:21; 1958b:51; 1969b:51; 1970:345; Utinomi
& Kikuchi. 1966:5.

Distribution: Southern Japan.
Euraphia rhizophorae (de Oliveira), 1940b:379
Synonymy/diagnosis: de Ohveira, 1941:26.
References: Lacombe & Monteiro. 1974:633; Pope. 1965:

40; Stubbings. 1963b:ll.
Distribution: Bahamas; Panama; Brazil.
Euraphia withersi (Pilsbry). 1916:312
Synonymy/diagnosis: Pope. 1965:39.
References: Barnes & Klepal. 1971:77 (pedicel of penis);

Broch. 1931; 131; Hiro. 1937b:49: Karande. 1967:1245

(fouUng); Karande & Palekar. 1963b: 130 (breeding);

1966:148; Kolosvary. 1941b:70; Longhurst. 1958:59. 85

(C. aestuarii): Morton. 1973:491; Nilsson-Cantell. 1921:

295; 1930b:8: 1931a:107: 1938b:31; RoseU. 1972:182;

1973b:74; Stubbings. 1963b;ll: 1967:259; Utinomi.

1968b: 168; Zevina & Tarasov. 1963:83.
Distribution: Mergui Arch.; Australia; Philippines; India;

Madagascar.

Subfamily Chthamalinae Darwin. 1854

Genus Chthamalus Ranzani, 1817

Chthamalus angustitergum Pilsbry. 1916:305

Synonymy/diagnosis: Ross. 1968:2; Southward. 1975:20.

References: Barnes & Klepal. 1971:77 (pedicel of penis);
Henry. 1954:444; Kolosvary. 1939c:161; 1941b:68; Mar-
shall. 1953:435 (as C stellatus): Newell et al. 1959:209:
Nilsson-Cantell. 1933:506; 1939a:3; Pilsbry. 1927:37;
Smith et al. 1950:134; Stephensen & Stephensen. 1950:
389 (as C stellatus): 1954:80 (as C stellatus): Voss &
Voss. 1960:102 (as C stellatus): WeUs. 1966:92 (as C.
stellatus): Werner. 1967:70 (as C. stellatus).

Distribution: Caribbean.
Chthamalus anisopoma Pilsbry. 1916:317

Synonymy: Ross. 1962:8.

Diagnosis: Pilsbry. 1916:317.

References: Barnes & Barnes, 1965b:392 (variation in
egg size); Henry, 1942:127; 1943:372; 1960:144; Kolosvarv.
1941b:70; Nilsson-Cantell. 1921:276.

Distribution: Gulf of Cahfornia.
Chthamalus antennatus Darwin. 1854:460

Synonymy diagnosis: Pope, 1965:45.

References: Anderson, 1969:183 (embryology and
phylogeny); Barnes & Klepal. 1971:77 (structure of the
penis); Bennett & Pope. 1953:105; 1960:182; Broch.
1916:14; 1922:305; Dakin et al. 1948:176; Endean et al.
1956:88; Gruvel. 1903b:113; 1905a:203; 1911:292. 1912a:
349; 1920:52; Guiler. 1952:20: Kolosvary. 1941b:70;
Nilsson-Cantell. 1921:277. 285; 1926:10; 1927a:781; Pils-
bry. 1916:296 (footnote); Pope. 1945:3.56; Rosell. 1972:
174; 1973b:74; Utinomi. 1968b:170; Weltner. 1897:271;
1900:308; Wisely & Blick. 1964:163 (nauplii): Womersley
& Edmonds. 1958:214 (ecology).

Disthibution: Australia; Tasmania.
Chthamalus antiquus PhiHppi. 1887:224

Synony.MY: Ortmann, 1902:250 (? = Balanus varians
Sowerby).

Distribution: Miocene, Chile.
Chthamalus belyaevi Zevina & Kurshakova. 1973:187

Distribution; Easter Is.; southeast Pacific.
Chthamalus challengeri Hoek. 1883:165

Synonymy: Hiro. 1932a:546.

Diagnosis: Nilsson-Cantell. 1921:279.

References: Barnes & Klepal. 1971:77 (pedicel of penis);
Bhatt & Bal. 1960:439; Broch. 1927d:136; 1931:53 (as C
challengeri forma krakatauensis nov.); 1947:5; Gruvel.
1903b:113; 1905a:203; Hiro. 1932b:469; 1935c:215. 227;
1937c:429; 1938c:1687 (resistance to salinity and insola-
tion); 1939a:128; 1939f:207; Kolosvary. 194ib:70; 1943a:
75; Kruger. 1911a:46; 1911b:460; Luckens. 1968:75
(breeding and settlement); 1969:251 (breeding and settle-
ment): 1970a:35 (predation and zonation); 1970b: 161
(seasonal distribution); Nilsson-Cantell. 1921:279; 1925:
23; 1927a: 781; 1932b:8; 1932e:2; 1938b:31; Pilsbry.
1916:307; Pope. 1965:52; Tarasov & Zevina, 1957:256;
Utinomi. 1949a:21; 1954:25; 1958b:51; 1962:215; 1969b:
51; 1970:345; Utinomi & Kikuchi 1966:5; Weltner. 1897:
272, Zevina & Litvinova, 1970:174; Zevina & Tarasov,
1963:79.

Distribution: Japan; Bonin Is.; Philippines; Indonesia;
Indian Ocean; Red Sea.
Chthamalus challengeri krakatauensis Broch. 1931:53

Synonymy: Hiro, 1939e:249 (= C. mora Pilsbry); Karande
& Palekar, 1963a:231 (= C. malayensis).
Chthamalus challengeri nipponensis Pilsbry, 1916:309

Synonymy: Nilsson-Cantell, 1921:279 (= C. c. challengeri).
Chthamalus cirratus Darwin, 1854b:461

Synonymy/diagnosis: Pilsbry, 1916:321.

References: Gruvel, 1903"b:113; 1905a:202; 1912a:349:
Kolosvarv, 1941b:70; 1943a:75; Nilsson-CanteU, 1921
277; 1957:16; Pilsbry, 1909:71; Weltner. 1897:272:
1898b:6; 1900:305; Zevina & Kurshakova. 1973:183.

Distribution: Chile; Peru; Ecuador.

42

Chthamalus dalli Pilsbry. 1916:316

Synonymy; Cornwall. 1955b:23.

Diagnosis: Pilsbry. 1916:316: Henry. 1940a:17.

Rkfkkencks: Barnes & Barnes. 1965a:392 Ivariation in
egg size): Barnes & Conor. 1958:194 (neurosecretory
cells): Barnes & Klepal. 1971:77 (pedicel of penis): Corn-
wall. 1925:472: 1937:232: 19.50:318; 1953:76 (nervous
system): 1955a:36: Dayton. 1971:351 (community organ-
ization): Henry. 1942:121; Hiro, 1932b:469: 1935c:215;
Kolosvary, 1941b:70: 1943a:76: Nilsson-Cantell. 1921:
277; Rice. 1930:249 (distribution in communities): South-
ward & Southward. 1967:8 (biology); Stallcup. 1953:143;
Tarasov & Zevina. 1957:256; Utinomi. 1970:345.

DlsiKdUTiiiN: Unalaska to central California; northern
Japan.
Chthamalus dentatus Krauss. 1848:135

Synony.my DI.AC.Ndsis: Stubbings. 1967:252.

Rkkkkencks: Barnard. 1924:97: Barnes & Barnes. 1965a;
392 (variation in egg size): Barnes & Klepal, 1971:77
(structure of the penis); Broch. 1924b:202; Darwin,
1854b:463: Day & Morgans. 1956:303: Gauld, 1957:10:
Gruvel. 1903b;il3: 1905a:204; 1912a:345; Hoek. 1883:164;
1913:xvii; Kolosvary. 1941b:68; Millard. 1950:270; Mil-
lard & Broekhuysen, 1970:298; Nilsson-CanteU. 1921:277,
282; 1931a:107; 1938a:176; Ritz & Foster. 1968:553
(temperature response): Sandison. 1954:94; Stebbing,
1910:574: Stubbings. 1961b:19; 1961c:183: 1963b:13
1964b:333; 1965:885; Utinomi. 1968b;169; Weltner.
1897:272.

Di.sTRlBUTlON: West coast of Africa as far north as Cape
Verde Is., southeastern coast of Africa north to
Madagascar and Mauritius.
Chthamalus fissus Darwin, 1854b:462

Synon"! .\iy; Ross. 1962:36.

Diagnosis: Henry. 1942:121.

Refkrenck.s: Augenfeld. 1967:92 (metaboUsm): Barnes &
Barnes, 1958a:550: 1959h:516 (metabolism); 1965a:392
(variation in egg size); Barnes & Klepal, 1971:77 (struc-
ture of the penis); Broch. 1922:308; ConneU. 1970:49
(predation); Gruvel. 1903b:113: 190.5a:202: Henry. 1943:
368; 1960:144; Kolosvary. 1941b;71: 1943a:75; 1947e:
361: 1951b:292; Nilsson-CanteU. 1921:276; Pilsbry. 1916
317. Weltner. 1897:273.

DisrujiH I'Ion: San Francisco, south into Gulf of Cahfornia.
Chthamalus fragilis Darwin. 1854b:456

Syn<inymy diagnosis: Pilsbry. 1916:297: Stubbings. 1967:
262; Southward. 1975:19.

Rekkkences: Barnes & Klepal. 1971:77 (pedicel of penis);
Bousfield, 1954:123: Broch, 1927c:19; Crisp & South-
ward, 1961c:271 (cirral activity); Gordon. 1969:139 (in-
fluence of salinity); Gruvel. 19d3b;113; Henry. 1954:444;
Johnson. 1958:205 (fungal parasite in ova): Kolosvary,
1941b:68: 1943a:74: McDougall. 1943:351; Nilsson-
Cantell. 1921:277; 1928a:30: 1933:505; 1939a:3: Pilsbry,
1927:37; Visscher, 1928a:327 (attachment); Visscher &
Luce, 1928:336 (cyprid reaction to light); Wells, 1966:88;
Weltner, 1897:273: Zullo. 1963b:8.

DisrHiHurioN: Cape Cod. Massachusetts south to West
Indies: West Africa.
Chthamalus imperatrix Pilsbry. 1916:320

Synonymy uiAiiNosis: Pilsbry. 1916:320.

Reeerences: Kolosvary. 1941b:70; Nilsson-Cantell, 1921:
276.

Distriuution: Panama.
Chthamalus ligusticus deAlessandri. 1895:306

SvNONY.MY diagnosis: deAlessandri, 1906:283; Withers,
1953:61.

DisrHiHLi'noN: Pliocene, Italy.
Chthamalus malayensis Pilsbry. 1916:310

Synonymy: Utinomi. 1954:18; Karande & Palekar, 1963a:
231; Pope. 1965:51.

Diagnosis: Pope. 1965:51.

Reeekeni'Es: Barnes & Klepal. 1971:77 (structure of the
penis); Broch, 1916:14 (as C. antennatus); 1922:307 (as

C. mora}; 1931:53 (as C. challengeri forma krakatauensis),
55. 56 (as C moro): Daniel. 1955c:34 (as C stellatus
slellatus); Darwin. 1854b:455 (as C. stellatus): Endean et
al. 1956:88 (ecology and distribution): 1956:317 (ecology
and distribution); Foster. 1974:42; Gruvel. 1912a:345 (as
C. antennatus): Hiro. 1937b:49 (as C mora): 1939e:249
(as C mora). 250; Hoek. 1913:267 (as C. stellatus):
Karande. 1966:148; 1967:1245 (fouUng); Karande &
Palekar. 1963a:231; 1963b:130 (breeding activity);
Kolosvary. 1941b:70: 1943a;76; Kruger. 1914:435 (as C.
stellatus var. communis): Nilsson-Cantell. 1921:277 (as
C. mora): 279 (C. challengeri in part); 1934b:50 (C. mom):
1938b:30 (as C. stellatus stellatus), 31: Pilsbry. 1916:
304 (as C. stellatus stellatus): 311 (f. mora): Stephenson
et al. 1958:268 (insular ecology); Southward. 1964b:252;
Stubbings. 1936:49 (as C stellatus): 1961a:171; 1963a:
328; Rosell. 1972:178 (questions synonymy of C. mora
with C malavensis): Utinomi. i949a:2.5; 1968b: 169;
1969:82: Zevina and Tarasov. 1963:80.

Distribl'TIon; From Persian Gulf. India. Pakistan. Malay
and South China Seas to Formosa; also Indonesia.
Philippines, Palau Is.
Chthamalus microtretus Cornwall. 1937:232

Synonymy diagnosis: Cornwall. 1951:319: Newman, 1976:
269 (= C fissus).

Rkeerence: Henry. 1942:127 (distribution hst).
Chthamalus panamensis Pilsbry. 1916:319

Synony.my dia(_;nosis: Pilsbry. 1916:319.

References: Nilsson-Canteli. 1921:277; Kolosvary. 1941b:
70.

Distribution: Quarantine I.. Panama.
Chthamalus permitini Zevini & Litvinova. 1970:178

Disiribu I'ION: Red Sea (possibly C. malayensis).
Chthamalus scabrosus Darwin. 1854b:468.

Synonymy: Nilsson-Cantell. 1921:278.

Diagnoses: Pilsbry. 1916:323.

Refehences: Gruvel. 1903b:113; 190.5a:205; 1912a;349:
Kolosvary, 1941b:70; 1943a:76; Nilsson-Cantell, 1957:6;
Pilsbry. 1909:72; Weltner. 1895:291; 1897:272; 1898b:6:
1900:305; Zevina & Kurshakova. 1973:183.

DiMKiBi.1 I'Ion: Peru to Tierra del Fuego; Patagonia;
Falkland Is.
Chthamalus stellatus (Poll). 1791:29

S^NON'l^n: Southward. 1964b:241.

DiAcAosis: Southward. 1964b:247; Utinomi. 1959a:392.

Refkkenges: Annandale. 1906:149; Barnes. 1956b:355
(growth rate): 1956c:309 (biometry); Barnes & Barnes,
1958a;550 (self-fertilization): 1959h:515 (metabohsm);
1964b:l (distribution and ecology); 1965a:391 (variation
in egg size); 1966a:83 (ecological and zoogeographical
observations); 1966b:247 (recovery from severe winter);
1968a:135 (egg number and variation): 1969b:36 (seasonal
changes in o.\ygen consumption); 1974:197 (embryonic
development & salinity): Barnes & Crisp 1956d:631 (self-
fertilization); Barnes et al. 1963f;233 (dessication/anaerobic
conditions); 1970:70 (resistance to impaction); 1971:173
(spermatozoa); 1972:89 (body weight and biochemical
composition); Barnes & Klepal. 1971:77 (structure of the
penis); Bassindale. 1936:57 (developmental stages); 1958;
381; 1961:485; 1964:36; Bhatnagar & Crisp, 1965:419
(salinity tolerance of larvae): Bocquet-Vedrine, 1956:2159
(tidal rhythym and growth); 1957:1545 (parasite of)
1958a:484 (parasite of): 1958b:2440 (parasite of); 1961
549 (parasite of): 1963:1350 (structure of the shell)
1965a:469 (parasite of); Bocquet & Ovechko, 1959:106
(salivary glands); Borradaile. 1916:135; Broch. 1924b
203; 1927e:19; 1927d:136; Carh. 1966b:115; Caziot. 1921
54; Connell, 1957:1 (competition); 1961b;710 (competition)
Crisp, 1950:311 (breeding and distribution): 1964a:208
(effect of severe winter); Crisp & Patel, 1958:1078 (breed-
ing and ecdysis); Crisp & Southward, 1961:271 (cirral
activity): Daniel. 1955a:97 (gregariousness); 1955c:34;
1957a:305 (effect of illumination); 1957b:866 (tidal in-
fluence); Darwin, 1854b;455; de Alessandri, 1895:304;

43

1906:283: Fischer, 1872:434: Fischer-Piette, 1955:37
(distribution): Fischer-Piette & Prenant, 1956:18: Fishel-
son, 1971:126: Foster, 1970:377 (response to sahnity):
1971a:12 (dessication): Groom, 1894:119 (early develop-
ment); Gruvel, 1905a:201: 1907d:5: 1912a:345; 1920:52:
Hatton & Fischer-Piette. 1932:1 (settlement and growth):
Hoek, 1875:58; 1909:272; 1913:267: Kitching, 19.50:820:
Klepal & Barnes. 1975:269 (ecology): Knight-Jones.
1953:583 (gregariousness); 1955:266 (gregariousness);
Kolosvary. 1939a:178: 1941a:41: 1941b:68: 1943a:73:
1947a:31; Kruger. 1911a:45. 1911b:460: 1914:435: 1927a:
13: 1927b:4; LeReste. 1965:53 (larvae): Monod. 1933:7;
Monterosso, 1927-1932:(see bibliography): Moore. 1936:
701 (biology): Moore & Kitchmg. 1939:521 (biology);
Moyse. 1960:120 (rearing larvae); Moyse & Nelson-Smith.
1963:15 (zonation); Nilsson-Cantell. 1921:277, 281;
1931a:107; 1938b:30; 1939c:92; O'Riordan. 1967:291;
Patel & Crisp. 1960a:667 (influence of of temperature):
1960b:104 (rate of embrvonic development): Petriconi.
1969:539 (mouth parts); Pilsbry, 1916:302; Pope. 1965:
24; Powell. 1954:688: Prenant & Teissier. 1923:172;
Rehni. 1964:397: Riedl. 1963:256; Rosell. 1972:172;
1973b:73; Southward, 1950:408; 1951:410: 1955b:403
(behavior); 195.5c:423 (behavior); 1957:323 (behavior);
1962:162 (behavior); 1964a:391 (cirral activity and
temperature); 1965:441 (metabolism and survival);
Southward & Crisp. 1952:416 (distribution); 1954a:163
(distribution); 19.56:211 (distribution): 1963:38 (fouUng
organisms); Stubbings. 19.36:49; 1961b:18; 1963b:6;
1964a:107; 1965:885: 1967:251; Summer. 1909:373: 1911:
128: Tarasov & Zevina. 1957:253: Tenerelli. 1952:122;
1958:263; 1959a:l (fertilization); 1959b:14 (female sex
apparatus); Utinomi. 1959a:381: Visscher. 1928b:193
(survival in freshwater): Wellner. 1895:291: 1897:272;
1898a:443; 1898b:9; Williams. 1950:311; Zevina, 1963:73.

Distribution: British Isles: coasts of France, Portugal and
Spain; Mediterranean and Black Seas; western coast of
Africa to Cape Verde. Scattered records from Indo-
Pacific need verification.
Chthamalus stellatus hisinuatus Pilsbry, 1916:306

S'lNDN'iMV iiiACNdsis: Pilsbrv. 1916:306: de Uliveira. 1941;
24; Southward. 1975:28.

References Kolosvary, 1941b:68; Lacombe & Monteiro,
1974:633; Nilsson-Cantell, 1931a:107; Stubbings, 1961b:
19: 1967:252; Wells, 1966:88.

DiSTRiHirrios: Rio de Janeiro and Santa Catarina Is..
Brazil; Lagos. Nigeria; St. Andrews Bay. Florida.
Chthamalus stellatus cornutus Nilsson-Cantell. 1925:25

Disi'iiiBUTlDN: St. Vincent. Brazil; Isla de Flores. Uruguay.
Chthamalus stellatus thompsoni Henry, 1958:220

DisiKiBUTiON: Bermuda.

Genus Jehlius Ross, 1971

Jehlius gilmorei Ross, 1971:271

Disi kibuikin: Islas San Ambrosio and San Felix. Chile
(Zevina & Kurshakova. 1973:184).

Genus Tetrachthamalus Newman. 1967

Tetrachthamalus oblitteratus Newman. 1967:425
References: Achituv. 1972:126 (zonation); Fishelson.
1971:113 (ecology): Morton. 1973:491; Southward. 1967:
437 (ecology and cirral activity): Taylor. 1968:146
(ecology): Zevina & Litvinova. 1970:174.
Distribution: Gulfs of Aqaba and Suez: Seychelles;
Mauritius; Aldabra.

Genus Chamaesipho Darwin, 1854

Chamaesipho brunnea Moore, 1944:320

Synonymy diagnosis: Moore, 1944:320.

References: Foster. 1967a:85; 1967b:33 (early stages);
Luckens. 1970c:497 (breeding and settlement); Pope,
1965:63: Ritz and Foster, 1968:545 (comparative tem-
perature responses).

Distribution: New Zealand.
Chamaesipho columna (Spengler). 1790:192

Synonymy uiAONosis: Moore. 1944:316.

Rkfehences; Anderson. 1969:183 (embryology); Barnes &
Klepal, 1971:79 (structure of the penis): Broch, 1922:308
Bennett & Pope 1953:105: 1960:182; Dakin et al, 1948
176; Darwin. 1954b:470: Endean et al. 1956:88; Filhol
1885:489; Foster, 1967a:84; 1967b:33 (early stages)
Gruvel, 1903b:159; 190.5a:282; Guiler, 1952:20: Hutton
1879:329; Jennings, 1918:63; Linzey, 1942a:280: Luckens,
1970c;497 (breeding and settlement); Moore, 1944:316
Nilsson-CanteU, 1926:11; Pope, 1945:357; Ritz and Foster,
1968:545 (comparative temperature responses): Weltner,
1897:273; 1899a:445; 1900:308; Wisely and Bhck, 1964:
162 (abundance of first stage nauplii); Womersley &
Edmonds, 1958:232 (ecology).

Distrihuiton: Australia: New Zealand.
Chamaesipho scutelliformis Darwin, 1854b:472

Synonymy uiAONosi.s: Darwin, 1854b:472; Zevina &
Tarasov, 1963:85.

References: Gruvel. 1903b:159; 1905a:283; Hoek. 1883:36;
Kruger. 19Ua:4; 1911b:461; Pope, 1965:64; Weltner,
1897:273.

Distribution: South China Sea.

Superfamily Balanomorphoidea n. superfam.

Family Coronulidae Leach, 1825

Subfamily Chelonibiinae Pilsbry, 1916

Genus Chelonibia Leach. 1817

Chelonibia capellini de Alessandri. 1895:300

Disiribution: Mio-Pliocene. Italy.
Chelonibia caretta (Spengler). 1790:185

Synonymy: Pilsbry. 1916:267.

Diagnosis: Darwin, 1854b:394.

References: Barnard, 1924:93; Borradaile, 1903:443;
Broch, 1924a: 16; Daniel, 195,5c:32; Dawydoff, 1952:129;
Gruvel, 1905a:269; Hinks, 1840:333 (as Balanus
cheltrypetes): Hiro, 1937b:69; Hoek, 1913:xvii
Kolosvarv, 1943a:99; Korschelt. 1933:13: Morch, 1852
67; Nilsson-Cantell, 1938b;14; Stubbings, 1967:297
Utinomi, 1969a:92: Wells, 1966:68; Weltner, 1897:254
Withers, 1928a:391; ZuUo. 1963b:13.

DisTKiBLi'iTos: Tropical Atlantic and Indo-West Pacific;
Miocene. Zanzibar.
Chelonibia depressa Seguenza. 1876:411

DisTRiBUi ion: Pliocene. Sicily.
Chelonibia hemisphaerica Rothpletz & Simonelli, 1890:724

Distribution: Pliocene, Grand Canary I.
Chelonibia manati Gruvel. 1903b:116

Synony.my diagnosis: Stubbings. 1965:894.

References: Broch. 1924b:203; Gruvel, 1905a:267; Hiro,
1936a:61 (commensahsm); Korschelt, 1933:17; Pilsbry,
1916:265; Stubbings, 1967:297: Utinomi. 1950:62.

Distribution: West Africa: on skin of manatees.
Chelonibia manati crenatibasis Pilsbry. 1916:266

Dlstribution: Unknown; probably from loggerhead turtle.
Chelonibia manati lobatobasis Pilsbry. 1916:266

References: Henry, 1954:444; Kolosvarv, 1942c:146:
WeUs. 1966:86.

Distribution: Florida; on turtles.
Chelonibia patula (Ranzani), 1818:86

Synonymy.'DIAGnosis: Pilsbry, 1916:268: Stubbings, 1967:
297.

References: Broch, 1924b:203; 1927d:136; 1935:2; 1947:7;
Crisp & Costlow. 1963:22 (sahnity tolerance); Daniel,
1955:32; Darwin, 1854b:396: Dawydoff, 1952:129: Ed-
mondson, 1933:231; Gauld, 1957:10; Gordon, 1970:90:
Gruvel, 1905a:268; 1907d:8; 1912a:346: Henry, 1954:444;
Hiro, 1936a:60 (commensalism); Hoek, 1913:xvii;
Kolosvary, 1943a:98; Korschelt, 1933:17; Kruger, 1911a:

44

4; 1911b:461; McDougall, 1943:343; NUsson-Cantell
1934b:61: 1938b:77; Pearse. 1947:327: 1952:7; Relini
1969:169; Ross, 1963b:225; Ross & Jackson, 1972:203
Ross and Newman, 1967:18; Sandeen & Costlow, 1961
192 (pigment activators); Southward & Crisp, 1963:26
Stubbings, 1961b:38; Utinomi. 1950:62; 1958a:309; WeUs,
1966:86; Weltner, 1897:254; Williams & Porter. 1964
150; Withers, 1929b:569; Zevina & Litvinova, 1970:174
ZuUo, 1963b:14.

Distribution: Tropical Atlantic to Indo-West Pacific.
Miocene, Paris Basin.
Ckelonibia patula dentata Henry, 1943:370

Reference: Henry, 1960:147.

Distribution: Sonora, Mexico; on crab.
Chelonibia ramosa Korschelt, 1933:2

Reference: Hiro, 1936a:61 (commensaUsm).
Chelonibia testudinaria (Linnaeus), 1757:668

Synonymy: Nilsson-Cantell, 1921:369.

Diagnosis Nilsson-Cantell, 1921:369; Daniel, 1955c:31.

References: Annandale, 1906:138; Barnard, 1924:92
Borradaile, 1903:443; Broch, 1916:14; 1924b:202; 1931
122; 1947:7; Caziot. 1921:51; Darwin, 1854b:392
Dawydoff, 1952:129; de Alessandri, 1895:391; 1906:314
Edmondson & Ingram, 1939:258; Fischer. 1884:355
Gauld, 1957:10; Gordon, 1970:94; Gruvel, 1903b:115
1905a:267; 1907d:8; Henry. 1941:105; 1943:371; 1954
444; 1960:147; Hiro. 1936a:61 (commensaUsm); 1937b
69; 1937c:470; 1939f:214; Hoek, 1913:xvii; Kolosvary
1942c:149; 1943a:99; 1951c:411; 1967b:392; Korschelt
1933:16; Kriiger, 1911a:57; 1911b:461; Lanchester, 1902
371; Linnaeus, 1767:1108; MacDonald, 1929:537; Morch
1852:67; Newman et al, 1969:R289; Nilsson-Cantell
1930b:19; 1931a:116; 1932d:258; 1938b:77; 1939a:5:
1957:7; Pillai, 1958:126 (larval stages); Pilsbry, 1916:264
1928:316; Relini, 1969:169; Riedl, 1963:258; Ross, 1963b:
227; Ross and Newman, 1967:18; Stubbings, 1965:893
1967:296; Utinomi. 1949a:24; 1958a:309; 1969a:92
1969b:53; 1970:359; Utinomi & Kikuchi, 1966:8; Weltner,
1895:298; 1897:254; 1899a:443; 1910:528; WeUs, 1966
86;ZuUo, 1963b:14.

Distribution: All temperate and tropical seas, attached
to turtles. Miocene, Cuba; Pliocene. Italy; Pleistocene,
Florida.
Chelonibia testudinaria solida Withers, 1929b:568

Synonymy: Ross, 1963b:230.

References: Ross & Newman, 1967:19.

Distribution: Mio- Pliocene, France; Pleistocene, Florida.

Platylepas hexastylos (Fabricius), 1798:35

Synonymy: Pilsbry. 1916:285.

Diagnosis: Pilsbry, 1916:285; Hiro, 1937c:472 (mouthparts)

References: Barnes & Klepal, 1971:89 (pedicel of penis)
Broch, 1924a: 18; 1924b:203; 1927c:30; Daniel, 1955c:33
Darwin, 1854b:428 (as P. bissexlobata): Fischer, 1884
359; Gruvel, 1903b:151; 1905a:276; Henry, 1954:444
Hiro, 1936a:62 (commensaUsm); 1936e:319; Kolosvary
1943a:101; 1951c:412; Korschelt, 1933:22; Kriiger, 1912:
13; ReUni, 1969:169; Richards, 1930:143; Schwartz
1960:116; Stubbings, 1965:899; 1967:300; Utinomi, 1950:
62; 1959a:384; Weltner, 1897:253; ZuUo. 1963b:15.

Distribution: All tropical and sub-tropical seas; on turtles,
manatees, dugongs.
Platylepas hexastylos ichthyophila Pilsbry, 1916:287

Reference: Ryder. 1879:453 (as P. decomta).

Distribution: On garfish; Florida.
Platylepas indicus Daniel, 1958b:755

Distribution: Madras, India; on sea snakes.
Platylepas krugeri (Kriiger), 1912:12

Synonymy: Ross, 1963a:153.

Distribution: Thailand.
Platylepas midtidecorata Daniel, 1962b:641

Distribution: Little Andaman I.; on green turtle.
Platylepas ophiophilus Lanchester, 1902:371

Synonymy/diagnosis: Utinomi, 1970:360.

References: Broch, 1931:122; Darwin, 1854b:430; Gruvel,
1905a:277; Hiro, 1936a:61 (commensaUsm); 1936e:319;
Korschelt, 1933:22; Kruger, 1912:12; Nilsson-CanteU.
1921:376; 1938b:77; Pilsbry, 1916:285; (renames Cry-
ptolepas ophiophlus Kruger as Platylepas krugeri).

Distribution: Sea of Japan; Indonesia; western AustraUa;
India; Arabian Sea; on sea snakes.
Platylepas wilsoni Ross, 1963a:153

Distribution: Pleistocene. Florida.

Genus Stomatolepas Pilsbry, 1910

Stomatolepas elegans (Costa), 1838:17

Synonymy: Hiro. 1937c:473.

Diagnosis: Pilsbry, 1916:289; Hiro. 1936e:314 (includes S.
praegustator Pilsbry 1916:289 and questionably S.
transversa Nilsson-CanteU. 1930a:2).

References: Henry, 1954:444; Hiro, 1936a:61 (com-
mensaUsm); Holthuis, 1969:44; Nilsson-CanteU, 1930b:
20; Pilsbry, 1910:304; ReUni, 1968a:223; 1969:169; Stub-
bings, 1965:902; 1967:300; Utinomi. 1970:363: WeUs,
1966:87; ZuUo & Bleakney, 1966:162.

Distribution: CosmopoUtan; soft skin and throat of sea
turtles.

Subfamily Emersoniinae Ross, 1967

Genus Emersonius Ross, 1967

Emersonius cybosyrinx Ross, in Ross & Newman, 1967c:8
Reference: Newman et al, 1969:R290.
Distribution: Upper Eocene, Florida.

Subfamily Platylepadinae n. subfam.

Genus Platylepas Gray, 1825

Platylepas decorata Darwin, 1854b:429
Synonymy/diagnosis: Nilsson-CanteU, 1921:376.
References Gruvel, 1905a:276; 1912a:350; Hiro, 1936a:61

(commensaUsm); 1936e:319; 1937b:70; Korschelt, 1933:22;

NUsson-CanteU, 1921:376; Utinomi, 1970:363; Weltner,

1897:253.
Distribution: Galapagos, through Pacific Oceania to

western coast of AustraUa; on turtles and sea snakes.

Genus Cylindrolepas Pilsbry, 1916

Cylindrolepas darwiniana Pilsbry, 1916:288
Reference: Hiro, 1936e:319.
Distribution: West Indies; in skin of sea turtle.

Genus Stephanolepas Fischer, 1886

Stephanolepas muricata Fischer, 1886:193
Synonymy/diagnosis Nilsson-CanteU, 1932d:258.
References Broch, 1947:7; Dawydoff, 1952:128; Gruvel,

1903b:151; 1905a:280; Hiro, 1936a:61 (commensaUsm);

1936e:318; Hoek, 1913:xvii; Nilsson-CanteU, 1938b:14;

Weltner, 1897:253.
Distribution: South China Sea; Ceylon; in skin of turtle.

Subfamily Coronulinae Leach, 1817

Genus Coronula Lamarck, 1802
Coronula aotea Fleming, 1959:243

45

Synonymy/diagnosis Beu, 1971:899.

Distribution: Pliocene. New Zealand.
Coronula barbara Darwin, 1854a:38

Synonymy/diagnosis Darwin, 1854b:421.

References Beu, 1971:900: Darwin, 1854a:38; de Ales-
sandri, 1895:303; 1906:317; Menesini, 1968a:395; Weis-
bord, 1971:91; Withers. 153:63; ZuUo, 1969a:21.

Distribution: Pliocene and early Pleistocene of Europe;
Pliocene, Southern California.
Coronula bifida Bronn, 1831:126

Synonymy/diagnosis: Darwin. 1854b:423.

References: de Alessandri. 1895:302; 1906:315; Menesini,
1968a:390; 1968b:584; Seguenza, 1876:324; Weisbord,
1971:94; Withers, 1953:63.

Distribution: Tertiary, Italy.
Coronula diadema (Linnaeus). 1767:1108

Synonymy: Pilsbry, 1916:273 (contains pre-Darwinian
references).

Diagnosis: Pilsbry, 1916:273: Cornwall, 1924a:421.

References: Barnard, 1924:94; Barnes & Klepal, 1971:88
(pedicel of penis); Bassindale, 1964:43; Beu, 1971:902
(Pleistocene, New Zealand); Borradaile, 1916:135; Broch,
1924a:91; Cornwall, 1927a:504; 1953:83 (nervous system);
1955a:51; 1955b:40; Crisp & Stubbings, 1957:179 (orien-
tation to water currents); Darwin, 1854b:417; Filhol,
1885:489; Fischer, 1872:433; Fleming, 1959:246 (fossil);
Gruvel, 1903b:152; 1905a:273; 1905b:308 (anatomy);
Guiler, 1956:3; Hatai, 1938:98; 1939a:262; Hayasaka,
1933:49; 1935:1; Henry. 1943:368; Hiro, 1935c:226;
1936e:318; 1937c:471; 1939f:214; Hoek, 1883:163; 1909:
271; Hutton, 1879:329; Jennings, 1918:62; Kolosvary,
1942a:138; 1942c:149; 1943a:99; 1967b:393; Korschelt,
1933:18; Mbrch, 1852:66; Newman & Ross, 1971:179;
Newman et al. 1969:R289; Nilsson-Cantell, 1921:371;
1930c:256; 1930d:212; 1931a:116; 1938b:14; 1939b:237;
1957:7; O'Riordan, 1967:294; PUsbry. 1916:273 (= Lepas
balaenaris Muller; Balanus bataena Da Costa; Diadema
vulgaris Schumacher; Diadema candidum Ranzani;
Polylepas kleinii Gray; Coronula biscayensis Van Beneden;
Diadema japonica Van Beneden; D. califomica Van
Beneden); Pilsbry & Olson, 1951:203 (= Diadema anti-
quum Philippi 1887:226); Scheffer, 1939:67; Stebbings,
1910:571; Stephensen, 1938:6; Tarasov & Zevina, 1957
241; Weisbord, 1971:94; Weltner, 1895:290; 1897:254
1898b:8; 1899b:102; 1900:302; 1922:86; Wolff, 1960:8:
ZuUo, 1963b:14.

Distribution: Cosmopolitan, on Humpback, Fin, Blue
and Sperm whales. Pliocene to Recent.
Coronula dormitor Pilsbry & Olson, 1951:202

Distribution: Pliocene, Ecuador.
Coronula ficarazzensis Gregorio, 1895:5

Distribution: Pleistocene, Italy.
Coronula macsotayi Weisbord, 1971:91

Distribution: Pliocene, Venezuela.
Coronula reginae Darwin, 1854b:419

Synonymy: Tarasov & Zevina, 1957:244.

Diagnosis: Cornwall, 1955b:43.

References: Barnard, 1924:94; Barnes & Klepal, 1971:88
(structure of the penis); Broch, 1924a:93; Cornwall,
1927a:507; 1955a:54; Gruvel, 1903b:152; 1905a:272;
Hiro, 1936e:318; Kolosvary. 1942a:140; 1942c:141;
1943a:99; 1967b:393; Kruger. 1927a:15; 1927b:5; New-
man & Ross, 1971:178; Nilsson-Cantell, 1926:15; 1939b:
238; 1957:8; Petriconi, 1969:539 (comparison of mouth
parts); Pilsbry, 1916:275; Stephensen, 1938:7; Weltner.
1897:254; 1898b:ll; Wolff, 1960:8; ZuUo, 1963b:14.

Distribution: Atlantic and Pacific Oceans; on Humpback
Whales.

Genus Cetopirus Ranzani, 1817

Cetopirus complanatus (Mbrch), 1852:67
Synonymy/diagnosis: Pilsbry, 1916:276.
References: Barnard, 1924:95; Broch, 1924b:204; Corn-
wall, 1953:83 (nervous system); Darwin, 1854b:415 (as

Coronula balaenaris); Gruvel, 1903b:152; 1905a:271
Guiler, 1956:3; Hiro, 1936e:318; Kolosvary, 1942a:141
1943a:100; Murray, 1895:449; Newman et al, 1969:R289:
Nilsson-Cantell, 1931a:116; 1938b:14; Stebbing, 1910:572
(as Coronula darwini); Stubbings, 1967:300; Tarasov &
Zevina, 1957:245; Weisbord, 1971:94; Weltner, 1897:
254; 1898b:8; 1900:307; ZuUo, 1961a:13.
Distribution: ChUe: Cape of Good Hope; AustraUa;
Tasmania; Kerguelen I., coast of Norway; Kei Is.

Genus Cetolepas ZuUo, 1969

Cetolepas hertleini ZuUo, 1969a: 17
Distribution: PUocene, San Diego.

Genus Cryptolepas DaU, 1872

Cryptolepas murata ZuUo, 1961a:14

Distribution: Pleistocene, San Quintin Bay, Baja
CaUfornia.
Cryptolepas rhachianecti DaU, 1872:300

Synonymy/diagnosis: Pilsbry, 1916:279.

References: Briggs & Morejohn, 1972:287; ComwaU,
1955a:49 (soft parts); 1955b:44; Gruvel, 1903b:153;
1905a:274; Hiro, 1935c:227; 1936a:62 (commensaUsm);
1936e:318; Hoek, 1883:7; Kasuya & Rice, 1970:42 (orien-
tation on whales); Kolosvary, 1943a:101; Korschelt,
1933:21; Scammon, 1874:22; Tarasov & Zevina, 1957:
246; Weltner, 1897:278; ZuUo, 1961a:13.

Distribution: Bering Sea to Lower CaUfornia; Korea;
Hawaiian Is.; on Grey whales.

Genus Tubicinella Lamarck, 1802

Tubicinella major Lamarck, 1802:463

Synonymy: Nilsson-CanteU, 1921:373 (includes pre-
Darwinian authors).

Diagnosis Darwin, 1854b:431.

References: Barnard, 1924:95 (as Tubicinella striata
Lamarck); Gruvel, 1903b:148 (as T. trachealis Darwin);
1905a:278; 1909a:225: Hiro, 1936a:62 (commensaUsm);
1936e:318; Hutton, 1879:330 (as T. trachealis): Kolosvary,
1943a: 100; Marloth. 1902:1 (mode of growth); Mbrch,
1852:66; Nilsson-CanteU, 1931a:116; 1957:8; Pilsbry,
1916:281; Stebbing, 1902:62 (as T. trachealis); 1910:573
(as T. striata); Weltner, 1897:253 (as T. trachealis);
1898b:7; 1900:307; ZuUo. 1963b:15.

Distribution: Southern Atlantic and Pacific Oceans; on
Southern Right Whales.

Genus Xeno6a/anus Steenstrup, 1851

Xenobalanus globicipitis Steenstrup, 1851:pl. 3, figs. 11-15;
1852:64

Synonymy/diagnosis: CornwaU, 1927a:510; Stubbings,
1965:902 (mouthparts).

References: Barnard, 1924:96; Barnes & Klepal, 1971:88
(pedicel of penis); Bassindale, 1964:43; Broch, 1924a:95;
Caiman, 1920:165; ComwaU, 1955a:56; 1955b:46; Darwin,
1854b:440; DoUfus, 1968:55; Gruvel, 1903b:159; 1905a:
280; 1920:55; Heldt, 1950:25; Hiro, 1936a:62 (com-
mensaUsm); 1936e:318: Hoek, 1909:271; Kolosvary,
1943a:100; Kruger, 1911a:59; Newman & Ross, 1971:
180; Nilsson-CanteU, 1921:375; 1930c:258; PiUerai, 1970:
248; Pilsbry, 1916:283; Pope, 1958:159; Richard, 1936:
55; Richard & NeuviUe, 1897:108; Stebbing, 1923:12 (as
X. natalensis); Stubbings, 1967:301; Tarasov & Zevina,
1957:250; Weltner, 1897:253; 1898b:ll; ZuUo, 1963b:15.

Distribution: World-wide; on porpoise, dolphin, and
Black Fish.

Family Bathylasmatidae Newman and Ross, 1971
Subfamily Bathylasmatinae n. status

Genus Bathylasma Newman and Ross, 1971

Bathylasma aucklandicum (Hector), 1887:440
Synonymy/diagnosis: Newman & Ross, 1971:151.
References: Benham, 1903:111; Clarke, 1905:419; Park,
1910:113; Utinomi, 1965:11; Withers, 1913:841; 1924:18;

46

1953:357.

Distribution: Miocene, New Zealand.
Batkylasma corolliforme (Hoek), 1883:155

Synonymy/diagnosis Newman & Ross, 1971:143.

References Bage, 1938:1; Borradaile, 1916:132 (as
Hexelasma antarcticum n. sp.|; Gruvel, 1903b:143;
1905a:255: Hoek, 1913:245; Kruger, 1911a:55 (see
Aaptolasma callistoderma); 1911b:460; Murray, 1895:421,
456; Nilsson-Cantell, 1930c:252; Pilsbry, 1916:330; South-
ward & Southward, 1958:635; Speden. 1962:746; Weis-
bord, 1965:1015 (in part); 1967:51 (in part); Weltner,
1897:271; 1900:305; Withers, 1924:22; Utinomi, 1965:13;
Zevina, 1968:93.

Distribution: Circum-Antarctic; to 1464m. Pleistocene,
to 70m above sea level.
Batkylasma hirsutum (Hoek). 1883:158

Synonymy/diagnosis: Newman & Ross, 1971:149.

References Crisp & Southward, 1961:271 (cirral activity);
Gruvel, 1903b:143; 1905a:256; 1920:55; Hoek, 1912:408;
1913:245; Jeffreys, 1878:414; Murray, 1895:456; NUsson-
Cantell, 1930c:252 (footnote 1); Pilsbry, 1916:330; South-
ward, 1957:323 (cirral activity); Southward & Southward,
1958:635; Utinomi, 1965:11; Weltner, 1897:271; 1898b:12.

Distribution: Northeast AtlEintic from Faeroe Is. south
to Azores; 944-1829m.

Genus Tessarelasma Withers, 1936

Tessarelasma pilsbryi Withers, 1936:591
Reference: Newman & Ross, 1971:155.
Distribution: Miocene, Pakistan.

Genus Tetrachaetasma Newman and Ross, 1971

Tetrachaetasma southwardi Newman & Ross, 1971:152
Synonymy/diagnosis: Newman & Ross, 1971:152.
References Borradaile, 1916:132 (in part); Weisbord,

1965:1015 (in parti; 1967:51 (in part).
Distribution: Antarctic Basin and off S. America; 1190-

2328m.

Subfamily Hexelasminae n. subfam.

Genus Hexelasma Hoek, 1913

Hexelasma arafurae Hoek, 1913:251
Synonymy/diagnosis Hoek, 1913:251.
References: Newman & Ross, 1971:155; Utinomi, 1965:

11.
Distribution: Arafura Sea; 560m.
Hexelasma fosteri Newman & Ross, 1971:155

Distribution: New Zealand; 538-676m.
Hexelasma velutinum Hoek, 1913:246
Synonymy: Utinomi, 1968a:30.
Diagnosis Hoek, 1913:246.
References Broch, 1931:53 {see Aaptolasma leptoderma);

Hiro, 1933:70; Newman & Ross, 1971:155; Withers,

1913:847.
Distribution: Japan; Phihppines to South China Sea;

204-390m.

Genus Aaptolasma Newman and Ross, 1971

Aaptolasma americanum (Pilsbry), 1916:330

Synonymy/diagnosis Newman & Ross, 1971:161.

Reference: Utinomi, 1965:12.

Distribution: Blake plateau, off Florida; 734-770m.
Aaptolasma brintoni Newman & Ross, 1971:162

Synonymy/diagnosis Newman & Ross, 1971:162.

Distribution: Off DaNang, Vietnam; 110-198m.
Aaptolasma callistoderma (Pilsbry), 1911:78

Synonymy/diagnosis Newman & Ross, 1971:159.

References Hoek, 1913:245; Kruger, 1911a:55 (as Balanus
coralliformis): 1911b:460; Pilsbry, 1916:332; Utinomi,
1958a:307; 1965:12.

Distribution: Japan; 115-141m.
Aaptolasma leptoderma Newman & Ross, 1971:165

Synonymy/diagnosis Newman & Ross, 1971:165.
References Broch, 1931:53 (as Hexelasma velutinum, in

part).
Distribution: Kei Is.; 290m.
Aaptolasma triderma Newman & Ross, 1971:164
Synonymy diagnosis Newman & Ross, 1971:164.
Distribution: Japan; 549m.

Family Tetraclitidae Gruvel, 1903
Subfamily Austrobalaninae n. subfam.

Genus Austrobalanus Pilsbry, 1916

Austrobalanus imperator (Darwin), 1854b:288
Synonymy/diagnosis Pope, 1945:364; Ross, 1971b:266 (as

Balanus (Austrobalanus) imperator).
References Barnes & Klepal, 1971:86 (pedicel of penis);
Dakin et al, 1948:176; Davadie, 1963:78; Endean et al,
1956:88 (ecology and distribution); Gruvel, 1905a:246;
Kolosvary, 1942c:140; 1943a:92; Kruger, 1940:466; Pils-
bry, 1916:219; Pope, 1959:117; Weltner, 1897:271;
Wisely & BUck, 1964:163 (nauphi).
Distribution: Australia.

Genus Epopella Ross 1970

Epopella breviscutum (Broch). 1922:337

Synonymy/diagnosis Ross, 1970:3.

Reference: Hiro, 1939e:275.

Distribution: Auckland Is.
Epopella plicatus (Gray), 1843:269

Synonymy/diagnosis Moore, 1944:326 (includes Elminius
rugosus Hutton 1879:328); Ross, 1970:9 (ex Elminius).

References Barnes & Klepal, 1971:87 (pedicel of penis);
Broch, 1922:341; Darwin, 1854b:351; Filhol, 1885:489;
Foster, 1967b:35 (larval stages); Gruvel, 1903b:163;
1905a:296, 297; 1906a:270; 1907d:l; 1909b:26; Jennings,
1918:62; Kolosvary, 1942c:140; Kruger, 1940:470;
Luckens, 1970c:497 (breeding and growth); Nilsson-
CanteU, 1930d:211; Pilsbry, 1916:261; Ritz & Foster,
1968:552 (temperature); Weltner, 1897:256; 1899a:443;
1900:307.

Distribution: Australia; New Zealand; Chatham, Snares
and Auckland Is.
Epopella simplex (Darwin), 1854b:353

Synonymy/diagnosis Pope, 1945:370; Ross, 1970:9 (ex
Elminius).

References Barnes & Klepal, 1971c:87 (structure of the
penis); Broch, 1922:342; Dakin et al, 1948:176; Gruvel,
1903b:163; 1905a:297; 1912a:350; 1909b:6; Guiler, 1952:
20; Kolosvary, 1942c:140; Kruger, 1914:429; 1940:470;
Linzey, 1942a:280; Moore. 1944:333; Nilsson-Cantell,
1938b:13; Pope, 1966:181; Weltner, 1897:256; 1900:307.

Distribution: Australia; Tasmania; Kermadec Is.

Subfamily Tetraclitellinae n. subfam.
Genus Tetraclitella Hiro, 1939

Tetraclitella chinensis (Nilsson-Cantell), 1921:359
Synonymy: Utinomi, 1970:347.
Diagnosis Nilsson-Cantell, 1921:359.

References Hiro, 1931:155 (as Tetraclita purpurascens
nipponensis n. subsp.); 1932b:473; 1937c:469; 1939e:273;
Ross, 1971a:217, 223; Utinomi, 1949a:36; 1954:23; Uti-
nomi and Kikuchi, 1966:8; Zevina & Tarasov, 1963:97.
Distribution: Southern Japan; China; Formosa.
Tetraclitella costata (Darwin), 1854b:339
Synonymy/diagnosis Darwin, 1854b:339; Nilsson-Cantell,

1930b:19.
References Gordon, 1970:98; Gruvel, 1905a:287; Hiro,
1916:259; 1928:316; Ross, 1971a:217, 233; Weltner,
1898:257.
Distribution: Phihppines; Indonesia.
Tetraclitella costata digita Resell, 1975:97
Distribution: Phihppines.

47

Tetraclitella darwini (Pilsbry), 1928:314
Synonymy: Utinomi. 1970:348.
Diagnosis Pilsbry. 1928:314.

References Hiro, 1937c:469: 1939e:277; 1939f:214; Kolo-
svary, 1943a:98; Nilsson-Cantell, 1931a:115; Ross. 1971a:
217. 223; Utinomi, 1949a:24; 1958a:304; 1962:237; 1969b:
53; Utinomi & Kikuchi, 1966:8.
Distribution: Japan; Formosa.
Tetraclitella divisa (Nilsson-Cantell), 1921:362
Synonymy: Ross. 1968:13.
Diagnosis Stubbings. 1967:291.

References Bassindale, 1961:485; Edmondson. 1933:231
(as T. purpurascens); Foster, 1974:45; Hiro, 1939e:275;
Pilsbry. 1928:316; Ross. 1961:210; 1968:13. (as T. d. sub-
quadrata n. subsp.); 1971a:217, 223; Utinomi. 1949a:25;
Zevina & Tarasov. 1963:96.
Distribution: Caribbean; West Africa; Sumatra; Formosa;
South China Sea; Pacific Oceania to Hawaii and Pitcaim.
Tetraclitella hyastina Rosell, 1974:7

Distribution: Mindanao. Philippines.
Tetraclitella karandei Ross, 1971a:217
References Ross, 1972:307; Karande, 1974a:249 (larvae).
Distribution: Bombay coast, India; Taiwan
Tetraclitella multicostata (Nilsson-Cantell), 1930a:2.
Synonymy/diagnosis Utinomi. 1962:231.
References Foster. 1974:46; Nilsson-Cantell, 1930b:18-

Ross, 1971a:217. 223.
Distribution: New Guinea; Fiji; Japan.
Tetraclitella pilsbryi (Utinomi), 1962:234
Synonymy/diagnosis Utinomi, 1962:234.
References Ross, 1971a:217, 223. Utinomi & Kikuchi

1966:8; 1970:348.
Distribution: Southern Japan.
Tetraclitella purpurascens (Wood). 1815:55
Synonymy: Nilsson-Cantell. 1921:358.
Diagnosis Pope. 1945:367.

References Anderson, 1969:183 (embryology and phyto-
geny); Barnes & Klepal. 1971:87 (pedicel of penis); Bhatt
and Bal. 1960:440; Broch. 1931:117; Dakin et al, 1948:176
Daniel. 1955c:30; Darwin, l854b:337; Dawydoff. 1952
128; Endean et al, 1956:88 (ecology and distribution)
Filhol, 1885:488; Foster, 1967a:83; 1967b:35 (larvae):
Gauld, 1957:10; Gordon, 1970:101; Gruvel. 1905a:285
Guiler. 1952:20; Button, 1879:328; Jennings, 1918:61
Karande. 1967:1245; 1966:147; Kolosvary. 1941a:42 (as
Tetraclita squamosa depressa); 1942c:140 (as Tetraclita
purpurascens darwini): 1943a:97 (var. darwini); 1941e:ll
(as Tetraclita radiata wagneri); Kruger. 1911a:4; Linzey.
1942a:279; Luckens. 1970c:497 (distribution and growth)!
Moore, 1944:333; NUsson-CanteU, 1931a:115; 1938b:13
Ritz & Foster, 1968:552 (temperature response); Ross
1971a:217. 223; Stubbings. 1967:293; Weltner, 1897:258
1899a:443; 1900:307; Wisely & BUck. 1964:163 (naupUi)
Distribution: Australia; New Zealand; Indonesia; India
Madagascar; East Africa.

Genus Newmanella Ross, 1969

Newmanella radiata (Bruguiere), 1789:168

Synonymy/diagnosis Ross, 1969:242

References de Blainville, 1824:378; 1825:598; 1827:plate
85; Bruguiere, 1791:plate 164; Chemnitz, in Martini &
Chemnitz, 1785:343; Darwin, 1854b:343; Deshayes, 1831
357; Gmelin. 1791:3213; Gruvel, 1903b:161; 1905a:291
Hoek. 1913:xvi; Jay, 1839:7; Kolosvary. 1943a:97:
Lamarck. 1818:393; Lamy & Andre, 1932:218; NUsson-
CanteU, 1931a:115; 1939a:5; PUsbry, 1916:259; 1927:38;
1953:27; Pope, 1945:368; Ranzani, 1818:75; 1820:39;
Ross. 1968:18; Southward. 1962:163 (behavior of cirrus
IV); Southward. 1975:17; Sowerby. 1823: (no pagination);
Spengler, 1790:172; Weltner. 1897:258.

Distribution: Florida through Caribbean to Venezuela.

Subfamily Tetraclitinae Gruvel, 1903

Genus Tesseropora Pilsbry. 1916
Tesseropora isseli (de Alessandri). 1895:296

Distribution: Oligocene. Italy (Withers. 1953:59).
Tesseropora pacifica (Pilsbry), 1928:312
Synonymy/diagnosis Pilsbry. 1928:312 (as T wireni

pacifica); Henry, 1957:33.
References Foster. 1974:44; Gordon. 1970:103; Hiro,
1936a:59 (commensalism); Kolosvary. 1962cl93- 1967b'
393; Ross. 1969:239.
Distribution: Insular. Indo-West Pacific (Fiji to Hawaii).
Tesseropora rosea (Krauss), 1848:136
Synonymy: Pilsbry, 1916:260.
Diagnosis Pope, 1945:366.

References Anderson. 1969:183 (embryology/phylogenv)-
Barnard. 1924:92; Dakin et al, 1948:176; Darwin 1854b'
335; Endean et al. 1956:88 (ecology and distribution)-
Gruvel. 1903b:161; 1905a:286; Hoek, 1883:161; Kolosvary
1943a:98; Linzey. 1942a:280; Moore, 1944:333; Nilsson-
CanteU, 1927a:786; 1938b:14; Stubbing, 1910-571- Wisely
& Blick, 1964:163 (nauplii); Weltner, 1897:258.
Distribution: Australia; Kermadec I.; South Africa.
Tesseropora wireni (Nilsson-Cantell), 1921:366
Synonymy/diagnosis Hiro, 1937b:68.
Reference: Hiro, 1936a:59 (commensahsm).
Distribution: Sumatra; Palau Is.
Tesseropora wireni africana (Nilsson-Cantell), 1932b:14
Synonymy/diagnosis Nilsson-Cantell. 1932b:14
References NUsson-CanteU. 1938b:14; Smith, 1971103
Distribution: Dar-es-Salaam and Diego Garcia, Indian
Ocean.

Genus Tesseroplax Ross, 1969

Tesseroplax unisemita (Zullo). 1968d:272
Reference: Ross. 1969:237.

Distribution: PUocene, Isla Angel de la Guardia, Gulf of
California.

Genus Tetraclita Schumacher, 1817
Tetraclita alba Nilsson-Cantell, 1932b: 11
Reference: Nilsson-Cantell, 1938b:13.
Distribution: Dar-es-Salaam.
Tetraclita coerulescens (Spengler), 1790:191
Synonymy: Nilsson-Cantell. 1938b:77.
Diagnosis Broch. 1931:116.

References Darwin, 1854b:342; Endean et al, 1956-88-
Gruvel, 1905a:290; Hiro, 1936b:635; 1937b:67- 1939c'
586; Hoek, 1883:161; 1913:257; PUsbry, 1916:259; Rosell,
1972:211; Stephenson et al, 1958 (insular ecology)'
Weltner, 1897:257.
Distribution: Philippines; Palau Is.; Sulu Arch.; Indo-
nesia; Bay of Bengal; Mergui Arch.; Australia.
Tetraclita dumortieri Fischer. 1865:434
Reference: de Alessandri. 1907:290.
Distribution: Miocene. France.
Tetraclita hentscheli Kolosvary, 1942c:141
Reference: Kolosvdry, 1943a:98.
Distribution: Puerto Cabello, Venezuela.
Tetraclita serrata Darwin, 1854b:334
Synonymy: Barnard, 1924:91.
Diagnosis Darwin. 1854b:334.

References: Annandale, 1906:144; Barnes & Barnes.
1965a:392 (variation in size); Day & Morgans. 1956-27d
(ecology); Gruvel, 1903b:161; 1905a:289; Hoek. 1913-254-
Millard, 1950:270; NUsson-Cantell. 1938b:13; Pichon,
1972:381; Ritz & Foster. 1968:545 (temperature re^
spouses); RoseU. 1972:208; Sandison, 1954:96 (nauplii)-
Stebbing, 1910:570; Weltner, 1897:258. '

Distribution: South Africa; Madagascar; Ceylon; Philip-
pines.

48

Tetraclita squamosa squamosa (Brugui^re), 1789:170

Synonymy: Henry. 1957:33.

Diagnosis Pilsbry, 1916:251; Kniger. 1911a:61.

References Barnard, 1924:90: Borradaile, 1900:799;
Broch. 1916:14: 1922:337; 1924b:204: 1931:116; 1947:7;
Crisp & Southward, 1961:272 (cirral activity); Darwin,
1854b:329 (as Tetraclita porosa var. viridis): Dawydoff,
1952:128; de Oliveira, 1941:6 (probably T. stalactifera);
Endean et al, 1956:88 (mainland ecology and distribu-
tion); 1956:317 (insular ecology and distribution); Foster,
1974:45 (as T. squamosa viridis): Gruvel. 1896a:43
(branchiae): 1896b:205 (anatomy); 1896e:186 (review);
1905a:288; 1909a:216, 225: 1909b:25; Hire, 1936b:635;
1937b:66; 1937c:467; 1939c:586; 1939e:271; Hoek, 1913:
254; Kolosvary. 1943a:96; 1951c:412; Kruger. 1911b:461;
1914:441; 1940:472; Moore. 1944:333; Morton. 1973:491;
Nilsson-CanteU. 1921:364; 1930b:17; 1931a:115; 1934a:
71; 1934b:61; 1938b:76; Nomura. 1938:87; Ooishi. 1964:
195; Pilsbry, 1916:249; Rosell, 1972:205; 1973b:94; Speng-
ler, 1790:192 (? Lepas mitra); Stebbing. 1910:570 [porosa
Gmelin. 1790 = squamosa Brugiere. 1789); Stephenson
et al. 1958 (insular ecology); Stubbings. 1967:284; Utinomi,
1942:10; 1949a:25; 1954:23; 1958a:304; 1968b:178; 1969b:
53; Weltner. 1895:289 (probably T. stalactifera): 1897:257;
1910:528; Zevina & Litvinova. 1970:174; Zevina &
Tarasov, 1963:95.

Distribution: Japan; Formosa; Philippines; Palau Is.;
Indonesia; Australia; Mergui Arch.; Andamans; Great
Nicobar I.; Red Sea; Rio de Janeiro; Cape Palmas,
W. Africa. Pliocene: Ryukyu I.
Tetraclita squamosa formosana Hire, 1939e:271

Synonymy/diagnosis Hiro, 1939e:271.

References Ooishi, 1964:195; Utinomi, 1949a:23; 1954:23;
1969b:53.

Distribution: Southern Japan; Formosa.
Tetraclita squamosa japonica Pilsbry. 1916:252

Synonymy: Hiro. 1932a:551; 1937c:469.

Diagnosis Pilsbry, 1916:252.

References Hiro, 1932b:473; 1938c:1687 (resistance to
exposure); 19391:213; Ikenouye, 1968:99 (spatial distribu-
tion); Kolosvary, 1943a:96; Kruger, 1911a:61 (as Tetraclita
porosa var. nigrescens): 1940:472; Mori, 1958:23 (rhyth-
mic activity); 1961:373 (rhythmic activity); Nilsson-
CanteU, 1927a:786; 1931a:115: 1932a:27; Pilsbry, 1911:
81 (as T porosa): Suzuki & Mori, 1963:1 (water content);
Tarasov & Zevina, 1957:236: Utinomi, 1949a:23; 1958a:
304; 1958b:51; 1969b:53; 1970:347; Utinomi & Kikuchi,
1966:6; Weltner, 1897:257 (as var. nigrescens): Zevina
& Tarasov. 1963:95.

Distribution: Japan and Korea.
Tetraclita squamosa milleporosa Pilsbry. 1916:257

Synonymy/diagnosis Pilsbry, 1916:257

References Hedgpeth, 1969:9 (as Tetraclita stalactifera
milleporosa): Zullo, 1966c:143.

Distribution: Galapagos Is.
Tetraclita squamosa panamensis Pilsbry. 1916:256

Synonymy/diagnosis Pilsbry. 1916:256.

References Hedgpeth. 1969:17; Kolosvary, 1943:97;
Nilsson-CanteU, 1957:10; Pilsbry, 1909:64; Zevina &
Kurshakova, 1973:183.

Distribution: Panama; Ecuador; Peru; Galapagos Is.
Tetraclita squamosa patellaris Darwin, 1854b:330

Synonymy: Pilsbry, 1916:248.

Diagnosis Darwin, 1854b:330.

References Gruvel, 1903b:161; 1905a:288; 1907d:8;
1909b:25; Nilsson-CanteU, 1938b:13; Weltner, 1897:258.

Distribution: Andaman Is.
Tetraclita squamosa perfecta Nilsson-CanteU, 1931a:133

Di.stribution: Santuao, China.
Tetraclita squamosa rubescens Darwin, 1854b:329

Synonymy: Ross. 1962:34.

Diagnosis CornwaU, 1951:312.

References Barnes, 1959a:233 (stomach contents);
Barnes & Barnes, 1959h:515 (metaboUsm); 1965a:392

(egg size); Barnes & Klepal, 1971:86 (pedicel of penis);
Broch, 1922:337; Darwin, 1854b:330 (as T. s. rubescens
forma elegans nov.); Emerson. 1956:339; Gruvel, 1903b:
161; 1905a:288; 1909b:25 (including forma elegans):
Henry, 1942:122, 123 (as elegans): 1960:147; Hewatt,
1935:250; 1946:199; Hoek, 1913:254; Kolosvary, 1943a:
96; Kruger, 1940:472; PUsbry, 1916:257, 258 (as elegans):
Rasmussen, in Shelford et al, 1935:307; Shimkin et al,
1951:650 (carcinogenic substances); Weltner, 1897:257,
258 (as T porosa var. 5, elegans): WiUett, 1937:383;
ZuUo, 1966c:141 (as T. squamosa elegans):.

Distribution: Central California to Cape San Lucas, Baja
California. Pleistocene: Los Angeles, CaUfornia and
Punta China, Baja California.
Tetraclita squamosa rufotincta PUsbry, 1916:253

Synonymy: Utinomi, 1968b:180.

Diagnosis Pilsbry. 1916:253.

References Achituv, 1972:73 (zonation); Barnes & Klepal,
1971c:86 (pedicel of penis): Fishelson, 1971:123 (ecology
and distribution in Red Sea); Kolosvary. 1943a:97
Kruger, 1940:472; Nilsson-CanteU, 1921:365; 1928a:35
1938b:13; Utinomi, 1969a:82; Zevina & Litvinova, 1970:
174.

Distribution: Red Sea; East Africa; Arabian coast, west
coast of India, and islands of the western Indian Ocean.
Tetraclita stalactifera (Lamarck), 1818:394

Synonymy/diagnosis Ross, 1968:8.

References Barnes & Klepal, 1971:86 (pedicel of penis);
Bigelow, 1902:180; Chenu, 1843:(no pagination); Cornwall,
in Steinbeck & Ricketts, 1941:430; Darwin, 1854b:329
(as Tetraclita porosa var. 1, communis and var. 2,
nigrescens): de OUveira, 1940a:138; 1941:7; Gruvel
1903b:161; 1905a:287; Henry, 1941:105; 1942:127; 1943:
367; 1954:444; 1958:224; 1960:147; Kolosvary, 1943a:97
Kruger, 1911a:61 (Tetraclita squamosa japonica accord-
ing to Pilsbry); 1911b:461 (T. s. japonica): 1940:472:
Lacombe & Monteiro, 1974:633; Lamy & Andre, 1932
222; Morch, 1852:67; NeweU et al, 1959:209; NUsson-
CanteU, 1933:508; 1939a;5; Pilsbry, 1916:254; 1927:38;
Ross, 1962:32; Smith et al, 1950:134; Stephensen &
Stephensen, 1950:388; 1952:8; Stubbings, 1936:49;
Utinomi, 1968b:179; VerrUl, 1901:22; Voss & Voss, 1960:
102;ZuUo, 1966c:141.

Distribution: Bermuda; S.E. United States; Gulf of
Mexico; West Indies to southern BrazU; Gulf of CaUfor-
nia to Acapulco, Mexico. Other locaUties: Arabian Sea
(reported by Stubbings as Tetraclita porosa var. com-
munis): Cape Province, S. Africa (reported by Utinomi
as Tetraclita squamosa stalactifera). PUo-Pleistocene:
Curacao. Pleistocene: Venezuela.
Tetraclita stalactifera confinis PUsbry, 1916:255

Synonymy: Ross, 1962:34.

Diagnosis Pilsbry, 1916:255.

References Barnes & Klepal, 1971:86 (pedicel of penis);
Henry, 1941:105; 1943:369; 1960:143; Hertlein &
Emerson, 1956:167.

Distribution: Gulf of CaUfornia. Pleistocene: Sonera,
Mexico.
Tetraclita stalactifera floridana PUsbry, 1916:255

References Barnes & Klepal, 1971:86 (pedicel of penis).

Distribution: Lake Worth Inlet, Florida.
Tetraclita vitiata Darwin, 1854b:340

Synonymy: Nilsson-CanteU, 1938b:76.

Diagnosis Broch, 1922:339

References Endean et al, 1956:88 (mainland ecology and
distribution); 1956:317 (insular ecology and distribution);
Gruvel, 1903b:161; 1905a:289; Hiro, 1936b:635; 1937b:
67; 1939c:586; Hoek, 1913:256; Pilsbry, 1916:259; RoseU,
1972:214; Stephenson, 1968:51; Stephenson et al, 1958:261
(insular ecology); Weltner, 1897:258.

Distribution: Philippines; Sulu Arch.; Indonesia; Great
Barrier Reef, Australia; Nicobar I.

49

Superfamily Balanoidea Leach, 1817 n. status

Family Archaeobalanidae n. fam.

Subfamily Archaeobalaninae n. subfam.

Genus Archaeobatanus Menesini, 1971

Archaeobalanus semicanaliculatus Menesini, 1971:28
Reference: Plaziat & Cavalier, 1973:2875.
Distribution: Eocene and OUgocene, Paris Basin.

Genus Actinobalanus Moroni, 1967

Actinobalanus actinomorphus (Moroni). 1952:73
Synonymy: Moroni, 1967:923.
Diagnosis: Moroni, 1952:73.
Distribution: Pliocene, Italy.
Actinobalanus bisculcatus (Darwin), 1854a:26
Synonymy': Ross & Newman, 1967:6.
Diagnosis: Darwin, 1854b:293.
References: Davadie, 1967:74; de Alessandri. 1907b:286;

Moroni, 1967:925; Withers, 1953:58, 62.
Distribution: Coralline Crag, England; Eocene- Pliocene,
Northern Europe.
Actinobalanus bisulcatus plicatus (Darwin), 1854a:26
Synonymy': Ross & Newman, 1967:6.
Reference: Darwin, 1854b:293; Davadie, 1963:74.
Distribution: Coralline Crag, England: Belgium.
Actinobalanus pantanelli (de Alessandri), 1895:293
Actinobalanus dolosus (Darwin), 1954a:28
Synonymy: Moroni, 1967:925.
Diagnosis: Darwin, 1854b:295; Davadie, 1963:76; Kolo-

svary, 1967b:391; Lecointre. 1910:138.
Distribution: Miocene, France; Pliocene, England and
Norway.
Actinobalanus inclusus (Darwin), 1854a:31
Synonymy: Moroni, 1967:925; Davadie. 1963:75; Kolo-

svary, 1967b:391.
Diagnosis: Darwin. 1854b:299.

Distribution: Miocene to Pleistocene, Northern Europe.
Distribution: Pliocene, Italy.
Actinobalanus stellaris (Brocchi), 1814:599
Synonymy: de Alessandri, 1906:302 (= B. corrugatus
Darwin, 1854b:254); Davadie, 1963:66 {B. s. var.
miocenicus Seguenza, 1876:453).
Distribution: OUgocene-Pliocene, Italy.

Genus Kathpalmeria Ross, 1965

Kathpalmeria georgiana Ross, in Ross and Newman,
1965a:61
Distribution: Eocene, Georgia.
Kathpalmeria hantkeni (Kolosvary), 1947b:305
Synonymy: Ross, 1965a:62.
Diagnosis: Kolosvary, 1947b:305.
Reference: Plaziat & Cavelier, 1973:2875 (as Balanus

(Austrobalanus) hantkeni).
Distribution: Eocene, Hungary.

Genus Armatobalanus Hoek, 1913

Armato balanus (Armatobalanus) allium (Darwin), 1854b:281

Synonymy: Zullo, 1963d:588.

Diagnosis; Utinomi, 1949a:30.

References: Annandale, 1906:148; 1924:62 (as Balanus
arcuatus); Broch, 1922:325, 333 (as Acasta madreporicola
n. sp.); 1931:78 (as Balanus arcuatus): Gruvel, 1905a:247;
Hiro, 1936a:38 (as Acasta madreporicola); Hoek, 1913:
210 (as Balanus arcuatus n. sp.); Kolosvary, 1951a:
288; Nilsson-Cantell, 1921:337 (as Balanus arcuatus);
1938b:52 (as Balanus arcuatus); PUsbry, 1916:228;
Utinomi, 1949a:32 {Balanus arcuatus and Acasta mad-
reporicola, discussion); 1962:217; Utinomi & Kikuchi,
1966:6; Weltner, 1897:271; ZuUo, 1967b:126.

Distribution: Great Barrier Reef; Indonesia; Southwest
Japan; Ceylon; Bay of Bengal; 9-55m.

Armatobalanus (Armatobalanus) allium truncatus (Utinomi),

1949a:32
Synonymy/diagnosis: Utinomi, 1949a:32.
Reference: Zullo, 1963d:588.
Distribution: Tanabe Bay, Japan.
Armatobalanus (Armatobalanus) catvertensis (Ross), 1965:334

Distribution: Miocene, Maryland.
Armatobalanus (Armatobalanus) cepa (Darwin), 1854b:283
Synonymy: Nilsson-Cantell, 1938b:52.
Diagnosis: Nilsson-Cantell, 1938b:52.
References: Borradaile, 1903:442 (as Walanus terebratus);

Broch. 1931:79; Gruvel. 1905a:251; Hiro, 1936b:625 (as

Balanus fujiyama); Kolosvary, 1943a;93 (as Balanus

fujiyama); 1947e:358 (as Balanus fujiyamaformis n. sp.);

Nilsson-Cantell, 1932c:6; Pilsbry, 1916:228; Utinomi,

1949a:29; ZuUo, 1963d:589.
Distribution: Indonesia; Australia; Mergui Arch.; Mal-
dives; southwest Japan; 50m.
Armatobalanus (Armatobalanus) circe (Kolosvary), 1947e:359
Synonymy/diagnosis: Kolosvary, 1947e:359.
References: Kolosvary, 1951a:288; Ross, 1965b:332;

ZuUo, 1963d:589.
Distribution: West Indies.
Armatobalanus (Armatobalanus) duvergieri (de Alessandri),

1922:223
Diagnosis/references: Withers, 1929a:562; 1953:57; 58;

ZuUo, 1961b:71.
Distribution: Miocene, France; on Porites incrustans.
Armatobalanus (Armatobalanus) filigranus (Broch) 1916:8
Synonymy/diagnosis: Broch, 1916:8.
References: Hiro, 1937b:56; ZuUo, 1963d:(errata).
Distribution: W. Australia; Palau Is.; 4-20m.
Armatobalanus (Armatobalanus) funiculorum (Annandale),

1906:145
Synonymy/diagnosis: Annandale, 1906:145.
References: Kolosvary, 1951b:229; ZuUo, 1963d:589.
Distribution: Gulf of Manaar.
Armatobalanus (Armatobalanus) nefrens (ZuUo), 1963d:590
Synonymy/diagnosis; ZuUo, 1963d:590.
Reference: Ross, 1965b:332.
Distribution: Monterey and Carmel Bays and Channel

Is., CaUfornia.
Armatobalanus (Armatobalanus) oryza Broch, 1931:82
Synonymy/diagnosis; Broch, 1931:82.
Reference: ZuUo, 1963d; 589.
Distribution: Banda Sea; 200m.
Armatobalanus (Armatobalanus) palaoensis Hiro, 1937b:60
Synonymy/diagnosis; Hiro, 1937b:60.
Reference: ZuUo, 1963d:589.
Distribution: Palau Is.
Armatobalanus (Armatobalanus) quadrivittatus Darwin,

1854b:284
Synonymy: ZuUo, 1963d:589.
Diagnosis: Hoek, 1913:213.
References: BorradaUe, 1903:442; Broch, 1947:8; Davadie,

1952:30; Dawydoff, 1952:128; Gruvel, 1903b:141; 1905a:

248; Hoek, 1907:xvi; Kolosvary, 1947d:425; 1951b:292;

Nilsson-CanteU, 1921:339; 1934b:60; 1938b:54; Pilsbry,

1916:229; Utinomi, 1962:217; Utinomi & Kikuchi, 1966:

6; Weltner, 1897:271.
Distribution: Maldives; Indonesia; Singapore; Viet Nam;

Mergui Arch.; Southwestern Japan; Philippines; 31-51m.

Miocene, Algeria.
Armatobalanus (Armatobalanus) quinquevittatus Hoek,

1913:216
Synonymy/diagnosis: Hoek, 1913:216.
Reference: ZuUo, 1963d:589.
Distribution: Off Ambon; 32m.
Armatobalanus (Armatobalanus) terebratus Darwin,

1854b:285
Synonymy: Nilsson-CanteU, 1938b:51.
Diagnosis: Hoek, 1913:207.
References: Annandale, 1906:148; Borradaile, 1903:442;

Broch, 1916:6; Dawydoff, 1952:128; Gruvel, 1905a:249;

50

Hiro, 1935a:l; 1937b:55; Korschelt, 1933:27; Weltner,
1897:271; Zullo, 1963d:590.

Distribution: Palau Is.; Kei Is.; western Australia;
Gulf of Siam; Madras, India; 0-55m.
Armatobalanus (Armatobalanus) terebratus radicifer (Annan-
dale), 1924:63

Synonymy/diagnosis Annandale, 1924:63.

References Hiro, 1937b:55; ZuUo, 1963d:590.

Distribution: Mergui Arch.

Subgenus Hexacreusia ZuUo, 1961

Armatobalanus (Hexacreusia) durhami (Zullo), 1961b:73
Synonymy/diagnosis: Zullo, 1961b:73.
References: Newman & Ladd, 1974:383; Ross, 1962:37;
Ross & Newman, 1973:148; ZuUo, 1967b:126; ZuUo &
Beach. 1973:13; ZuUo et al, 1972:72.
Distribution: Pliocene to Recent: Gulf of CaUfornia, on
Pontes.
Armatobalanus (Hexacreusia) straeleni (ZuUo & Beach),
1973:11
Distribution: Galapagos, on ahermatypic coral; 55-90m.

Genus Chirona Gray, 1835

Chirona (Chirona) bimanicus (Withers), 1923:288

Distribution: Miocene, Burma.
Chirona (Chirona) evermanni (Pilsbry), 1907d:203

Synonymy: Tarasov & Zevina, 1957:230 (includes B. (Meta-
balanus) hoekianus Pilsbry, 1911:77).

Diagnosis: Pilsbry, 1916:201 (as Balanus hoekianus);
Pilsbry, 1916:210.

References: Barnes & Klepal, 1971:85 (pedicel of penis);
Henry, 1942:126; Hiro, 1935c:227; Hoek, 1913:151, 246
[as Hexelasma hoekianum); Kruger, 1911b:460; Newman
& Ross, 1971:171; PUsbry, 1911:76 (evermanni). 11 (hoek-
ianus).

Distribution: Gulf of Alaska; Aleutians and Bering Sea;
140-490m.
Chirona (Chirona) hameri (Ascanius), 1767:8

Synonymy: Pilsbry, 1916:205.

Diagnosis: Pilsbry, 1916:205.

References: Barnes & Barnes, 1965a:391 (variation in egg
size); Barnes & Klepal, 1971:85 (pedicel of penis); Bas-
sindale, 1964:39; Bousfield. 1954:121; Broch, 1924a:88;
Crisp, 1962b: 123 (larval stages); Crisp & Southward,
1961:271 (cirral activity); Darwin, 1854a:24; 1854b:277;
Davadie, 1963:71; Foster, 1970:377 (acclimation to
saUnity); Gruvel, 1903b:141; 1905a:245; Hoek, 1875:60:
1909:270; Kolosvary, 1967b:392; Kruger, 1927a:14
1927b:5; 1940:464; Moore, 1935a:57 (growth rate)
Morch, 1852:68: O'Riordan, 1967:293; Poulsen, 1935:16:
Rzhepishevskii, 1968:36; Southward & Crisp, 1963:32
(fouUng); Stephensen, 1938:6; Tarasov, 1937:52; Walker,
1970:239 (cement apparatus); 1972:429 (cement composi-
tion); 1973b:455 (frontal horns and gland ceUs); Weltner,
1897:270; 1898a:443; 1898b:12; ZuUo, 1963b:13.

Distribution: North Atlantic; Chesapeake Bay; Barents
and North Seas; England; 29-305m. PUo-Pleistocene,
Northern Europe and North America.
Chirona (Chirona) sublaevis (Sowerby), 1840:327

References: Withers, 1923:285.

Distribution: Soomrow, India.
Chirona (Chirona) unguiformis (Sowerby), 1846:pl. 648

Synonymy: Darwin, 1854a:29.

References: Darwin, 1854b:296; Davadie, 1963:73 (thin
sections); de Alessandri, 1907b:288; Plaziat & CavaUer,
1973:2875 (paleo-ecology); Ross & Newman, 1967:4;
Withers, 1953:48 et seq.

Distribution: Eocene-OUgocene, England and Paris Basin.
Eocene, Southeastern U.S.
Chirona (Chirona) varians (Sowerby), 1846:pl. 2

References: Darwin. 1854b:298; 1897:622; Ortmann,
1902:250 (probably includes Chthamalus antiquus
(PhilUppi); Withers, 1953:141.

Distribution: Tertiary, Patagonia and Tierra del Fuego.

Subgenus Smafo6a/anus Hoek, 1913

Chirona (Striatobalanus) amaryllis (Darwin), 1854b:279

Synonymy diagnosis: Darwin, 1854b:279; Hoek, 1913:179.

References: Annandale, 1906:147: Barnes & Klepal,
1971:85 (pedicel of penis); Broch, 1916:6; 1922:321 (as
forma euamaryllis nov.); 1931:66, 67 (as forma laevis
nov.); 1947:5, 6; Daniel, 1955:25; Darwin, 1854b:279
|var. a (= Balanus roseus Lamarck, 1818); var. b.];
Dawydoff, 1952:128; Endean et al, 1956:88; Gruvel,
1903b:141; 1905a:250 (as var. niveus nov. = var. b.
Darwin, 1854b); Hiro. 1936b:624; 1939d:243; Hoek, 1883:
153; 1912:408; 1913:179: Karande. 1967:1245; Karande
& Palekar, 1966:147; Kruger, 1911a:4; 1911b:460; 1940:
464; Lanchester, 1902:369 (as Balanus amaryllis dis-
similis n. subsp.); Nilsson-CanteU, 1921:329; 1927a:785;
1930b:10; 1931a:114; 1934a:68; 1934b:58; 1938b:46;
Pope, 1945:364; Stubbings, 1936:41; 1961a:174; Utinomi.
1962:216; 1968b:174; 1969a:88: Utinomi & Kikuchi,
1966:6; Weltner, 1897:270.

Distribution: Indo-west Pacific: East Africa to PhiUppines
and Northeast AustraUa: 5-500m., and on ships.
Chirona (Striatobalanus) bimae (Hoek), 1913:182

Synonymy/diagnosis: Hoek, 1913:182.

References: Broch, 1931:70; Nilsson-CanteU, 1934b:58;
1938b:48.

Distribution: Java Sea; 12-35m.
Chirona (Striatobalanus) krugeri (PUsbry), 1916:214

Synonymy/diagnosis: PUsbry, 1916:214.

References: Broch, 1931:71; Hiro, 1933:72; 1937c:440;
1939b:56; Utinomi, 1949b:96: 1958a:308.

Distribution: Japan; Moluccas; 100-250m.
Chirona (Striatobalanus) maculatus (Hoek), 1913:187

Distribution: Java Sea.
Chirona (Striatobalanus) taiwanensis (Hiro), 1939e:264

Distribution: Formosa.
Chirona (Striatobalanus) tenuis (Hoek), 1883:154.

Synonymy: Hiro, 1937c:439.

Diagnosis: Hoek, 1913:185 (as Balanus albus n.sp.).

References: Barnard, 1924:74; Broch, 1931:70: Daniel,
1955:24; Gruvel, 1905a:247; Hoek, 1912:408; Nilsson-
CanteU, 1925:34; 1927a:785; 1938b:46; Pilsbry, 1916:
216; Stubbings, 1936:41 (as albus): 1940:390; Utinomi,
1950:63; 1962:216; 1968b:174; 1969a:88; Utinomi and
Kikuchi, 1966:6; Weltner, 1897:271.

Distribution: Indo-west pacific: South Africa, Persian
Gulf to PhiUppines and Japan: 7-500m.
Chirona (Striatobalanus) tuberculatus (RoseU), 1974:2

Distribution: Mindanao, PhiUppines.
Chirona (Striatobalanus) zealandicus (Withers), 1924:35

Distribution: Miocene, New Zealand (Withers, 1953:78
et seq.).

Genus Solidobalanus Hoek, 1913

Solidobalanus (Solidobalanus) astacophilus (Barnard),
1926:128

Synonymy/diagnosis; Barnard, 1926:128.

References: Henry & McLaughMn, 1967:47; ZuUo & New-
man, 1964:368.

Distribution: South Africa; 420m.
Solidobalanus (Solidobalanus) auricoma (Hoek), 1913:198

Synonymy: ZuUo & Newman, 1964:368.

Diagnosis: Hoek, 1913:198; Nilsson-CanteU, 1938b:49.

References: Broch, 1922:323; 1931:71; Henry &
McLaughlin, 1967:46; Nilsson-CanteU, 1934a:70;
Utinomi, 1969:82.

Distribution: Persian Gulf; Moluccas; southwest
AustraUa; 27-320m.
Solidobalanus (Solidobalanus) ciliatus (Hoek), 1913:199.

Synonymy: ZuUo & Newman, 1964:368.

Diagnosis: Hoek, 1913:199; Nilsson-CanteU, 1934a:68.

References; Annandale, 1906:148 (as Balanus maldiven-
sis Borradaile, 1903); Broch, 1931:72; 1947:6; Dawydoff,
1952:128; Henry &. McLaughUn, 1967:47; NUsson-
CanteU, 1925:38; 1934b:59; 1938b;49; Stubbings,
1936:43; Utinomi, 1969a:90.

51

Distribution: Indo-west Pacific: Gulfs of Aden, Persia

and Manaar; Red Sea; India; Indonesia: 13-220m.

Solidobalanus ISolidobalanus) compressus (Hoek), 1913:202

References: Henry & McLaughlin, 1967:47: ZuUo & New-
man, 1964:369.

Distribution: Banda Sea; 75-1 12m.
Solidobalanus ISolidobalanus) echinoplacis (Stubbingsl,
1936:45

References: Henry & McLaughlin, 1967:47; Zullo & New-
man, 1964:369.

Distribution: Zanzibar; 225m.
Solidobalanus (Solidobalanus) hawaiensis (Pilsbry), 1916:222

References: Gordon, 1970:81; Henry & McLaughlin,
1967:47 {hawaiiensis |sic)); Utinomi, 1949b:96; Zullo &
Newman, 1964:369.

Distribution: Hawaiian Is.; 21-222m.
Solidobalanus (Solidobalanus) maldivensis (Borradaile),
1903:442

Synonymy/diagnosis: Hoek, 1913:195.

References: Annandale, 1906:148; Henry & McLaughlin,
1967:47; Zullo & Newman, 1964:369.

Distribution: Maldives and Flores Sea; 69-390m.
Solidobalanus (Solidobalanus) masignotus (Henry &
McLaughlin), 1967:47

Reference: McLaughlin & Henry, 1972:14 (complemental
males).

Distribution: Central Baja Cahfornia; Mazatlan, Mexico;
Costa Rica; Ecuador; subUttoral.
Solidobalanus (Solidobalanus) mylensis (Seguenza), 1876:308

Synonymy: Moroni. 1967:5 {milensis of some authors);
Withers, 1953:61,62.

Distribution: Neogene, Italy.
Solidobalanus (Solidobalanus) nascanus (Zullo, in Zullo &
Newman), 1964:366

References: Henry & McLaughhn, 1967:47; Zevina &
Kurshakova, 1973:183.

Distribution: Eastern Pacific; Nasca Ridge; 228m.
Solidobalanus (Solidobalanus) pseudauricoma (Broch),
1931:72

Synonymy/diagnosis: Utinomi, 1949b:97.

References: Henry & McLaughlin, 1967:47; Zullo &
Newman, 1964:369.

Distribution: Celebes; Japan; 70-500m.
Solidobalanus (Solidobalanus) socialis (Hoek), 1883:150

Synonymy: Zullo & Newman, 1964:369.

Diagnosis: Hoek, 1883:150; 1913:192.

References: Annandale, 1906:148 (as Balanus aeneas);
Gruvel, 1905a:226,252 (as Balanus aeneas): Henry &
McLaughlin, 1967:46; Hiro, 1932b:473; 1937c:442; Hoek,
1913:150,154 (as Balanus aeneas); Lanchester, 1902:370
(as Balanus aeneas n.sp.); Stubbings, 1963a:334
Utinomi, 1949a:22; 1962:217; 1968b:175; 1969a:89:
1970:359; Utinomi & Kikuchi, 1966:6; Weltner, 1897
267.

Distribution: Indo-west Pacific: Gulfs of Persia and
Manaar; Bay of Bengal; Indonesia; Southeast coast of
Japan; to 91m.
Solidobalanus (Solidobalanus) solidus (Broch), 1931:76

References; Henry & McLaughlin, 1967:47; Zullo &
Newman, 1964:369.

Distribution: Japan; 300m.
Solidobalanus (Solidobalanus) tantillus (Pilsbry), 1916:224

References: Henry & McLaughlin, 1967:47; Zullo & New-
man, 1964:369.

Distribution: Sulu Arch.; 100m.
Solidobalanus (Solidobalanus) thompsoni (Stubbings),
1936:43

References: Henry & McLaughlin, 1967:47; Nilsson-
CanteU, 1938b:13; ZuUo & Newman, 1964:369.

Distribution: Gulf of Aden; 73-220m.

Subgenus Hesperibalanus Pilsbry, 1916

Solidobalanus (Hesperibalanus) comwalli (Zullo) 1966a:200
Distribution: Eocene, Washington.

Solidobalanus (Hesperibalanus) elizabetfiae (Barnard)
1924:72

Synonymy/diagnosis: Millard, 1950:267.

References: Nilsson-Cantell, 1932c:8 (as Balanus emk-
weniensis n.sp.); Henry & McLaughhn, 1967:47.

Distribution: South Africa.
Solidobalanus (Hesperibalanus) engbergi (Pilsbry), 1921:113

Synonymy: Cornwall, 1955b:31.

Diagnosis: Pilsbry, 1921:113.

References: Barnes & Klepal, 1971:84 (pedicel of penis);
Cornwall, 1955a:31; Henry, 1940:33; 1942:107; Henry &
McLaughlin, 1967:47; Tarasov & Zevina, 1957:227;
ZuUo, 1969b:351.

Distribution: Alaska to Oregon; 28-80m. Pleistocene,
Oregon.
Solidobalanus (Hesperibalanus) fallax (Broch), 1927:26

Synonymy'DIagnosls: Stubbings, 1963b:30.

References: Barnes & Klepal, 1971:86 (pedicel of Penis);
Bassindale, 1961:485; Henry & McLaughlin, 1967:47;
Nilsson-CanteU, 1939c:93; Stubbings, 1961b:34 (as
Balanus (Solidobalanus) occidentalis n.sp.); 1961c:189
(as occidentalis): 1963:30; 1965:892; 1967:287; Utinomi,
1959b:402.

Distribution: Algeria and West Africa; 7-220m.
Solidobalanus (Hesperibalanus) hesperius hesperius (Pils-
bry), 1916:193

Synonymy diagnosis: Pilsbry, 1916:193.

References: Barnes & Barnes, 1959d:237 (naupliar
stages); 1965a:391 (size variation); Barnes & Klepal,
1971:85 (pedicel of penis); Broch, 1922:321; CornwaU,
1955b:35; Henry, 1942:127: Henry & McLaughlin, 1967:
47; Hiro, 1935c:225; 1939f:212; Kolosvary, 1943a:92;
Kriiger, 1940:464; Tarasov & Zevina, 1957:228; Utinomi,
1970:359; ZuUo, 1969b:351.

Distribution: North Pacific: Japan; Bering Sea; Alaska;
British Columbia; 60- 180m. Pleistocene, Oregon.
Solidobalanus (Hesperibalanus) hesperius laevidomiformis
(Kolosvary), 1941e:9

References: Barnes & Klepal, 1971:85 (pedicel of penis);
Kolosvary, 1943a:92; Henry & McLaughhn, 1967:47 (as
incertae sedis)

Distribution: Panama.
Solidobalanus (Hesperibalanus) hesperius laevidomus (Pils-
bry), 1916:196

Synonymy/diagnosis: Henry, 1940:31.

References: Barnes & Gonor, 1958:194 (neurosecretory
ceUs); CornwaU, 1955a:34; 1955b:35; Henry, 1942:110;
Henry & McLaughhn, 1967:47; Hiro, 1935c;227; Kolos-
vary, 1943a:92; Newman, 1975:269; Nilsson-Cantell,
1927a:785; Pilsbry, 1921:113; Tarasov & Zevina,
1957:228.

Distribution: Alaska to San Francisco; 8-338m.
Solidobalanus (Hesperibalanus) hesperius nipponensis (Pils-
bry), 1916:199

Synonymy/diagnosis: Pilsbry, 1916:199.

References: Henry & McLaughhn, 1967:47; Tarasov &
Zevina, 1957:228; Utinomi, 1958a:308.

Distribution: Pacific coast of Japan; 50m.
Solidobalanus (Hesperibalanus) parahesperius (Menesini),
1971:22

Reference: Plaziat & Caveher, 1973:2875 (paleo-ecology).

Distribution: Eocene and Ohgocene, Paris Basin.
Solidobalanus (Hesperibalanus) phineus (Kolosvary),
1956:187

Distribution: Eocene, Hungary.
Solidobalanus (Hesperibalanus) proinus (Woodring, in
Woodring & Bramlette), 1950:92

Synonymy: ZuUo, 1969a:16.

Distribution: PUocene, Southern California.
Solidobalanus (Hesperibalanus) sookensis (Cornwall),
1927b:400

Distribution: Miocene, Vancouver I., Canada.
Solidobalanus (Hesperibalanus) stenonotus (Pilsbry &
Olson), 1951:201

Distribution: Ohgocene, Ecuador.

52

SoUdobalanus (Hesperibalanus) varians (Sowerby), 1846:pl. 2
References Chapman, 1914:54.67; Darwin. 1854b:298;

Ortmann. 1902:250.
Distribution: Eocene, Patagonia.
SoUdobalanus (Hesperibalanus) uialovi (Kolosvary).
1961c:150
Distribution: Eocene, USSR.

Snhgeims Bathybalanus Hoek, 1913

SoUdobalanus (Bathybalanusj pentacrini (Hoek), 1913:230
Synonymy diagnosis: Hoek. 1913:230.
References: Hiro. 1936a:59; Newman & Ross. 1971:173.
Distribution: Banda Sea; 240-304m.

Genus Notobalanus n.gen

Notobalanus flosculus (Darwin). 1854b:290
Synonymy/diagnosis: Pilsbry. 1916:219.
References: Gruvel. 1905a:252; Kolosvary, 1943a:93;

Korschelt. 1933:27; Kriiger. 1940:466; Nilsson-Cantell.

1957:21; Weltner. 1895:291; 1897:271; Zevina & Kur-

shakova. 1973:183.
Distribution: Peru; Chile; Tierra del Fuego.
Notobalanus flosculus var. sordidus (Darwin), 1854b:290
Synonymy/diagnosis: Nilsson-Cantell. 1921:330.
References: Davadie, 1952:30; Fletcher, 1938:115;

Gruvel. 1903b: 141; 1905a:252; 1905b:345; Kolosvary,

1943a:93; Newman & Ross, 1971:169; Nilsson-Cantell,

1957:21; Weltner, 1895:291; 1897:271; 1898b:5; 1900:305.
Distribution: Chile to Tierra del Fuego. Miocene. Algeria.

Late Cenozoic. Kerguelen I.
Notobalanus vestitus (Darwin), 1854:286
Synonymy diagnosis: Newman & Ross, 1971:169.
Refere.wes: Broch. 1922:322; 1931:71; Filhol. 1885:487;

Foster, 1967a:83; 1967b:35; Gruvel, 1905a:248; Hutton,

1879:328; Kriiger, 1940:464.466; Moore. 1944:333;

PUsbry. 1916:219; Weltner. 1897:271; 1899a:445; 1900:

307; Withers, 1924:36; 1953:77,98.
Distribution: Australia; New Zealand; Auckland I.;

0-51m. Lower Ohgocene. Chatham 1.

Genus Elminius Leach, 1825

Elminius cristallinus Gruvel. 1907a: 106

Synonymy/diagnosis: Gruvel, 1909a:216 (probably intro-
duced E. modestus Darwin).

Reference: Kriiger, 1940:470.

Distribution: Azores.
Elminius kingii Gray, 1831:13

Synonymy/diagnosis: Darwin, 1854b:348; Nilsson-Cantell,
1921:348.

References: Gruvel. 1903b:163; 1905a:294; 1911:292
Hoek, 1907:4; Kolosvary, 1943a:95; Kriiger. 1940:470:
NUsson-CanteU, 1930c:255; 1957:22; PUsbry, 1916:260
Weltner. 1895:289; 1897:256; 1898b:5; 1900:305.

Distribution: Chile, below 30°S; Cape Horn to Punta
Arenas, Argentina; Falkland Is.
Elminius modestus Darwin, 1854b:350

Synonymy/diagnosis: Moore, 1944:329 (includes E. sinu-
atus Hutton, 1879:328).

References: Austin et al, 1958:497 (chromosome num-
bers); Barnes & Barnes, 1960:137 (recent spread in NW
Europe); 1961a:121 (spread in SW Scotland); 1965b:23
(further European records); 1966a:83 (coasts of western
Europe); 1968a:135 (egg numbers); 1968b:261 (French
Atlantic coast); 1969a:156 (in France); 1974:197 (embry-
onic development and sahnity); Barnes & Klepal. 1971:
87 (pedicel of penis); Barnes, Barnes & Klepal, 1972:187
(French Atlantic coast); Barnes, Klepal & Munn. 1971:
173 (spermatozoa); Barnes & Powell, 1966:107 (at
Arcachon, France); Barnes & Stone, 1972b:309 (western
Scotland); Bassindale, 1947:223; 1958:381; 1964:42;
Beard, 1957:1145; Bhatnagar & Crisp, 1965:419 (salinity
tolerance); Bishop, 1947:501 (first report outside Aus-

traha); 1951:531 (spread in European waters); 1954:1145
(in France); Bishop & Crisp. 1957:482 (in France); 1958:
109 (France); Bishop et al. 1957:1 (on French Atlantic
coast); Bocquet-Vedrine. 1964:5060 (relationship be-
tween growth and molting); 1965b:30 (molting);
Boschma. 1948:403 (Netherlands); Boulton et al. 1971:
487 (fouhng); Broch. 1922:342 (as Elminius sinuatus);
ConneU, 1955:954 (Scotland); Corner et al, 1968:29
(toxicity); Crisp, 1955:569 (cyprid behavior); 1958:483;
1959b:37; 1960a:681 (northern limit); 1961:421 (territor-
ial behavior); 1964a:179 (severe winter); Crisp & Austin,
1960:787 (fouhng); Crisp & Barnes. 1954:142 (orienta-
tion); Crisp & Chipperfield. 1948:64 (British waters);
Crisp & Christie, 1966:59 (toxicity); Crisp & Davies,
1955:357 (breeding); Crisp & Meadows. 1962c:500 (gre-
gariousness); Crisp & Patel, 1958:1078; 1961:105
(growth rates); 1967:612 (contour of substratum); Crisp
& Ritz, 1967b:236 (temperature accUmation); Crisp &
Southward, 1959:429 (spread in British Isles); 1961:271
(cirral activity); Crisp & Stubbings, 1957:179 (orienta-
tion); Den Hartog, 1953:9 (in North Sea); 1956:141 (in
France); Evans, 1968:260; Filhol. 1885:489; Fischer-
Piette. 1963:176 (French/Spanish border); 1964:500
(France); 1965:466 (Fr. Atlantic coast); Fischer-Piette &
Prenant. 1956:7 (NW coast Spain); Foster. 1967a:84;
1967b:33 (early stages); 1969:326 (tolerance of high-
temperatures); 1970:380 (acclimation to salinity);
197 la: 12 (dessication); 1971b:33 (upper limit); Foster &
Nott, 1969:340 (sensory structures); Gruvel, 1903b:163
1905a:295 (as Elminius sinuatus), 296; Guiler, 1952:20
Hutton, 1879:328 (as E. sinuatus): Jennings, 1918:62
Jones, 1961:103 (SE coast of Scotland); Knight-Jones
1948:201 (British harbors); 1953:583 (gregariousness)
1955:266 (gregariousness); Knight-Jones & Crisp, 1953
1109 (gregariousness); Knight-Jones & Morgan, 1966:
267 (hydrostatic pressure); Knight-Jones & Stephenson
1950:281 (gregariousness); Knight-Jones & Waugh
1949:413 (larval development); Kolosvary, 1967b:393
Kriiger, 1940:470; Kuhl, 1963:99 (German coast); 1967
965 (Elbe estuary); Leloup & Lefevre, 1952:1 (Belgian
coast); Meadows, 1969a:273 (fouhng); 1969b:65 (settle-
ment, growth); Moore, 1944:329; Moyse, 1963:175 (food
for larvae); Moyse & Nelson-Smith, 1963:1 (zonation);
Nilsson-Cantell, 1921:351; 1925:42 (as var. laeuis n.var.);
1926:13; 1927a:786; 1930d:212; O'Riordan, 1967:294
(Ireland); Patel & Crisp, 1960a:667 (influence of tem-
perature); 1961:89 (breeding and molting); Pope, 1945:
368; 1966:181; PoweU, 1960:119 (Scotland); Ritz &
Foster, 1968:552 (temperature); Roskell, 1962:263 (on
Littorina shells); Sandison, 1950:79 (S. Africa); Singa-
rajah et al, 1967:144 (phototactic behavior); Skerman,
1958:224 (fouling); 1960:610 (predation); Southward,
1955b:403 (behavior); 1955c:423 (behavior); Southward &
Crisp, 1952:416 (changes in distribution); 1963:24 (foul-
ing); Stubbings, 1950:277; Utinomi. 1968b:178; Walker,
1970:239 (cement apparatus); 1973b:455 (frontal horns
and gland cells); Weltner, 1897:256,257 (as Elminius sin-
uatus); 1900:307; Wisely & BUck, 1964:162 (naupUi);
Zevina, 1963:73 (Black Sea).

Distribution: Austraha, Tasmania, New Zealand; intro-
duced to South Africa, possibly Azores (see E. cristal-
Unus Gruvel), Black Sea and Northern Europe.
Elminius modestus molluscorum Kolosvary. 1942c: 147

Reference: Kolosvary. 1943a:95.

Distribution: Auckland, New Zealand.

Genus Membranobalanus Pilsbry. 1916

Membranobalanus brachialis (RoseU), 1973:184
Distribution: Puerto Galera. Phihppines.

Membranobalanus cuneiformis (Hiro). 1936b:627
Synonymy/diagnosis: Hiro, 1939c:243.
Distribution: Arafura Sea and Japan; 15m.

53

Membranobalanus declivis (Darwin), 1854b:275

Synonvmv/diagnosis: Barnes & Klepal, 1971:86 (pedicel of
penis); Pilsbry, 1916:230.

References Gruvel. 1905a:244; Henry, 1954:443; 1958:
215; Hiro, 1936:631; VerriU, 1901:22 (as Balanus
declivus cuspidatus n.subsp.); Wells, 1966:83: Weltner,
1897:270,

Distribution: Bermuda: Florida; West Indies.
Membranobalanus longirostrum (Hoek), 1913:205

Synonymy: Utinomi, 1968b:176.

Diagnosis Hoek, 1913:205; Utinomi, 1968b:176.

References Broch, 1931:85; 1947:6; Daniel, 1955:26 (as
Balanus longirostrum var. krusadaiensis nov.); Dawyd-
off, 1952:128; Daniel & Prem-Kumar. 1968:147 (as
Balanus (Membranobalanus) roonwali n.sp.); Hiro,
1936b:631; Nilsson-Cantell, 1921:340; 1938b:54;
Suhaimi, 1966:65 (as Balanus (M.) basicupula n.sp.);
Weltner, 1897:270 (as IBalanus declivis).

Distribution: East coast of India to Singapore; 6-36m.
Membranobalanus nebrias (Zullo & Beach), 1973:2

Distribution: Galapagos Is.
Membranobalanus orcutti (Pilsbry), 1907b:361

Synonymy/diagnosis Pilsbry, 1907b:361

References Barnard, 1924:75; Henry, 1942:127; Hiro,
1936b:631; Pilsbry, 1916:233.

Distribution: Monterey to Baja California; South Africa;
Sulu Arch.; to 52m.
Membranobalanus orcuttiformis (Kolosvary), 1941:189

Distribution: Indian Ocean.

Genus Acasta Leach, 1817

Acasta aculeata Nilsson-Cantell, 1921:342

Distribution: Gulf of Siam.
Acasta alba Barnard, 1924:83

Distribution: Off Natal, South Africa; 90-180m.
Acasta alcyonicola Utinomi, 1953:142

Synonymy/diagnosis Utinomi, 1953:142.

Reference: Utinomi, 1959c:313.

Distribution: Tanabe Bay, Japan.
Acasta angusticalcar Broch, 1931:106

Synonymy/diagnosis Broch, 1931:106.

Distribution: Kei Is.; 20m.
Acasta antipathidis Broch, 1916:13

Synonymy/diagnosis Broch, 1916:13.

References: Hiro. 1936a:58; 1937b:53 (possibly a
Conopea).

Distribution: Cape Jaubert, western Australia.
Acasta armata Gravier, 1921a:353

Distribution: Off Djibouti; 20m.
Acasta cancellorum Hiro, 1931:151

Synonymy/diagnosis Hiro, 1937c:459.

References Nilsson-Cantell, 1938b:58.

Distribution: Seto, Japan.
Acasta conica Hoek, 1913:235

Synonymy/diagnosis Hoek, 1913:235.

References Broch. 1922:330.

Distribution: Celebes; Sulu Arch.; 40-60m.
Acasta coriobasis Broch, 1947:25

Synonymy/diagnosis Utinomi, 1953:139.

Reference: Dawydoff, 1952:139.

Distribution: Indochina; Hiroshima Bay.
/I casta crassa Broch, 1931:109

Distribution: Saparua Bay, Moluccas; 20-30m.
Acasta ctenodentia Rosell, 1972:200

Distribution: Puerto Galera, Philippines.
Acasta cyathus Darwin, 1854b:312

Synonymy: Darwin, 1854b:312.

Diagnosis Pilsbry, 1916:244.

References Annandale, 1906:144; Barnard, 1924:82:
Broch, 1922:330; 1927c:30; 1931:95; Gravier, 1921a:357:
Gruvel, 1903b:121; 1905a:259; Henry, 1954:443; Hoek,
1913:xvi,xix; Nilsson-CanteU, 1921:342; 1938:13; Pilsbry,
1916:244; 1952:27; Stubbings, 1936:48; WeUs, 1966:85;
Weltner, 1897:258; 1910:528; 1922:85; Zevina & Lit-

vinova, 1970:174.
Distribution: Florida; Caribbean; Madeira; Morocco;

East Africa; Red Sea; Gulf of Manaar; Singapore; Kei

Is.; Sulu Arch.; Philippines; western Australia; 15-180m.
Acasta denticulata Hiro, 1931:153
Synonymy;diagnosis Hiro, 1931:153.
Reference: Utinomi, 1958a:309.
Distribution: Sagami Bay, Japan.
Acasta dolfleini Kriiger, 1911a:56
Snonymy: Hiro, 1937d:454.
Diagnosis Broch. 1922:330.
References Broch, 1931:96; Dawydoff, 1952:129; Hiro,

1931:151 (as Acasta aperta n.sp.); Kriiger, 1911b:461;

Nilsson-Cantell, 1921:341; Pilsbry, 1916:247; • Rosell,

1972:198; Utinomi, 1950:64; 1958a:309; 1962:221; 1968b:

178; 1969b:53.
Distribution: Southern Japan; Indonesia; Sulu Arch.;

24-280m.
Acasta echinata Hiro, 1937a:70
Synonymy: Utinomi, 1962:224.
Diagnosis Broch, 1947:6; Utinomi, 1949a:32.
References Dawydoff, 1952:129; Utinomi, 1959c:313.
Distribution: Southern Japan; southeast Asia; 15-20m.
Acasta fenestrata Darwin, 1854b:316
Synonymy: Rosell, 1972:194.
Diagnosis Darwin, 1854b:316.
References Gruvel, 1905a:262; Hiro, 1939d:243; Hoek,

1883:160; 1913:233; Kriiger, 1911a:4; 1911b:461;

Nilsson-CanteU, 1938b:57; Weltner, 1897:259; 1922:104.
Distribution: Red Sea; Bay of Bengal; Philippines;

Seto, Japan; to 51m.
Acasta fischeri Locard, 1877:18

Distribution: Miocene, Sicily, Sardinia and Corsica.
Acasta flexuosa Nilsson-CanteU, 1931a:130
Synonymy: Hiro, 1937c:457.

Diagnosis: Hiro, 1931:153 (as Acasta amakusana n.sp.).
References Utinomi, 1949a:32.
Distribution: Amakusa and Seto, Japan.
Acasta foraminifera Broch, 1931:98

Distribution: Amboina Bay, Kei Is.; 100-140m.
Acasta formae de Alessandri, 1897:46

Reference: de Alessandri, 1906:312; Withers, 1953:61.
Distribution: Miocene. Italy.
Acasta fossata Barnard, 1924:84

Distribution: Seal I., S. Africa; 24-50m.
Acasta glans Lamarck, 1818:398
Synonymy: Utinomi, 1969a:91.
Diagnosis Hoek, 1913:241.
References Broch, 1931:133; Darwin, 1854b:314: Gruvel,

1903b:121; 1905a:261; Hoek, 1913:233; Kolosvary,

1943a:94; NUsson-CanteU, 1938b:56; Pilsbry, 1916:242;

Weltner, 1897:258.
Distribution: Gulf of Iran; Bay of Bengal; southwest

AustraUa; Indonesia; PhiUppines; 15-55m.
Acasta gregaria Utinomi, 1959c:314
Distribution: Tanabe Bay, Japan.
Acasta hirsuta Broch, 1916:10
Synonymy/diagnosis Broch, 1916:10.
References Broch, 1931:96; Hiro, 1936a:58; Utinomi,

1959c:317.
Distribution: Cape Jaubert, AustraUa; Amboina; Moluc-
cas; 100-140m.
Acasta idiopoma Pilsbry, 1912:294
Reference: Pilsbry, 1916:247.
Distribution: Mindanao, Philippines; 40m.
Acasta japonica Pilsbry, 1911:80
Synonymy: Utinomi, 1962:221.
Diagnosis Pilsbry, 1911:80.
References Broch, 1922:330; 1931:96; 1947:6; Dawydoff,

1952:129; Hiro, 1939f:213; Kruger, 1940:460; NUsson-

CanteU, 1938b: 13; Pilsbry, 1916:243; Utinomi, 1970:359;

ZuUo, 1968a;227.
Distribution: Southern Japan; Taiwan; southeast Asia;

10-800m.

54

Acasta laevigata Gray, 1825:103
Synonymy: Nilsson-Cantell. 1938b:57.
Diagnosis Darwin, 1854b:315; Hiro, 1937b:62.
References Gruvel, 1903b:121; 1905a:261; Hoek, 1913:
233; Resell, 1972:197; Weltner, 1897:259; Zevina &
Litvinova, 1970:174.
Distribution: Red Sea; Zanzibar; Andaman Is., Phil-
ippines; Palau Is.
Acasta membranacea Barnard, 1924:88
Reference: Nilsson-Cantell, 1938b:13.
Distribution: South Africa; 28-180m.
Acasta microforamina Rosell, 1970:105

Distribution: Philippines.
Acasta muricata Seguenza, 1876:312

Distribution: Tertiary, Italy (Withers, 1953:62).
Acasta pectinipes Pilsbry, 1912:294
Synonymy: Hiro, 1937c:463.
Diagnosis Nilsson-Cantell, 1938b:57.
References Barnard, 1924:87; 1925:1; Broch, 1922:330;
Hiro, 1931:149 (as Acasta komaii n.sp.); Hoek, 1913:237
(as Acasta nitida n.sp.); Kriiger, 1914:438; Pilsbry, 1916:
247; Utinomi, 1949a:23; 1962:221; 1969b:53; ZuUo,
1968a:227.
Distribution: Japan; Philippines; Sulu Arch.; Java Sea;
Andaman Is.; South Africa; 35-170m.
Acasta porata Nilsson-Cantell, 1921:348
Synonymy: Nilsson-Cantell, 1938b:56.
Diagnosis Broch, 1931:96.
References Broch, 1947:6; Dawydoff, 1952:129, RoseU,

1972:203.
Distribution: Philippines; Viet Nam; Bay of Bengal;
l-55m.
Acasta purpurata Darwin, 1854b:318
Synonymy/diagnosis Darwin, 1854b:318.
References Gruvel, 1905a:262; Hiro, 1937c:450; Hoek,
1913:234; Nilsson-Cantell, 1938b:13; Utinomi, 1959c:
313; Weltner, 1897:259.
Distribution: Sumatra; PhiUppines.
Acasta sarda de Alessandri, 1895:64
References de Alessandri, 1906:311; Withers, 1953:59.
Distribution: Ohgocene, Sicily.
Acasta schdfferi de Alessandri. 1910:124

Distribution: Miocene, Austria (Withers, 1953:70).
Acasta sculptura Broch, 1931:101
Reference: Utinomi, 1959c:313.
Distribution: Java Sea; 49m.
Acasta scuticosta Weltner, 1887:102
Synonymy/diagnosis Weltner, 1887:102.
References Gruvel, 1905a:260; Hiro, 1931:152; Hoek,

1913:233; Weltner, 1897:259.
Distribution: Cartagena, Spain.
Acasta semota Hiro, 1933:73

Distribution: East coast of Japan; 33m.
Acasta serrata Hiro, 1937b:64

Distribution: Palau Is.
Acasta spinitergum Broch, 1931:112
Synonymy/diagnosis Broch, 1931:112.
References Hiro, 1936a:58; Kolosvary, 1943a:95;

Utinomi, 1959c:313.
Distribution: Java Sea; Philippines; 35m.
Acasta spinifera Utinomi, 1967:222

Distribution: Tokyo Bay; 70m.
Acasta spinosa Hiro, 1939e:267

Distribution: Formosa.
Acasta spongites Poll, 1791:25
Synonymy: Darwin, 1854b:308.
Diagnosis Utinomi, 1958a:300.

References Barnard, 1924:80; Bassindale, 1964:42;
Bocquet-Vedrine, 1966a:2733; 1966b:337 (structure and
growth of operculum); 1966c:693; Crisp & Southward
1961:271 (cirral acivity); Dawydoff, 1952:129; de Ales
sandri, 1907b:289; Fischer, 1872:433; Gruvel, 1903b:121
1905a:263; Hiro, 1931:154; 1935c:227; Hoek, 1875:60
1909:271; Kolosvary, 1939a:176; 1941c:156; 1943a:94

1947a:22; Kriiger, 1911a:4; 1911b:459; 1914:439; 1940:
459; LeReste, 1965:65 (larvae); Moroni-Ruggieri, 1952:
77; Moyse, 1960:120 (laboratory rearing); 1961:371
(larval stages); Nilsson-Cantell, 1938b:13; Pilsbry, 1916
242; ReUni. 1969:169; Riedl, 1963:258; Stebbings, 1910
570; Utinomi, 1969a:82; Weltner, 1897:259; 1898:11
Zevina & Litvinova, 1970:174.
Distribution: British Isles; France; Portugal; Mediterran-
ean; Red Sea; South Africa; Australia; Japan. Pliocene:
France and Italy.
i4casfa sporittus Darwin, 1854b:319
Synonymy/diagnosis Darwin, 1854b:319.
References Gruvel, 1905a:260; Hiro, 1931:150; Hoek,

1913:233; Weltner, 1897:259,
Distribution: Sulu Arch.
Acasta striata Gruvel, 1901:262
Synonymy DIAGNOSIS Gruvel, 1905a:264.
Reference: Hoek, 1913:233.
Distribution: Off Madeira; 400m.
Acasta sulcata Lamarck, 1818:398
Synonymy: Darwin, 1854b:310; Hiro, 1937c:451.
Diagnosis Stubbings, 1961a:174.

References Borradaile, 1903:442; Broch, 1947:6; Dawyd-
off, 1952:129; Gruvel, 1903b:121; 1905a:263; Kolosvary,
1947e:361; Kriiger, 1911a:56; 1911b:461; Nilsson-CanteU,
1938b:13; Utinomi, 1950:64; 1958a:309; 1969a:82: Welt-
ner, 1897:259; Zevina & Litvinova, 1970:174.
Distribution: Red Sea; Persian Gulf; Maldives; Gulf of
Siam; South China Sea; Viet Nam; Austraha; PhiUp-
pines; Japan; 5-25m.
Acasta sulcata anchoris Barnard, 1924:81

References Nilsson-Cantell, 1938b:13; Zevina & Lit-
vinova, 1970:174.
Distribution: Natal, South Africa; 28m.
Acasta sulcata spinosa Daniel, 1955:29

Distribution: Off Madras, Bay of Bengal.
Acasta tenuivalvata Broch, 1947:28
Reference: Dawydoff, 1952:129.
Distribution: Viet Nam; 15m.
Acasta tulipa Hiro, 1933:76

Distribution: Off southern Japan; 126m.
Acasta undulata Darwin, 1854a:34
Distribution: Upper Pliocene, England (Withers,
1953:53).
Acasta umitosaka Utinomi, 1962:224

Distribution: Nomosaki, Japan.
Acasta zuiho Hiro, 1936b:632
Distribution: Off Port Darwin, Australia.

Genus Pseudoacasta Nilsson-Cantell. 1930

Pseudoacasta libera Nilsson-Cantell, 1930a:l
Synonymy/diagnosis Nilsson-Cantell, 1930b:ll.
Distribution: Moluccas.

Genus Conopea Say, 1822

Conopea acuta (Nilsson-Cantell), 1921:334

Distribution: Kyusyu, Japan.
Conopea calceola (Ellis), 1758:853

Synonymy: Hiro, 1937c:443.

Diagnosis McLaughlin & Henry, 1972:24.

References de Alessandri, 1895:281; 1906:299; Broch,
1922:325; 1924b:202; 1927c:29; 1931:85; Daniel, 1955:28;
Darwin, 1854a:15; 1854b:218; Davadie, 1963:35; Gauld,
1957:10; Gruvel, 1903b:130; 1905a:221; 1907b:164;
1909b:25; Hoek, 1913:221; Kolosvary, 1947e:361;
Kriiger, 1911a:4; 1911b:460; 1940:459; Nilsson-Cantell,
1928a:34; 1938a:180; 1938b:55; Pilsbry, 1916:238; Relini,
1969:175; Stebbing, 1910:568; Stubbings, 1963b:36;
1964:343; 1967:290; Utinomi, 1949a:23; 1950:60; 1958a:
296; 1959b:403; 1962:218; 1969a:91; 1969b:53; Utinomi
& Kikuchi, 1966:6; Weltner, 1897:262; Withers, 1953:61.

Distribution: Mediterranean to South Africa; Indian
Ocean; Persian Gulf to western Australia and Japan;

55

18-250m. Miocene to Pleistocene, Italy; Coralline Crag,
England.
Conopea comuta (Hoek), 1913:227

References Broch, 1931:87; Utinomi, 1962:219; Utinomi
& Kikuchi, 1966:6.

Distribution: Banda Sea to Japan; 32-140m.
Conopea cymbiformis (Darwin), 1854b:221

Synonymy: Utinomi, 1962:219.

Diagnosis Darwin, 1854b:221.

References Broch, 1922:326; 1931:85; Daniel, 1955:28;
Dawydoff, 1952:128; Foster, 1974:48; Gruvel, 1905a:222;
Hiro! 1935a:25; Hoek, 1913:228 (as Balanus proripiens
n.sp.), 2G2 {as Pyrgoma jedani n.sp.); Kruger, 1911a:4
1911b:460; 1940A:464; Nilsson-Cantell, 1921:331; Stub-
bings, 1935:48; Utinonii, 1949a:23; 1958a:297; Utinomi
& Kikuchi, 1966:6.

Distribution: Indo-west Pacific: Gulf of Aden; India;
Indonesia; Phihppines; Japan; 27-453m.
Conopea dentifer {Broch). 1922:326

References Broch, 1931:88; Kolosvary, 1967b:392;
Kruger, 1940:464.

Distribution: Tonga I.; Japan; Kei Is., 180m.
Conopea foUiculus Hiro, 1937b:53

Distribution: Marianas Is.; on antipatharian.
Conopea fragilis (Broch), 1931:92

References Kruger, 1940:464.

Distribution: Amboina Bay; 100-140m.
Conopea galeata (Linnaeus), 1771:544

Synonymy: Ross, 1962:31.

Diagnosis Pilsbry, 1916:236.

References Caziot, 1921:52; Cornwall, 1951:325; Darwin,
1854b;220; Gomez, 1973:163 (setthng sites); 1975:105
(sex determination); Gomez et al, 1973:813 (effect of
juvenile hormone mimics on metamorphosis); Gravier,
1921b:430; Gruvel, 1903b: 130; 1905a:222; Henry, 1942:
126; 1954:443; Kolosvary, 1943a:93; Kruger, 1940:464;
McDougaU, 1943:343; McLaughlin & Henry, 1972:13
(comparative morphology of complemental males)
Molenock & Gomez, 1972:100 (larval stages); Morch
1852:67; Nilsson-Cantell, 1931a:114; 1939a:3; Patton
1963:522; PUsbry, 1907d:204; 1927:37; 1953:25; Ramen
ofsky, et al, 1974:172 (juvenile hormones and metamor
phosis); Say, 1822:323 (as Conopea elongata); Wells
1966:84; Weltner, 1897:262; Zevina & Kurshakova
1973:183; ZuUo, 1966b:237.

Distribution: North Carolina through West Indies and
Gulf of Mexico to Venezuela; Southern CaUfornia to
Panama and Galapagos Is.; 2-540m.
Conopea granulata (Hiro), 1937c:444

Synonymy/diagnosis: Hiro, 1937c:444.

References Hiro, 1939e:266; Utinomi, 1949a:23; 1950:64;
1958a:307; 1962:220; 1970:359; Utinomi & Kikuchi,
1966:6.

Distribution: Japan; Taiwan; 90-200m.
Conopea investita (Hoek), 1913:244

References Pilsbry, 1916:235.

Distribution: Java and Banda Seas; 73-90m.
Conopea tongibasis Hiro, 1937b:52

Distribution: Palao Is.
Conopea merrilli Zullo, 1966b:237

References McLaughlin & Henry, 1972:13 (comple-
mental males).

Distribution: North Carolina; Gulf coast of Florida;
Puerto Rico; 2-46m.
Conopea mojbergi Borch, 1916:7

Distribution: Cape Jaubert, Australia; on Echinogorgia.
Conopea navicula (Darwin), 1854b:221

Synonymy: Utinonii, 1962:218.

Diagnosis Hoek, 1913:223.

References Dawydoff, 1952:128; Gruvel, 1905b:222;
Kruger, 1940:464; Nilsson-Cantell, 1938b:55; Stubbings,
1936:48; Utinonu, 1969a:91; Utinomi & Kikuchi, 1966:6.

Distribution: Indo-west Pacific: Gulfs of Aden, Persia
and Siam; Indonesia; southern Japan; 45-220m.

Conopea pygmaea (Broch), 1931:88

Distribution: Banda Sea; 85m.
Conopea scandens (Pilsbry), 1916:239

References Barnard, 1924:76; Kriiger, 1940:464;
Nilsson-Cantell, 1921:334; Utinomi, 1958a:308.

Distribution: Japan; South Africa; 110-250m.

Genus Eoceratoconcha Newman and Ladd, 1974

Eoceratoconcha kugleri Newman & Ladd, 1974:387

Distribution: Middle Miocene, Trinidad.
Eoceratoconcha renzi Newman & Ladd, 1974:389

Distribution: Middle Miocene, Trinidad.

Subfamily Semibalaninae n. subfam.

Genus Semibalanus, Pilsbry, 1916

Semibalanus balanoides (Linnaeus), 1767:1108
Synonymy: Darwin, 1854b:267; Pilsbry, 1916:183; Nilsson-

CanteU, 1921:328.
Diagnosis Darwin, 1854b:267; Pilsbry, 1916:183; Stub-
bings, 1975:1.
References Allison & Cole, 1935:34; Arnold, 1970:1045
(response to lowered saUnity); Arvy & Lacombe,
1968:1326; Arvy c& Liguori, 1968:817; Arvy et al,
1968:817; Arvy & Nigrelli, 1969:95 (epizoic peritriches
in branchial cavity); Arvy et al, 1969:351; Aurivillius,
1898b:29; Austin et al, 1958:497 (chromosomes); Barnes,
1950:73 (larvae); 1952-53:104 (rate of growth); 1953b:328
(lowered salinity); 1953d:429 (southern Umits); 1953e:297
(size variations); 1955a:109 (growth rate); 1955b:114
(hatching); 1955c:341 (rugophilic behavior); 1956a:72
(larval population); 1957a: 1 (northern Umits); 1957b:67
(spawning); 1958:139 (southern limits); 1959c:234 (tem-
perature and hfe cycle); 1961a:592 (observations on
southern Umit); 1961b:427 (growth rate); 1962b:462
(anecdysis); 1963a:717 (breeding); 1965:321 (egg bio-
chemistry); Barnes & Barnes, 1958b:160 (opening re-
sponse); 1958c:29 (rate of larval development); 1959a:l
(growth patterns); 1959b:19 (stimulation of nauplii)
1959e:242 (egg mass development); 1959g:581 (growth)
1962:1 (distribution); 1963:93 (egg development); 1965a
391 (variation in egg size); 1966a:83 (observations on
western European mainland); 1967:1 (starvation);
1968a:135 (variation in egg production); 1969b:36 (oxy-
gen consumption); 1969c:136 (Umits in France); 1974:197
(embryonic development and saUnity); Barnes et al,
1963:213 (metaboUsm); 1970:70 (behavior on impaction);
1971:173 (spermatozoa); 1972:189 (on French Atlantic
coast); Barnes & Finlayson, 1963:185 (seasonal changes);
Barnes & Healy, 1965:779 (biometrical studies); Barnes
& Klepal, 1971:85 (Pedicel of penis); Barnes & PoweU,
1950a:175 (development, morphology and elmination);
1966:107 (at Arcachon, France); Barnes & Stone, 1972a:
303 (penis development); 1974:275 (food, temperature,
photoperiod and molting); Bassindale, 1936:57 (develop-
ment); 1958:381 (in England); Belyaev, 1949:901
(osmoregulation); Bhatnagar & Crisp, 1965:419 (salinity
tolerance of larvae); Bishop et al, 1957:3 (in France);
Blom & Nyhohn. 1961:149 (settUng time, Sweden):
Bourget & Crisp, 1975a:231 (shell deposition); 1975b;221
(early changes in sheU form); Bousfield, 1954:118
1955a: 1 (ecology — Miramichi estuary); 1955b:763
Brattstrom, 1957:5; Brocchi, 1814:598; Broch, 1924a:84
1927b:22; Caziot, 1921:52; Chipperfield, 1948:13 (breed
ing and settlement); 1949:17 (environmental conditions)
Ciurea et al, 1933:6; Cole, 1929:599 (temperature and
pedal rhythm); 1932a:611 (stimulation); 1932b:143 (stim-
ulation); Cole & AUison, 1935:25 (stimulation); 1937:405
(electrolytes); ConneU, 1957:1; 1959:226 (recruitment and
mortaUty); Cook et al, 1972:409 (amino acid composi-
tion); Cook & Gabbott, 1970:11 (glycerol level); Cook &

56

Lewis. 1971:26 (cold tolerance); Crisp, 1953:331 (changes
in orientation); 1955:569 (cyprid behavior); 1956:263
(hatching); 1959a:275 (breeding); 1959c:119 (embryo
development); 1960b:95 (growth); 1960c:1208 (mobility);
1961:429 (behavior); 1962a:207 (planktonic stages);
1964a:165 (effect of severe winter); 1964b:33 (racial
differences); 1968a:2633 (difference in N. American and
European populations); 1968b:1161 (distribution of para-
sitic isopod); 1969:1037 (hatching substance); Crisp &
Austin, 1960:787 (fouling): Crisp & Barnes, 1954:142
(orientation); Crisp & Clegg, 1960:265 (induction of
breeding); Crisp & Knight-Jones, 1953:360 (aggregation);
Crisp & Meadows, 1962:500 (chemical basis of gregari-
ousness); 1963:364 (stimulus to settlement); Crisp &
Patel, 1958:1078 (breeding and ecdysis); 1960:31 (molt-
ing); 1967:612 (contour of substratum); 1969:283 (control
of breeding); Crisp & Ritz, 1967a:98 (temperature toler-
ance); 1967b:236 (temperature acclimation); 1974:327
(larval response to light); Crisp & Southward, 1961:271
(cirral activity); Crisp & Spencer, 1958:278 (hatching);
Crisp & Stubbings, 1957:179 (orientation); Crisp et al,
1967:629 (toxic action); Daniel, 1955c:23; Davadie, 1963:
70; Dawson & Barnes, 1966:249 (lipid composition);
Fales, 1928:534 (light receptive organs); Fischer, 1929:10
(distribution in English Channel); Fischer, 1872:433
(southwestern coast of France); Fischer, 1943:65 (distri-
bution — North Sea) Fischer-Piette, 1930:39 (St. Servan);
Fischer- Piette and Prenant, 1956:8 (northern Spain);
Forbes et al, 1971:539 (orientation to hght); Foster,
1969:326 (tolerance of high temperatures); 1970:377
(acchmation to salinity); 1971a;12 (dessication); 1971b:33
(upper limits of intertidal distribution); Gabbott &
Larman, 1971:143 (electrophoretic examination); Gibson
& Nott, 1971:227 (larvae); Gordon, 1969:139 (salinity &
distribution); Grainger & NeweU, 1965:469 (aerial respi-
ration); Groom, 1894b:81; 1895a:l; 1895b:269 (cyprid
stage); Gruvel, 1903b:139; 1905a:241; 1909a:225;
Gutmann, 1960:1 (morphology); 1962:193 (breeding/
molting); Hatai, 1939b:267; Hatton & Fischer-Piette,
1932:1 (settUng and growth); Haven, 1973:97 (ecology);
Henry, 1942:100; Hiro, 1935d:222; Hoek, 1875:37 (Neth-
erlands coast); 1884:519; Kauri, 1962:131 (nauplius eye);
1966:115 (sensory papilla X-organ); Kaye, 1964:580 (as
index of sea level changes); Klepal & Barnes, 1974:205
(penis regeneration); Klugh & Newcombe, 1935:39 (light
control); Knight-Jones, 1953:583 (gregariousness); 1955:
266 (gregariousness); Knight-Jones & Crisp, 1953:1109
(gregariousness-fouhng); Knight-Jones & Morgan, 1964:
29 (barosensitivity); 1966:267 (hydrostatic pressure);
Kolosvary, 1943a:91; 1962d:201; Kruger, 1927a:14;
1927b:5; 1940:464; Kuhl, 1963:99; 1965:113; 1967:965;
1968:1; Lacombe, 1970:164 (cement glands); Meadows,
1969a:273 (fouling communities); 1969b:65; Mohammad,
1962:488; Moore, 1934a:101 (growth rate); 1934b:851
(growth); 1935b:264 (soft parts); 1935c:279 (ecology);
1936:701 (distribution); Moore & Kitching, 1939:521
(comparison with C. stellatus); Moore & Parke, 1935:49
(algal infection); Morch, 1852:68; Moyse, 1960:120 (labo-
ratory rearing); 1963:176 (food for larvae); Muller, 1940:
113 (sensitivity to poisons); Munn & Barnes, 1970a:277
(spermatozoa); 1970b:261 (spermatozoa); Munn, Klepal
& Barnes, 1974:89 (structure and function penis sen-
sory setae); Neu, 1935:169 (growth forms); Newell &
Northcroft, 1965:387 (cirral activity); Norris et al,
1951:444 (larval stages); Nott, 1969:251; Nott & Foster,
1969:115 (antennular attachment organ); O'Riordan,
1967:292; Patel & Crisp, 1960b:104 (embryo develop-
ment); 1961:89 (breeding and molting); Petersen, 1962:1
(distribution); 1966:1 (natural history); Poulsen, 1935:17;
Prenant & Teissier, 1923:172 (Roscoff); Pyefinch, 1948a:
451 (identification of larvae); 1948b:464 (biology); Ritz &
Crisp, 1970:223 (feeding); Rosenberg, 1972a:313 (effect
of chlorinated hydrocarbons); 1972b:ll (salinity toler-
ance); Roskell, 1962:263 (epizoic on Littorina); Runn-

strom. 1925:1 (biology); Rusanova, 1959:568 (two popu-
lations); Rzhepishevskii, 1968:37 (Barents Sea); Schafer,
1938a:304 (boring organisms); 1938b:323 (paleontology);
1938c:564; 1948:74; 1952:240 (settling); Schwarz, 1932:
437 (influence of Ught); Sizer, 1937:327 (stimulation by
acids); Sneh, 1972:3; Southward, 1955b:403 (cirral activ-
ity); Southward & Crisp, 1954a:163 (distribution British
Isles); 1956:211 (fluctuation in distribution); 1963:35;
Stephenson, 1938:5 (Iceland); 1943:20 (E. Greenland);
Tarasov, 1937:53; Tarasov & Zevina, 1957:216; Tighe-
Ford, 1967:920 (breeding); 1968:225 (techniques); Tighe-
Ford & Vaile, 1972a: 19 (molting hormone); 1972b:202
(molting hormone); Trusheim, 1932:70 (paleontology);
Visscher, 1928b: 193 (resistance to fresh water); Walley,
1964:314 (metamorphosis); 1965:115 (oviducal gland);
1967:151 (epidermal gland); 1969:237 (larval structure
and metamorphosis); WaUey et al, 1971:489 (sperm);
Walker, 1970:239 (cement apparatus); 1971:205 (larval
cement apparatus); 1973a:305 (early development of
cement apparatus); 1973b:455 (frontal horns and gland
cells); Wells, 1960:578 (southern Umit); Weltner, 1897:
269; 1898a:442; 1898b:8,ll; 1900:302; ZuUo, 1963b:12.
Distribution: Atlantic: boreo-arctic, to northern Spain
and Cape Hatteras; North Pacific from Unalaska to
British Columbia. Miocene, Japan; Pliocene, England;
fossil, Caspian Sea region.
Semibalanus balanoides calcaratus Pilsbry, 1916:188
Synonymy/diagnosis Pilsbry, 1916:188.
References Henry, 1942:126; Hiro, 1935c:227.
Distribution: Shelikof Strait and Sitka, Alaska.
Semibalanus cariosus (Pallas), 1788:240
Synonymy/diagnosis: Pilsbry, 1916:189.
References Barnes, 1959a:231 (stomach contents);
Barnes & Barnes, 1959h:515 (metabolism); Barnes &
Klepal, 1971:71 (pedicel of penis); BatzU, 1969:531 (dis-
tribution of biomass); Connell, 1970:49 (predation); Corn-
waU, 1924b:41; 1925:462; 1951:322; 1955a:22; 1955b:26;
Darwin, 1854b:273; Fahrenbach, 1965:234 (photorecep-
tors); Gruvel, 1903b: 140; 1905a:243; GwiUiam, 1965:244
(physiology); Gwilliam & Bradbury, 1971:502; Hatai,
1938:96; Henry, 1940:13; 1942:102; Hiro, 1932b:472;
1935c:223; 1939f:211; Hoek, 1913:154, 155; Kolosvary,
1967b:391; Kruger, 1911a:54; 1911b:459; Millecchia &
Gwilliam. 1972:438 (electrophysiology); Pilsbry, 1911:76;
1921:112; Rice, 1930:249 (peculiarities in distribution);
Southward & Crisp, 1965:161 (activity rhythms); Towler,
1930:225 (communities); Tarasov & Zevina, 1957:211;
Utinomi, 1955a:119; 1969b:51; 1970:358; Weltner,
1897:270; 1898b:ll; 1900:302; Worley, 1939:233 (correla-
tion study); Yamaguchi, 1971:122; ZuUo, 1969b:351.
Distribution: North Pacific; Japan, Korea, Bering Sea;
Unalaska to Central California. Miocene, Japan; Pleisto-
cene, Japan and Oregon.
Semibalanus madrasensis (Daniel), 1958:305
Diagnosis Daniel, 1958:305.
Distribution: Bay of Bengal; on local craft.
Semibalanus sinnurensis (Daniel), 1962a:193
Diagnosis Daniel, 1962a:193.
Distribution: Porto Novo, India; on Murex sp.

Family Pyrgomatidae Gray. 1825
Subfamily Pyrgomatinae Gray. 1825

Genus Cantellius Ross and Newman, 1973

Cantellius acutum (Hiro). 1938d:398

Synonymy: Utinomi, 1962:227. Ross & Newman. 1973:150.

Diagnosis Hiro. 1938d:398.

References Darwin. 1854b:379 (as Creusia spinulosa var.
6. subvar. 2); Foster. 1974:49; Gruvel. 1905a:300; Hiro,
1935a:25; Kolosvary, 1947e:365; Nilsson-Cantell,
1921:352.

Distribution: Philippines; Palau Is.; Japan.

57

Cantellius arcuatum (Hirol, 1938d:395
Synonymy/diagnosis: Hiro, 1938d:395.
References: Kolosvary, 1947d:426; 1947e;364; Ross &

Newman, 1973:150.
Distribution: Palau Is.
Cantellius brevitergum (Hiro), 1938d:397
Synonymy/diagnosis: Hiro, 1938d:397.
Reference: Ross & Newman, 1973:150.
Distribution: Palau Is.
Cantellius euspinulosum (Broch), 1931:118
Synonymy: Utinomi, 1962:226.
Diagnosis: Darwin, 1854b:377: Hiro, 1935a:5.
References: Annandale, 1924:64: Barnes & Klepal, 1971
87 (pedicel of penis); Darwin, 1854b:377 (as Creusia
spinulosa var. 1): Foster, 1974:48; Gruvel, 1903b:164:
1905a:299; Hiro, 1937c;465; 1938d: 393; Kolosvary
1947e:365; Nilsson-Cantell, 1938b:59; Ross & Newman,
1973:150; Utinomi, 1949a:23.

Distribution: Pacific coast of Japan; Palau Is.; Sulu
Arch.; Indonesia; Singapore; Mergui Arch.; Andamans.
Cantellius gregarius (Sowerby), 1823:no pagination
Synonymy: Ross & Newman, 1973:150.
Diagnosis: Darwin, 1854b:378; Nilsson-Cantell, 1938b:30.
References: Broch, 1931:118; Darwin, 1854b:378 (as
Creusia spinulosa var. 3); Gruvel, 1905a:299; Hiro.
1935a:25; 1938d:403; Kolosvary, 1947e:362; 1951b:292.
Distribution: Banda Sea; Singapore; Bay of Bengal; to
70m.
Cantellius iwayama (Hiro), 1938d:393
Synonymy/diagnosis: Hiro, 1938d:393.
Reference: Ross & Newman, 1973:150.
Distribution: Palau Is.
Cantellius madreporum (Borradaile), 1903:443
Synonymy: Ross & Newman, 1973:150.
Diagnosis: Borradaile, 1903:443.
References: Dawydoff, 1952:128: Hiro, 1935a:25; Hoek,

1913:xvi; Nilsson-Cantell, 1938b:13, 65 (footnote).
Distribution: Gulf of Siam (?); Maldives.
Cantellius octavus Ross & Newman, 1973:152
Synonymy: Ross & Newman, 1973:152.
Diagnosis: Darwin, 1854b:380; {as Creusia spinulosa var. 8).
Reference: Gruvel, 1905a:300.
Distribution: Unknown.
Cantellius pallidas (Broch), 1931:118
Synonymy/diagnosis: Hiro, 1935a:6.
References: Kolosvary, 1947d:'i25; 1947e:364; Ross &

Newman. 1973:152.
Distribution: Tanabe Bay, Japan; Singapore; Philippines;
Fiji; Banda Sea.
Cantellius pseudopallidum (Kolosvary), 1947e:362
Synonymy: Ross & Newman, 1973:153.
Diagnosis: Kolosvary, 1947e:362.
Distribution: Pacific area.
Cantellius quintus Ross & Newman, 1973:153
Diagnosis: Darwin, 1854b:379 (as Creusia spinulosa var.

5).
Reference: Gruvel, 1905a:300.
Distribution: Unknown.
Cantellius secundus (Broch), 1931:118
Synonymy: Utinomi, 1962:227; Ross & Newman, 1973:153.
Diagnosis: Darwin, 1854b:378 (as Creusia spinulosa var. 2).
References: Fishelson, 1971:122; Gruvel, 1903b:164;
1905a:299; Hiro, 1935a:25; 1938a:397; Kolosvary, 1947d:
425; Nilsson-Cantell, 1938b:60; Utinomi & Kikuchi,
1966:7; Zevina & Litvinova, 1970:175.
Distribution: Japan; China; Palau Is.; Kei Is.; Singa-
pore; Andaman Is.; Red Sea; to 20m.
Cantellius septimus (Hiro), 1938d:395
Synonymy: Ross & Newman. 1973:153.
Diagnosis: Darwin, 1854b:380 {asCreusia spinulosavar.l).
References: Kolosvary, 1941e:9; 1943a:104; 1947d:426;

1947e:364; Nilsson-Cantell, 1921:354.
Distribution: Philippines; Palau Is.; Indian Ocean.

Cantellius sextus (Hiro), 1938:398

Synonymy: Ross & Newman, 1973:153.

Diagnosis: Darwin, 1854b:379 (as Creusia spinulosa var.
6. subvar. 3).

Reference: Gruvel. 1905a:300.

Distribution: Philippines; Palau Is.
Cantellius sumbawae (Hoekl. 1913:265

Synonymy: Ross & Newman. 1973:153.

Diagnosis: Hoek. 1913:265.

Distribution: Sunda Is.; to 36m.
Cantellius transversalis (Nilsson-Cantell). 1938a:61

Synonymy: Ross & Newman, 1973:153.

Diagnosis: Nilsson-Cantell. 1938a:61.

References: Darwin. 1854b:379 (as Creusia spinulosa var.
6, subvar. 1); Gruvel, 1903b:164; 1905a:300; Nilsson-
Cantell, 1921:352.

Distribution: Philippines; Andaman Is.
Cantellius tredecimus (Kolosvary), 1947d:426

Synonymy: Ross & Newman. 1973:153.

Diagnosis: Kolosvary. 1947d:426.

Reference: Kolosvary, 1947e:365.

Distribution: Singapore.

Genus Hiroa Ross and Newman. 1973

Hiroa stubbingsi Ross & Newman. 1973:153
Distribution: Truk, Caroline Is.

Genus Savignium Leach. 1825

Savignium crenatum (Sowerby). 1823:no pagination

Snonymy: Ross & Newman. 1973:159.

Diagnosis: Darwin. 1854b:370.

References: Annandale. 1924:66 (as Pyrgoma crenatum
phase tridacophvlliae nov.); Broch. 1931:120; Edmond-
son, 1951:187; Gruvel, 1905a:304; Hiro, 1935a:14; 1937c:
468; 1938d:399; Kolosvary, 1943a:95; 1947d:427; 1947e:
366; 1951a:287 (as Pyrgoma crenatiformis n. sp.);
Nilsson-CanteU, 1938b:i3; Pilsbry, 1916:262; Utinomi.
1949a:23; Utinomi & Kikuchi. 1966:7; Weltner. 1897:256.

Distribution: Japan; Philippines; Line Is.; Palau Is.;
Singapore; Mergui Arch.
Savignium dentatum (Darwin). 1854b:369

Synonymy/diagnosis: Hiro. 1935a:12.

References: Dawydoff. 1952:128; Gruvel. 1905a:305;
1912a:350; Hiro. 1931:154; 1937c:467; 1938d:400; Kolos-
vary. 1947e:366; Ross & Newman. 1973:159; Weltner.
1897:256.

Distribution: Red Sea; Gulf of Siam; New Guinea; Palau
Is.; Japan.
Savignium elongatum (Hiro). 1931:154

Synonymy/diagnosis: Hiro. 1931:154.

References: Dawydoff. 1952:128; Hiro. 1935a:19; 1937c:
468; 1938d:400; Ross & Newman. 1973:159.

Distribution: Japan; Palau Is.; Gulf of Siam.
Savignium milliporum (Darwin). 1854b:367

Synonymy: Ross & Newman. 1973:159.

Diagnosis: Darwin. 1854b:367.

References: Barnes & Klepal, 1971:87 (pedicel of penis)
Broch. 1931:120; Foster. 1974:49; Gruvel, 1905a:306
Hiro, 1935a:25; 1936a:58; 1938d:401; Hoek, 1913:257
Kolosvary, 1950:292 (as Pyrgoma milleporae forma
typica nov.; as Pyrgoma milleporae forma snelliusi nov.);
Nilsson-Cantell. 1921:355; 1938b:70; Weltner. 1897:256.

Distribution: Indo-west Pacific, east to Fiji and Palau Is.

Genus Creusia Leach, 1817

Creusia decima Ross & Newman, 1973:154

Synonymy/diagnosis: Darwin. 1854b:381 (as Creusia
spinulosa var. 10).

Distribution: Unknown.
Creusia indicum (Annandale). 1924:64

Synonymy/diagnosis: Utinomi. 1967:227.

References: Annandale, 1924:65 (as Pyrgoma indicum

58

phase merulinae nov. and phase symphylliae nov.);
Baluk & Radwariski, 1967b:482; Broch, 1931:118 (as C
spinulosa angustiradiata nov.); Darwin, 1854b:381 (as
Creusia spinulosa var. 11): Hiro. 1935a:7: 1937c;466:
1938d:399: Hoek, 1913:265: Nilsson-CanteU, 1938b:62,63
(as C s. angustiterga Broch \sic\y, Ross & Newman,
1973:154; Utinomi, 1943:16 (juvenile stages); 1962:227;
Utinomi & Kikuchi, 1966:7.

Distribution: Japan; Palau Is.; Kei Is.; Singapore;
Mergui Arch.; to 52m.
Creusia spinulosa Leach, 1818:171

Synonymy: Ross & Newman, 1973:154.

Diagnosis: Darwin, 1854b:380 iasCreusia spinulosa var. 9).

References: Annandale, 1906:143; 1924:64; Gruvel,
1903b:164; 1909b:26: Kolosvary, 1959:197, 198; Ladd,
1959:963 iPaleocreusia devonica — not a barnacle);
Weltner, 1897:255.

Distribution: Recent, unknown. Miocene, Hungary.

Genus Nobia Sowerby, 1839

Nobia conjugatum (Darwin), 1854b:364
Synonymy: Ross & Newman, 1973:155.
Diagnosis; Hiro, 1935a:15.

References: Annandale, 1906:143; Broch, 1922:344;
1947:7; Gruvel, 1905a:306; Hiro, 1931:154; 1937c:468:
Hoek, 1913:264; Kolosvary, 1947d:427; Nilsson-CanteU,
1938b:13: Weltner, 1897:255.
Distribution: Red Sea; Ceylon; Mergui Arch.; Gulf of
Siam; Sulu Arch.; Singapore; Japan.
Nobia grandis Sowerby, 1839:71
Synonymy: Darwin, 1854b:365; Nilsson-CanteU, 1938b:68.
Diagnosis: Darwin, 1854b:365.

References: Annandale, 1924:66; Borradaile, 1903:443;
Barnes & Klepal, 1971:88 (pedicel of penis); Broch,
1931:120; Darwin, 1854b:365 (? = Balanus duptoconus
Lamarck, 1818 = Duplocona laevigata Schliiter, 1838);
Dawydoff, 1952:128; Gruvel, 1905a:307; Hiro, 1931:154;
1935a:16: 1937c:468; 1938d:401; Hoek, 1913:258; Kolos-
vary, 1947d:427; 1947e:366; Korschelt, 1933:26; Nilsson-
CanteU, 1921:357; Ross & Newman, 1973:155; Weltner,
1897:256.
Distribution: Mergui Arch.; Maldives; Singapore; Indo-
nesia; Kei Is.; Gulf of Siam; Palau Is.; Japan.
Nobia halomitrae (Kolosvary), 1947e:363
Synonymy diagnosis- Kolosvary, 1947e:363.
Reference: Ross & Newman, 1973:155.
Distribution: Unknown.
Nobia kuri (Hoek), 1913:259
Synonymy diagnosis: Hoek, 1913:259.
References Hiro, 1931:155; 1935a:25; Ross & Newman,

1973:155.
Distribution: Kei Is.; Banda Sea; 204m.
Nobia orbicellae (Hiro), 1934:367
Synonymy/diagnosis: Nilsson-CanteU, 1938b:73.
References: Hiro, 1935a:17; 1937c:468: 1938d:401: Kolo-
svary, 1943a:96; 1947q:427; Ross & Newman, 1973:155.
Distribution: Japan; Palau Is.; Fiji; Singapore; Mergui
Arch.
Nobia projectum (Nilsson-CanteU), 1938b:70
Synonymy/diagnosis: NUsson-CanteU, 1938b:70.
References: Ross & Newman, 1973:155; Utinomi,

1969a:82.
Distribution: Persian Gulf; 24m.

Genus Pygroma Leach, 1817

Pyrgoma cancellata Leach, 1818:171
Synonymy; Hiro, 1935a:10.
Diagnosis: NUsson-CanteU. 1938b:67.
References: BorradaUe. 1903:443; Darwin, 1854b:362;

Dawydoff, 1952:128; Gruvel, 1905b:303; Hiro, 1937c:467;

1938d;399; Hoek, 1913:257, 264; Kruger, 1911a:4 (var.

japonica); 1911b;461; Ross & Newman, 1973:156;

Utinomi, 1958a:309; 1962:227: Utinomi & Kikuchi,
1966:7; Weltner, 1897:255 (as var. japonica nov.).
Dlstribution: Pacific coast of central to southern Japan:
Philippine Sea; Palau Is.; Gulf of Siam; Mergui Arch.;

Maldives.

Genus Pyrgopsella ZuUo, 1967

Pyrgopsella annandalei (Gruvel), 1906b: 1558

Synonymy: Ross & Newman, 1973:163.

Diagnosis: Gruvel. 1907d:8.

References: ZuUo. 1967a:123 (replacement name for
Pyrgopsis Gruvel).

Distribution: Andaman Is.
Pyrgopsella stellula RoseU. 1973a:5

Distribution: Sulu Arch.

Genus Hoekia Ross and Newman, 1973

Hoekia monticulariae (Gray), 1831:6

Synonymy: Baluk & Radwanski, 1967b:487.

Diagnosis: Ross & Newman, 1969:161.

Reference.s: Annandale, 1924:67; Darwin, 1854b:372:
Gruvel, 1905a:308; Hiro, 1931:155; 1935a:18; 1937c:
468; Hoek, 1913:264; Kolosvary, 1943a:95; 1947d:427
Nilsson-CanteU, 1938b:66; Robertson, 1970:44.

Distribution: Mauritius; Bay of Bengal; Singapore
Japan.

Subfamily Ceratoconchinae n. subfam.

Genus Ceratoconcha Kramberger-Gorjanovic, 1889

Ceratoconcha barbadensis (Withers), 1926:2
Reference: Nilsson-CanteU, 1938b:63: Ross & Newman,

1973:166.
Distribution: Pleistocene, Barbados, West Indies.
Ceratoconcha conicocystata Newman & Ladd, 1974:391
Distribution: Upper Miocene, Dominican RepubUc: Mid-
dle Miocene, Trinidad.
Ceratoconcha costata (Seguenza), 1876:316
Synonymy: Baluk & Radwanski, 1967b:477; Ross & New-
man, 1973:166.
Diagnosis: Seguenza, 1876:316.

References; Bogsch, 1957:25; de Alessandri, 1895:299
(as Pyrgoma costatum): 1906:322; 1910:115 (as Pyrgoma
cf. anglicum): Duvergier, in de Alessandri, 1922:228;
Kolosvary, 1949:1 (as Creusia spinulosa iorma praespin-
ulosa, nov., fig. 5 only); 1962a:86 (as Creusia spinulosa
forma kojumdgievae nov.) 1967b:393; Moroni, 1967b:17;
Prochazka, 1893:20 (as Creusia moravica n.sp.); Withers,
1953:61, 63.
Distribution: Miocene to Pleistocene of Italy; Miocene
of Bulgaria (?) and Hungary.
Ceratoconcha creusioides Newman & Ladd, 1974:392
Distribution: Lower Miocene, Jamaica; Middle Miocene,
Trinidad.
Ceratoconcha darwiniana (Prochazka), 1893:23
Synonymy/diagnosis: Prochazka, 1893:23.
References: Baluk & Radwanski, 1967b:480; Ross &

Newman, 1973:166.
Distribution: Miocene, Austria,
Ceratoconcha diplocona (Seguenza), 1876:322
Synonymy/diagnosis: Seguenza, 1876:322.
Reference: Ross & Newman, 1973:166; Withers, 1953:61.
Distribution: Phocene, Italy.
Ceratoconcha domingensis (Des MouUns), 1866:307
Synonymy/diagnosis: ZuUo et al, 1972:71.
Reference: Ross & Newman, 1973:166.
Distribution: Haiti; Dry Tortugas; Florida; Bermuda.
Ceratoconcha floridana (PUsbry), 1931:81
Synonymy/diagnosis: PUsbry, 1931:81.
References: Henry, 1954:444; Hiro, 1935a:25; Kolosvdry,

59

1951b:294: Ross & Newman, 1973:166; WeUs. 1966:86.
Distribution: West coast of Florida.
Ceratoconcha jungi Newman & Ladd, 1974:395

Distribution: Lower Miocene, Jamaica.
Ceratoconcha krambergeri (Baluk & Radwanski), 1967a:145
Synonymy/diagnosis: Baluk & Radwanski, 1967a:145.
References: Abel, 1920:fig. 136; 1928:13; 1935:535; Baluk
& Radwanski, 1967b:480; Bogsch, 1957:29; Brooks &
Ross. 1960:362; Dacque, 1921:fig. 91b; Kolosvary,
1949:111; Kramberger-Gorjanovic, 1889a:50 (as Cerato-
concha costata n. sp.); 1889b:231; 1889c:142: Prochazka,
1893:19; Ross & Newman, 1973:166; Stromer, 1912:fig.
232d; Termier, 1953:fig. 19; Withers, 1926:5.
Distribution: Miocene, Yugoslavia.
Ceratoconcha minuta Newman & Ladd, 1974:394

Distribution: Middle Miocene, Trinidad.
Ceratoconcha miocaenica (Prochazka), 1893:22
Synonymy/diagnosis: Baluk & Radwanski, 1967a:138.
References: Baluk & Radwanski, 1967b:479; de Alessan-
dri, 1910:125 (as Pyrgoma cf. anglicum). Ross & New-
man, 1973:167.
Distribution: Miocene, Austria and Yogoslavia.
Ceratoconcha noszkyi {Kolosvary), 1949:114.
References: Baluk & Radwanski, 1967b:476; Kolosvary,
1949:114 (as Andromacheia noszkyi n. sp.); 1951b:295;
Ross & Newman, 1973:167.
Distribution: Miocene, Hungary.
Ceratoconcha prefloridana (Brooks & Ross), 1960:355
References: Baluk & Radwanski, 1967b:484; Weisbord,

1972:60 (as Creusia neogenica n. sp.).
Distribution: Pliocene, Florida.
Ceratoconcha quadratoradiata Newman & Ladd, 1974:393

Distribution: Middle Miocene, Trinidad.
Ceratoconcha quarta (Kolosvary), 1947d:427
Synonymy: Ross & Newman, 1973:167.
Diagnosis: Darwin, 1854b:378(asCreusiaspinu/osa var. 4).
Reference: Utinomi, 1949a:35; 1962:231.
Distribution: West Indies.
Ceratoconcha rangi rangi (Des Moulins), 1866:302
Synonymy: Ross & Newman, 1973:167.
Diagnosis: Des Mouhns, 1866:302.

References: Kolosvary, 1949:111 (as Creusia spinulosa
forma praespinulosa n. f., figs 2, 3 only; and Creusia
spinulosa forma cladangiae n.f., both from Hungarian
Miocene); 1962a:86; 1967b:393; Prochazka, 1893:18 (as
Creusia fuchsi n. sp.); Seguenza, 1873:319 (as Pyrgoma
multicostatum n. sp.); Withers, 1953:57, 58, 68.
Distribution: Miocene, France, Hungary and Bulgaria.
Ceratoconcha rangi latum (Seguenza), 1876:321
Reference: Ross & Newman. 1973:167.
Distribution: Miocene, Italy.
Cera«oconc/iasanctacracens!s(Baluk&Radwanski),1967b:468
Synonymy: Ross & Newman, 1973:167.
Diagnosis/reference: Baluk & Radwanski, 1967b:468 (as

Creusia sanctacrucensis n. sp.).
Distribution: Miocene, Poland.
Ceratoconcha sturi (Prochazka), 1893:15
Reference: Baluk & Radwanski, 1967b:479; Ross & New-
man. 1973:167.
Distribution: Miocene, Czechoslovakia.
Ceratoconcha trolli (Abel), 1927:101
Reference: Abel, 1928:13.
Distribution: Miocene, Austria.

Subfamily Bosciinae n. subfam.

Genus Boscia Ferussac, 1822

Boscia anglica (Sowerby), 1823:no pagination
Synonymy: Darwin, 1854b:360.
Diagnosis: Broch, 1927:30.

References: Baluk & Radwanski, 1967c:693; Barnes &
Klepal, 1971:88 (pedicel of penis); Bassindale, 1964:43;

Crisp & Southward, 1961:271 (cirral activity); Darwin,
1854a:36; de Alessandri, 1895:297; 1906:320; Fischer,
1872:433; Gauld, 1957:10; Gruvel, 1905a:302; Hiro,
1935a:9; 1937c:467; Hoek, 1875:60; 1909:271; 1913:257;
Holdsworth, 1860:7111; Kruger, 1940:460; Le Reste.
1965:66 (larvae); Moyse, 1960:120 (rearing); 1961:371
(larval stages); 1971:125(settlementand growth); Nilsson-
CanteU, 1938b:66; O'Riordan, 1967:294; Pilsbry, 1916:
292; Rees. 1962:411; Relini. 1969:177; Ross c& Newman,
1973:164 (= Pyrgoma stokesii Gray , 1825:103);Seguenza,
1876:314; Stubbings, 1964a:lll; 1967:294; Utinomi,
1958a:309; Weltner, 1897:255; 1898b:ll; Withers, 1953:
39 et seq.

Distribution: England and Ireland; France; Sicily;
Madeira; Cape Verde Is.; West Africa; sublittoral to
450m. Pliocene, England; Plio- Pleistocene. Italy, Malta.
Boscia madreporarae (Bosc), 1812:66

Synonymy: Ross & Newman, 1973:164.

Diagnosis: Darwin, 1854b:361.

References: Gruvel, 1905a:303; 1912a:350; Hiro, 1935a:
25; Kruger, 1940:382; Pilsbry, 1916:262; Southward
1975:18; Utinomi, 1967:232.

Distribution; West Indies.
Boscia oulastreae (Utinomi), 1962:227

Synonymy: Utinomi, 1967:229.

Diagnosis: Utinomi, 1962:227.

References: Utinomi, 1949a:35 (as Creusia spinulosa
forma quarta): 1967:229 (as Megatrema oulastrea)
Utinomi & Kikuchi, 1966:8; Ross & Newman, 1973:164;
Sakakura, 1934:578.

Distribution: Tanabe Bay, Japan; Pleistocene, Japan.
Boscia seguemai (Baluk & Radwanski), 1967c:691

Reference: Baluk & Radwanski, 1967c:691 {as Pyrgomina
seguenzai): Ross & Newman, 1973:164.

Distribution: Phocene, Crete.

Family Balanidae Leach, 1817

Genus Balanus Da Costa, 1778
Group ot Balanus balanus

Balanus balanus (Linnaeus), 1758:667
Synonymy/diagnosis: Pilsbry, 1916:149 (= Balanus porca-
tus da Costa, 1778:249; includes pre-Darwinian refer-
ences).
References: Aurivillius, 1898a:30; Ballowitz, 1908:421
(sperm); Barnes, 1953a:141 (orientation and aggregation);
1953c:128 (effect of parasitism); 1955a:114 (hatching);
1959a:232 (stomach contents); 1962a:353 (oxygen uptake
and metabolism); 1963b:587 (seminal plasma); 1965:321
(biochemistry of eggs); Barnes & Barnes, 1954:63 (bio-
logy); 1965a:391 (variation in egg size); 1968a:135 (varia-
tion in egg production); 1969b:36 (seasonal changes in
oxygen consumption); 1974:197 (embryonic development
and salinity); Barnes & Blackstock, 1974:35 (constituents
of body fluids); 1974b:47 (composition of seminal plas-
ma): Barnes & Costlow, 1961:59 (larval stages); Barnes
& Dawson, 1966:263 (lipids); Barnes & Finlayson, 1962:
98 (ascorbic acid in semen); 1963:185 (seasonal changes);
Barnes & Healy. 1969:51 (biometrical studies); Barnes et
al. 1970:70 (effect of impaction); 1971:173 (spermatozoa);
Bassindale. 1964:39; Belyaev, 1949:902; Bertelsen, 1937:
38 (as B. crenatus according to Stephensen, 1943);
Bousfield, 1955b:766; Brattstrom, 1957:12; Brocchi,
1814:598; Broch. 1924a:73; 1927b:21; 1936:3 (as B. b.
artica); Chilton. 1909:670 (as B. porcatus, Auckland Is.
and Australia); 1920:53 (as B. porcatus); Cornwall,
1955a:32; 1955b:25 (= B. b. pugetensis Pilsbry, 1916:
163); Crisp, 1954: 473 (breeding); 1964a: 193 (effect of
severe winter); Crisp & Patel, 1969:284 (control of breed-
ing); Crisp & Southward, 1961:271 (cirral activity); Crisp
& Spencer, 1958:290 (control of hatching); Darwin,
1854a:21; 1854b:256; Davadie, 1963:68; Dawson &
Barnes, 1966:249 (biochemistry of eggs); de Alessandri,

60

1895:290; 1906:304; Filhol, 1885:487; Foster, 1970:390
(accUmation to saUnity); Gruvel. 1903b:137; 1905a:237;
1920:54 (as B. crenatus according to Stephensen, 1943);
Gutman, 1961:171 (colonization); Henry, 1942:101 [as B.
b. pugetensis Pilsbry); Hiro, 1935c:227; Hoek, 1875:60;
1909:271; Hutton, 1879:328; Jennings, 1918:61; Kolos-
vary, 1943a:89; 1967b:391; Korschelt, 1933:23; Kruger,
1911a:3; 1911b:460; 1927a:14; 1927b:5; Linnaeus, 1767:
1107; Menesini, 1966:124; Moore, 1934a:101 (growth
rate); Morch, 1852:68; Munn & Barnes, 1970a:277 (sper-
matozoa); 1970b:261 (spermatozoa); Nilsson-Cantell,
1931a:113; O'Riordan, 1967:293; Patel & Crisp, 1960b:
104 (rate of embryonic development); 1961:89 (breeding
and molting); Pilsbry, 1916:11 (Protobalanus, not a
barnacle); 1916:149; Poulsen, 1936:13; Remy, 1928:231
(as B. crenatus according to Stephensen, 1943); Ruede-
mann, 1918:382 {Eobalanus. not a barnacle); Rzhepishev-
skii, 1968:38; Schafer, 1952:238 (settling); Sommer,
1972a:271 (motor activity); 1972b:177 (periodicity);
1972c:1449 (mechanisms of pressure sensitivity); 1972d:
352 (physiology of pressure perception); Southward,
1957:327 (behavior); 1965:442 (metabolism and survival
at high temperatures); Southward & Crisp, 1963:31 (foul-
ing); Stephensen, 1938:4; 1943:18; Sumner, 1911:128;
Tarasov, 1932:60; 1936:46; 1937:40; Tarasov & Zevina,
1957:194; Visscher, 1928b:193 (fouling); Weltner, 1897:
267; 1898b:12; 1900:303; Zevina & Tarasov, 1964:239;
ZuUo, 1963b:10; 1968:6; 1969b:351.
Distribution: North Atlantic and North Pacific; low water
to 180m. PUocene; England and Italy; Pleistocene: Ore-
gon, Maine, Canada, Sweden, and Iceland.
Batanus crenatus Brugiere, 1789:168

Synonymy/diagnosis Cornwall, 1925:476; Pilsbry, 1916:

165 (includes pre-Darwin references not listed below).
References Abel, 1926:250; Addicott, 1966:04; Aurivil-
lius, 1898b:30; Austin et al, 1958:497 (chromosome num-
ber); Barnard, 1924:70; Barnes, 1950:74 (larvae); 1952-53:
104 (effect of light); 1953b:328 (effect of lowered saUnity);
1953e:297 (variations in cyprids); 1959a:231 (stomach
contents); Barnes & Bagenal, 1951b:369 (on Nephrops
norvegicus): Barnes & Barnes, 1965a:391 (egg/naupUus
size variation); 1974:194 (embryonic development and
salinity); Barnes & Crisp, 1956:631 (self-fertilization);
Barnes & Healy, 1969:51 (biometrical studies); Barnes &
Klepal, 1971:83 (pedicel of penis); Barnes & Powell,
1950a:175 (development, morphology); 1953a:107 (growth
under submersion); Barnes et al, 1951:227 (orientation);
1963:233 (effect of dessication); 1970 (behavior or impac-
tion); 1971:173 (spermatozoa); Bassindale, 1964:38;
Belyaev, 1949:902 (osmoregulation); Bishop etal, 1957:6;
Blom & Nyholm, 1961:153 (settling times); Bocquet-
Vedrine, 1970a:506 (cement glands); 1970b;963 (cement
glands); 1970c:521 (cement glands); Bohart, 1929:353
(attachment of cyprids); Bousfield, 1954:119; 1955a:19;
1955b;764; Brattstrom, 1957:10; Broch, 1922:321; 1924:
78; 1927b:22; 1936:4; Chilton, 1920:53; Ciurea et al,
1933:6; Cornwall, 1925:476; 1951:329; 1955a:25; 1955b:
28; Crisp, 1955:569 (behavior of cyprids); 1964a:181
(effects of severe winter); Crisp & Barnes, 1954:142
(orientation and settlement); Crisp & Patel, 1958:1078
(breeding and ecdysis); 1969:283 (environmental control
of breeding); Crisp & Southward, 1961:271 (cirral activ-
ity); Crisp & Stubbings, 1957:179 (orientation to water
currents); Darwin, 1854a:23; 1854b:261; Davadie, 1963:
63; de Alessandri, 1906:305; 1907b:284; Ellis & Solander,
1786:198 (as Balanus clavatus): Fischer-Piette, 1930:41;
1932:8; Fischer-Piette & Prenant, 1956:12; Foster, 1969:
326 (temperature tolerance); 1970:386 (salinity); 1971a:12
(dessication); Gruvel, 1903b: 139; 1905a;240; 1909b:25;
Henry, 1942:105; 1954:443; Herz. 1933:432 (morphology
of later stages); Hiro, 1935c:219; Hoek. 1875:35; 1884:
517; 1909:270; Jennings, 1918:61; Kauri, 1962:131
(nauplius eye); 1966:115 (X-organ); Knight-Jones,
1955:266; Knight-Jones & Crisp, 1953:1109 (gregarious-

ness); Kolosvary, 1943a:89; 1951c:411; 1956:187; 1959:
197; 1962a:85; 1963a:174; 1967b:392; Kruger, 1911a:52;
1911b:460; 1927:17; 1940:464; Kuhl, 1963:99; 1965:121;
1967:967 (ecology in Elbe estuary); Lecointre, 1910:139;
Meadows, 1969a:278 (fouUng); 1969b:65 (settlement,
growth and competition); Moore, 1934a:101; 1936:703;
Moyse, 1963:175; Muller, 1940:113 (sensitivity to poi-
sons); Nilsson-Cantell, 1921:326 1931a:113; O'Riordan,
1967:291; Patel & Crisp, 1960b:104 (embryo develop-
ment); 1961:89 (relationship between breeding and
moulting); Pilsbry, 1911a:75; 1921:112; Poulsen, 1935:21;
Prenant & Teissier, 1923:176; Pyefinch, 1948a:451;
1948b:464; 1948c:916 (larvae); Schafer, 1938b:323
(paleontology); 1952:242 (settling); Schwarz, 1932:437
(influence of light); Sneli, 1972:3; Southward, 1955b;403
(relation of activities to temperature); 1965:443 (metabo-
lism and survival); Southward & Crisp, 1963:36; 1965:
161 (activity rhythms); Stebbing, 1910:569; Stephensen,
1938:5; 1943:19; Stubbings, 1961b:33; Tarasov, 1936:48;
1937:50; Tarasov & Zevina, 1957:205; Trusheim, 1932:70
(paleontology); Utinomi, 1970:358; Walker, 1972:429
(chemical composition of cement); Weltner, 1897:268;
1898a:442; 1898b:ll; 1900:298; Withers, 1953:58, 61, 70;
Zevina & Tarasov, 1964:239; Zullo. 1963b:10; 1969b;315.
Distribution: North Pacific, south to Santa Barbara;
Arctic; North Atlantic south to Florida. Unverified
localities: S. Africa, Australia, West Indies, Peru, south-
ern China; intertidal to 250m. OUgocene to Pliocene,
Mediterranean Basin; Pleistocene, North America.
Balanus crenatus curviscutum Pilsbry, 1916:175
Synonymy/diagnosis: Pilsbry, 1916:175.
References: Henry, 1942:126; Hiro, 1935c:221; Utinomi,

1958a:308.
Distribution: North Pacific; Japan to Northwest America.
Balanus crenatus delicatus Pilsbry, 1916:177
Reference: Henry, 1942:126.
Distribution: Humboldt Bay, California.
Balanus glandula Darwin, 1854b:265
Synonymy: Pilsbry, 1916:178.
Diagnosis: Cornwall, 1925:438.

References: Augenfeld, 1967:92 (respiration); Barnes &
Barnes, 1956a:415 (biology); 1959h:515 (metabolism);
1965a:391 (variation in egg and nauplius size); Barnes &
Conor, 1958:194 (neurosecretory cells); Barnes & Healy,
1969:62 (biometrical studies); Barnes & Klepal, 1971:83
(pedicel of penis); Batzli, 1969:535 (distribution of bio-
mass); Bergen, 1968:229; Broch, 1922:321; ConneU,
1970:49 (predation by Thais): Cornwall, 1951:326; 1955a:
27; 1955b:33; Dayton, 1971:351 (competition); Glynn,
1965:109 {Endocladia-Balanus associations); Gruvel,
1905a:238; Haven, 1973:97; Henry, 1942:108; Johnson
& Miller, 1935:12 (settlement); Kolosvary, 1943a:91;
Newman, 1967:1038 (biology); Nilsson-Cantell, 1921:326;
Pilsbry, 1907d:201; Rice, 1930:249 (distribution in com-
munities); Tarasov & Zevina, 1957:202; Weltner, 1897:
269; 1898b:7; Worley, 1939:233 (correlation between
salinity, size and abundance), Zullo, 1969b:351.
Distribution: Aleutians to Baja California; Rio de Janeiro
(Spivak, in litt.). Pleistocene, Oregon.
Balanus withersi Pilsbry, 1930:429
Distribution: Miocene, New Jersey.

Group of Balanus nubilus

Balanus connelli Cornwall, 1927b:402

Dustrubution: Miocene, British Columbia.
Balanus nubilus Darwin, 1854b:253

Synonymy/diagnosis: Ross, 1962:24; Henry, 1942:112.

References: Abel, 1926:246; Addicott, 1966c:4; Arvy and
Lacombe, 1968:1326 (cement apparatus); Arvy & Ligouri,
1968:817 (muscular cytochrome oxidase activity); Arvy
et al, 1968:817 (alkaline phosphatase activity); Barnes,
1959a:234 (stomach contents); 1959b:607 (note on spel-
ling of nubilus); Barnes & Barnes, 1959b:19 (stimulation

61

of nauplii); 1959c:15 (naupliar stages); 1965a:391 (varia-
tion in egg, nauplius size); Barnes & Conor, 1958:194
(neurosecretory cells); CardereUi, 1968:1 (barnacle ce-
ment); CornwaU, 1925:479; 1927b:408; 1936:471 (as
Balanus altissimus); 1951:334 (as Balanus flos), 335;
1953:78, 80; 1955a:23; 1955b:36; 1958:81; 1959:404;
Emerson & Hertlein, 1960:7; Fitzgerald, 1968:1055 (cal-
cium and pH dependency); Gruvel, 1903b:130; 1905a:226;
Hagiwara & Nakajima, 1966:807 (effects of Ca ion con-
centration); Hagiwara & Takahashi, 1967:583 (surface
density of calcium in muscle fiber); Hagiwara et al,
1968:773 (effects of pH changes); Harnden, 1968:303
(digestive carbohydrates); Hatai, 1938:96; Henry, 1940
29; Hoyle & Smythe, 1963:49 (giant muscle fibers)
Hughes, 1914:213; Kaminer & Kimura, 1972:406 (cal
cium release in muscle); Kolosvary, 1943a:89; 1959:197
Kruger, 1940:464; Lacombe, 1970:164 (cement glands)
Nilsson-Cantell, 1931a:112; Pilsbry, 1907d:201 (as
Balanus flos n. sp.); 1916:131, 135; 1921:112; Shelford
et al, 1935:281; Tait & Emmons, 1925:42 (movement of
operculum); Whitney, 1970:229 (sterol biosynthesis);
ZuUo, 1969a:8; 1969b:351.

Distribution: Southern Alaska to San Quintin, Baja
California. Oligocene, Vancouver Is., B.C.; Miocene,
Japan and Hungary; Plio-Pleistocene, southern and
Baja California; Pleistocene. Oregon.
Balanus rostratus Hoek, 1883:152

Synonymy: Cornwall, 1955b:38.

Diagnosis Pilsbry, 1916:138 (as B. r. rostratus), 141 (as
B. rostratus alaskensis n. subsp.), 142 (as B. rostratus
heteropus n. subsp.), 147 (as B. rostratus dalli n. subsp.).

References; Addicott, 1966:C4; Barnes, 1959a:233;
Barnes & Barnes, 1959h:515; Barnes &. Conor, 1958:194
(neurosecretory cells); Barnes & Klepal, 1971:83 (pedicel
of penis); Broch. 1922:320; Cornwall. 1925:484; 1955a:29:
Gruvel, 1905a:239; Hatai, 1938:97; Henry, 1940:21
1942:117, 127 (as apertus and dalli); Hiro, 1932a:550:
1933:71; 1935c:217, 218 (as dalli), 227 (as apertus)
19391:210, 211 (as dalli); Kolosvary, 1943a:89 (as dalli)
1961a:78; 1962b:210; 1962d:202; 1967b:392; Kruger
1911a:52; 1911b:463 (as apertus); 1940:464; Nilsson
Cantell, 1932a:20 (as spiniferus); Pilsbry, 1911a:73, 74
(as B. r. apertus n. subsp.); 1916:144 (as apertus), 147 (as
dalli), 148 (as B. rostratus dalli form suturalis); Tarasov
& Zevina, 1957:199. 200, 201 (as apertus), 202 (as dalli)
Utinomi, 1958a:294, 295 (as apertus); 1969b:51; 1970:357
Weltner, 1897:269; 1900:296; Yamaguchi, 1971:122
ZuUo, 1969b:351 {as B. rostratus apertus).

Distribution: Japan; Siberia; Bering Sea; Alaska and
Puget Sound; 0-128m. Miocene. USSR and Japan;
Pleistocene, Central California, Oregon and Japan.
Balanus tamiamiensis Ross, 1964b:272

Distribution: Miocene, Florida.

Group of Balanus concavus

Balanus aquila Pilsbry, 1907a: 199

Synonymy/diagnosis: Pilsbry, 1916:127.

References; Baskin et al, 1969:471 (filaments from
myosin); CornwaU, 1951:333; 1960:831; Henry, 1942:100;
ZuUo, 1966c: 141.

Distribution: San Francisco to San Diego, CaUfornia;
intertidal to 18m.
Balanus bloxhamensis Weisbord, 1966:48

Distribution: Miocene, Florida.
Balanus concavus concavus Bronn, 1831:127

Synonymy/diagnosis: Menesini, 1965:110; Utinomi,
1969a:83.

References: Arnold, 1907a:543; 1907b:422; Beal, 1948:64
Darwin, 1854a:17; 1854b:235; Davadie, 1963:52; Davadie
Suaudeau, 1952:17; de Alessandri, 1895:282; 1906:295
1907b:280; DeLong, 1941:243; Emerson & Hertlein
1960:7; Gruvel, 1903b:136; 1905a:232; Kolosvary, 1943a
85; 1955:183; 1959:197; 1960:590; 1961a:78; 1961b:99
1961c:150; 1962b:206; 1962d:202; 1967b:391; Menesini
1963:5; 1966:116; 1967b:219; 1968b:580; 1972:40;NUsson

CanteU, 1939a:6; Nomland. 1917:301; Pilsbry, 1916:100;
Ross, 1962:14; 1964a:489; Sequenza, 1876:296; Utinomi,
1969a:83 (includes Balanus concavus sinensis Broch,
1931:63 and B. c. indicus Nilsson-CanteU. 1932b:2);
Weaver, 1949:104; Weisbord, 1966:32; Weltner, 1897:
264; ZuUo, 1969a:6.
Distribution: China; India; Persian Gulf. OUgocene-
Pleistocene. Mediterranean Basin; Miocene, Eastern
United States and Britain; PUocene, Venezuela.
Balanus concavus alloplax Pilsbry & Olson. 1951:200

Distribution: Oligocene, Ecuador.
Balanus concavus chesapeakensis PUsbry, 1916:103
Synonymy/diagnosis: Pilsbry, 1916:103.
References: Kolosvary, 1943a:85; Martin, 1904:94.
Distribution: Miocene, Maryland.
Balanus concavus coosensis DaU. 1909:138
Synonymy/diagnosis: Pilsbry, 1916:108 (= B. tintinnabu-

lum coosensis DaU).
Distribution: Miocene, Coos Bay, Oregon.
Balanus concavus dallonii Davadie-Suaudeau, 1952:20

Distribution: PUocene. Algeria.
Balanus concavus eseptatus Pilsbry, 1924:1

Distribution: Miocene, Haiti.
Balanus concavus finchii Lea 1833:211
Synonymy: Pilsbry. 1930:432.
Distribution: Miocene. Maryland.
Balanus concavus glyptopoma Pilsbry, 1916:102
Reference: Cones, 1968:61; Kolosvary, 1943a:85. Pilsbry,

1918:185.
Distribution: Miocene, eastern United States. PUocene,
Panama and east Mexico.
Balanus concavus mexicanus Henry, 1941:100
References: Henry, 1942:126; 1960:141.
Distribution: West coast of Baja CaUfornia to Mazatlan,
Mexico.
Balanus concavus oligoseptatus Kolosvary, 1961c:149

Distribution: Upper OUgocene, U.S.S.R.
Balanus concavus proteus Conrad, 1834:134
Synonymy/diagnosis: Ross, 1964a:486.
References: Davadie, 1963:53, Kolosvary, 1943a:86;

PUsbry, 1916:103.
Distribution: Mio- PUocene, eastern United States.
Balanus concavus raphanoides Moroni- Ruggieri, 1952:71

Distribution: PUocene, Italy.
Balanus concavus rariseptatus Pilsbry, 1918:186

Distribution: Miocene, Panama.
Balanus concavus rubescens Seguenza, 1876:450

Distribution: Tertiary, Italy.
Balanus concavus scutorum Seguenza, 1876:74
Synonymy/diagnosis: Moroni-Ruggieri, 1952:67.
Reference: de Alessandri, 1906:292 (as Balanus spongi-

cola).
Distribution: PUocene, Italy.
Balanus concavus sinensis Broch (see B. c. concavus)
Balanus eyerdami Henry, 1960:139
Reference: Ross, 1962:17.
Distribution: Gulf of CaUfornia; 0-85m.
Balanus gregarius (Conrad), 1856:315
Synonymy/references: ZuUo, 1969a:6 (includes Tamio-
soma gregaria, Radiolites gregaria, Balanus estrellanus
and Balanus concavus concavus, Ross, 1962:14).
Distribution: Mio- PUocene, Central and Southern CaUfor-
nia; PUocene, Baja CaUfornia.
Balanus indicus Withers, 1923:291

Distribution: Miocene, Pakistan.
Balanus polyporus PUsbry, 1924:2

Distribution: Miocene, Haiti.
Balanus regalis PUsbry, 1916:108
Synonymy/diagnosis: Ross, 1962:19.
References: CornwaU. in Steinbeck & Ricketts, 1941:430;
CornwaU, 1959:403; Henry, 1942:100; 1943:368; 1960:144
(as subsp. of B. aquila); Kolosvary, 1942c:139; 1943a:85.
Distribution: Southern and Baja CaUfornia.
Balanus talquinensis Weisbord, 1966:37
Distribution: Miocene, Florida.

62

Balanus vadaszi Kolosvary, 1949a:2
Reference: Davadie, 1963:57; Menesini, 1972:42.
Distribution: Miocene, Europe.

Group of Balanus amphitrite

Balanus albicostatus albicostatus Pilsbry, 1916:90
Synonymy/diagnosis Utinomi. 1967:209.
Rerences Gruvel, 1903b:133 (as Balanus violaceus);
1905a:227; Hirano, 1953:1 (rearing); Hirano & Okushi.
1952:639 (attachment and growth rate); Hiro. 1937c:432;
1938a;303; 1938c:1687 (resistance to exposure); 1939a:
128; 1939e:261; 1939f:209; Hudinaga & Kasahara, 1942:
108; Ishida & Yasgui, 1937:1659 (free-swimming stages);
Kawahara. 1961:70 (fouUng); Kawahara & lizima,
1960-584 (fouUng); Kolosvary, 1943a:84; 1951c:411;
1961b:100; 1961c:150; 1962c:202; 1967b:391; Kruger,
1911a:51 (as Balanus amphitrite communis); Mawatan,
1967-99 (fouhng); Mawatari et al. 1962:93; 1963:95
(water conduit fouling); Nilsson-CanteU, 1921:314; Nishi-
kawa, 1960:355 (chromosomes); Ooishi, 1964:195; Pils-
bry 1916:90 (as Balanus amphitrite albicostatus): Stub-
bings, 1967:277; Tarasov & Zevina, 1957:183; Utinomi,
1949a:22; 1955b:124 (geology); 1958a:308; 1958b:51;
1962:216; 1969b:52; 1970:356; Utinomi &Kikuchi, 1966:5;
Zevina & Tarasov, 1963:91.
Distribution: Japan; China; Formosa; Korea. Miocene;
Turkistan, U.S.S.R.
Balanus albicostatus formosanus Hiro, 1938a:306
Synonymy/diagnosis: Utinomi, 1967:212.
References: Hiro, 1939e:261; Kolosvary, 1961b:100;

1962:d:199; 1967b:391.
Distribution: Formosa: Miocene, U.S.S.R.
Balanus amphitrite abundantus Kolosvary, 1948:106

Distribution: Miocene, Hungary.
Balanus amphitrite amphitrite Darwin, 1854b:240
Synonymy: Utinomi, 1967:200; Southward, 1975:6.
Diagnosis: Harding, 1962:274; Stubbings, 1967:271. (In-
cludes communis Darwin, denticulata Broch, hawaiien-
sis Broch, carenatus Gruvel, franciscanus Rodgers,
herzi Rodgers, saltonensis Rodgers).
References: Annandale, 1907:40; 1911:170 (growth rate);
1915:138; BaU, 1937:534; 1950:283; Barnard, 1924:69;
Barnes & Barnes, 1959f:438 (osciUatory respiration);
1965a:392 (variation in egg/nauplius size); 1968a:146
(variation in egg numbers); Barnes et al, 1970:70 (impac-
tion); 1971:173 (spermatozoa); Bassindale, 1964:40; Bhatt
& Bal, 1960:439; Bishop, 1950:409; Bishop et al, 1957:8;
Bookhout & Costlow. 1959:212 (feeding, molting,
growth); Borghouts-Biersteker, 1969:4; Borradaile,
1903:441; Broch, 1916:5; 1922:314; 1924b:203; 1927d:133;
1931:58; 1935:2; CaUame, 1965:413 (effect of light);
Ciurea et al, 1963:6; Costlow & Bookhout, 1956:107
(as B. a. niveus — moulting and shell growth); 1958a:284
(larval development); 1958b:271 (moulting and respira-
tion); Crisp & Costlow, 1963:22 (embryo tolerance to
salinity and temperature); Crisp & Molesworth, 1951:489;
Crisp & Southward, 1961:271; Daniel, 1955c:20; Darwin,
1854b:493 (= B. pic'us Miinster); Davadie, 1963:44;
Davis et al, 1973:310 \interval cycle); Day & Morgans,
1956:303; de Alessandri, 1895:286; 1906:301; 1907b:282;
deOUveira, 1941:17; 1947:733; DePabna, 1963:15; Ed-
mondson & Ingram, 1939:257; Fahrenbach, 1965:233
(photoreceptors); Filhol, 1885:486; Fischer, 1929:10;
Fischer, 1872:432; Fischer-Piette & Prenant, 1956:15;
Fishelson, 1971:128; Foster, 1967a:83; Freiberger &
Cologer, 1966:881 (rearing in laboratory); Gordon, 1970:
69; Graham & Gay, 1945:381 (attachment and growth);
Gruvel, 1903b:137; 1905a:232; 1907d:6; 1912a:346;
Henry. 1958:222; 1959:192; 1960:142; 1973:968; Hirano.
1953:139 (rearing); 1962:77 (rearing); Hirano & Okushi,
1952:639 (seasonal variation in attachment and growth
rate)- Hiro, 1933:71; 1936a:59 (commensalism); 1936b:
624; 1937c:432; 1938a:301; 1938c:1687; 1939c:590;
1939e:263; 1939f:208; Hoek, 1913:172; Hudinaga &

Kasahara, 1942:108 (rearing); Karande, 1967:1245 (foul-
ingl- 1973:56 (larvae); 1974b:229 (larval comparison with
B. uariegatus): Karande & Palekar, 1966:143: Karande &
Thomas, 1971:109 (laboratory rearing); Kawahara, 1961:
67 (fouUng); Kawahara & lizima, 1960:584 (fouling);
Kolosvary 1939b:129; 19411:173 (as B. pictus): 1943a:83;
1943C-129- 1944:33; 1947c:424; 1947d;425; 1951b:292;
1959-197; 1961a;78; 1961b;100; 1961c;150; 1962b:205;
1962d:202; 1963a:173; 1963b:175; 1967b:391; Kruger,
1911a:51; 1911b:460; 1940:464; LaCombe, 1970:164 (ce-
ment glands); LaCombe & Monteiro. 1974:633; Lan-
chester, 1902:369; LeReste, 1965:65; Mawatari, 1970:80
(fouUng); Mawatari et al, 1954a:46 (settlement and
growth); 1954b:48 (settlement and growth); 1962:91
(fouUng); 1963:93 (fouUng); 1968:24 (propagation by
ships)- Mawatari & Kitamura, 1970:67 (fouUng); Mawa-
tari & Kobayashi, 1954a:37 (fouUng); 1954b;l (fouUng);
Menesini, 1965:102; Millard, 1950:266; MiUard & Broek-
huysen, 1970:299; Monod, 1937:15; Moore & Frue,
1959:431; Moore, 1944:333; Moyse, 1960:120; Newman,
1967-1038 (biology); Newman et al, 1967:170; Nilsson-
CanteU, 1921:311; 1930b:10; 1931a;122; 1934b:32; 1938b:
36- 1949:43; Norris et al, 1951:444 (variabiUty in larval
stages); Patel & Crisp, 1960a:667 (influence of tempera-
ture); PiUai, 1958:117 (development); Pilsbry, 1907c:190;
1916:89 (= B. carenatus Gruvel, 1907d;6); 1928:312;
Pope 1945:362; Por. 1972:112; Por & Ferber, 1972:151;
Prenant 1929:212; Prenant & Teissier, 1923:170; ReUni,
1962:3; 1964:408; 1966:179; 1968b:186; 1969:169; ReUm
& Giordano, 1969:251 (vertical distribution); Riedl,
1963-257- Ritz & Foster, 1968:545 (temperature re-
sponses); Rogers, 1949:5; RoseU, 1973b:82 (as B. a.
hawaiensis): Ross, 1962:12; Sandison, 1954:86; Seguenza,
1876:300; Shatoury, 1958:790; Southward, 1957:323
(influence of temperature); 1962:163 (influence of tem-
perature); Southward & Crisp, 1963:27 (fouUng); Steb-
bing 1910:568; Stubbings, 1936:41; 1961a:173; 1961b;22;
1963b-14; 1965:886; Suzuki & Konno, 1970:9 (fouUng);
Tarasov & Zevina, 1957:179; Utinomi, 1949a:22; 1950:63;
1955b- 125; 1958a:308; 1960:43 (synonymy of hawaiensis
and denticulata): 1962:215; 1969a:86; 1969b:52; 1970a:
355- Utinomi & Kikuchi, 1966a:5; Visscher. 1928a:327;
1928b:193 (fouUng); Visscher & Luce; 1928:336 (reaction
to Ught); Weiss, 1947a:56 (tolerance to copper and mer-
cury)- 1948:116 (abnormal growth); WeUs, 1966:83;
WeUs et al, 1964:567; Weltner, 1887:102: 1897:264;
1910-528- Wisely & BUck, 1964:163 (seasonal abundance
of larvae); Withers, 1923:290; 1924:32; Yasuda, 1968:27
(fouUng); Zevina, 1963:73; Zevina & Litvinova, 1970:173;
ZuUo, 1963b;8, 9 ( B. amphitrite subsp. n. = B. improvi-
sus Darwin according to Henry, 1973:968).
Distribution: CosmopoUtan. in warm and temperate seas.
Numerous FossUs attributed to this species; OUgocene
to Pleistocene.
Balanus amphitrite acutus Withers. 1924:30
Distribution: Miocene, New Zealand (Withers, 1953:77
et seq.).
Balanus amphitrite aeratus deOUveira. 1941:22

Distribution: Rio de Janeiro.
Balanus amphitrite archi-inexpectatus Kolosvary, 1948:108

Distribution: Miocene, Hungary.
Balanus amphitrite cochinensis Nilsson-Cantell, 1938b:43
References: Karande, 1966:144; 1967:1245.
Distribution: Bombay.
Balanus amphitrite columnarius Tarasov & Zevma. 1957:184

Distribution: Vladivostok, on Japanese Fishing boat.
Balanus amphitrite fluminensis deOUveira. 1941:21

Distribution: Rio de Janeiro.
Balanus amphitrite helenae Kolosvary. 1949a:7

Distribution: Miocene. Turkistan.
Balanus amphitrite hungaricus Kolosvary. 1948:108

Distribution: Miocene. Hungary.
Balanus amphitrite inexpectatus Pilsbry. 1916:97
Synonymy/diagnosis: Henry. 1943:368.

63

References Davadie, 1963:44; Henry, 1942:126; 1959:199
1960:142; 1973:983; Kolosvary. 1943a:84; 1947a:20
1951c:411; 1967b:391; Nilsson-CanteU, 1933:506; 1939a:3:
Tarasov & Zevina, 1957:187.
Distribution: Gulf of California; Bonaire; Red, Adriatic,
and Mediterranean Seas, (Kolosvary). Pliocene, Florida.
Balanus amphitrite insignis Nilsson-Cantell, 1938b:41
Reference: Karande, 1966:145; 1967:1245.
Distribution: Bombay.
Balanus amphitrite karakumiensis Kolosvary, 1961b:100

Distribution: Miocene, Hungary.
Balanus amphitrite kondakovi Tarasov & Zevina, 1957:191
References: Henry, 1973:991; Rosell, 1973b:88; Zevina &

Tarasov, 1963:94.
Distribution: Mainland coast southeast Asia.
Balanus amphitrite litoralis Kolosvary, 1948:106

Distribution: Miocene, Hungary.
Balanus amphitrite merklini Kolosvary, 1962d:199

Distribution: Miocene, U.S.S.R.
Balanus amphitrite peruvianus Pilsbry, 1909:69
Synonymy/diagnosis: Pilsbry, 1916:97.
References: Henry, 1973:983; Kolosvary, 1943a:84; Nils-
son-Cantell, 1957:10.
Distribution: Costa Rica to Peru; often on mangroves.
Balanus amphitrite poecilosculpta Broch, 1931:59
Synonymy/diagnosis: Nilsson-Cantell, 1934a:61.
References: Broch, 1947:5; Dawydoff, 1952:128; Hiro,

1937c:435; Utinomi, 1958a:294.
Distribution: Indonesia; South China Sea; 33-85m.
Balanus amphitrite rafflesi Nilsson-Cantell, 1934a:64

Distribution: Singapore; on mangroves.
Balanus amphitrite tongaensis Kolosvary, 1962c: 193
References; Kolosvary, 1967b:391.
Distribution: Tonga Is.
Balanus amphitrite vladivostokensis Tarasov & Zevina.
1957:184
Reference: Utinomi, 1967:214 (possibly a synonym of B.

variegatus cirratus).
Distribution: Vladivostok.
Balanus caboblanquensis Weisbord, 1966:26

Distribution: Pliocene, Venezuela.
Balanus caribensis Weisbord, 1966:23

Distribution: Venezuela.
Balanus citerosum Henry, 1973:976
Reference: Southward, 1975:42 (probably equals B.

pallidus)
Distribution: Rio de Janeiro to Santa Catarina.
Balanus dentivarians Henry, 1973:992

Distribution: Southwest Mexico to Ecuador.
Balanus eburneus Gould, 1841:15
Synonymy/diagnosis: Pilsbry, 1916:80.
References: Arvy and LaCombe, 1968:1326 (cement
apparatus); Arvy & Ligouri, 1968:817 (cytochrome oxi-
dase activity); Arvy & Nigrelli, 1969:95 (parasites);
Arvy et al, 1968:817 (alkaline phosphatase activity);
1969:351 (parasites); Bacon, 1971:187 (populations in
relation to salinity); Barnes & Barnes, 1965a:391; (varia-
tion in egg and naupUus size); Barnes & Healy, 1971:83
(biometrical studies); Barnes & Klepal, 1971:81 (pedicel
of penis); Barnes et al, 1970:70 (behavior on impaction);
1971:173 (spermatozoa); 1972:192 (distribution); Bishop,
1951:531; Bishop et al, 1957:7; Bookhout & Costlow,
1959:212 (feeding, molting, growth); Bousfield, 1954:122
(distribution and spawning season); Broch, 1924a: 11 2;
Ciurea et al, 1933:6; Clarke, 1947:73 (poisoning and re-
covery); Cones, 1968:61 (selectivity in fossil preserva-
tion); Costlow, 1963:254 (central nervous system); Cost-
low & Bookhout, 1957a:313 (larval development in
laboratory); Crisp & Costlow, 1963:22 (embryonic toler-
ance to salinity and temperature); Crisp & Southward,
1961:271 (cirral activity); Daniel, 1955c:18; Darwin,
1854b:248; Davadie, 1963:59; deOliveira, 1941:19 (in
part, Henry, 1973:968, 982); DePahna, 1963:15 (fouUng);
Edmondson, 1933:231; Fales, 1928:534 (Ught-receptive

organs); Fischer- Piette & Prenant, 1956:13; Freiberger
& Cologer, 1966:881 (rearing in laboratory); Freiberger
et al, 1969:469 (fouhng and anti-fouhng studies); Gordon,
1969:139 (influence of sahnity on distribution); Gordon
et al, 1970:461 (environmental influence); Grave, 1933:
378 (growth rate); Gregg, 1945:44 (attachment of cy-
prids); 1948:161 (replication of substrate detail); Gruvel,
1903b:137; 1905a:234; Gwilliam, 1963:470; 1965:244
(shadow reflex); Henry, 1954:443 (distribution); 1959:194;
1973:968, 982; Kolosvary, 1943a:81; 1943c:129; 1944:33;
1947a:21; 1965:272; 1967b:392, Kruger, 1940:464; La-
combe, 1970:164 (cement glands); Lacombe & Monteiro,
1974:633; Matsui et al, 1964:141; Mawatari, 1967:99
(harbor fouUng); McDougall, 1943:344; Moore & Frue,
1959:432 (settlement & growth); Neu, 1935b:92 (fouling);
Nilsson-Cantell, 1921:309; 1928a:32; 1931a:109; 1938b:
35; Ostroumoff, 1892:160; Pilsbry, 1918:185; 1924:1;
Pomerate & Reiner, 1942:14 (influence of surface angle
and hght on attachment); ReUni, 1962:3; 1964:406;
1966:179; 1968b:186; 1969:175; Rehni & Giordano, 1969:
250 (vertical distribution and settlement); Riedl, 1963:
258; Sandeen & Costlow, 1961:192 (central nervous sys-
tem); Shaw, 1972:145 (lateral eye); Shimony & Nigrelli,
1971:662 (cement apparatus); 1972:349; Smith, 1946:51
(effects of water currents); Southward, 1962:163 (tem-
perature on cirral activity); Southward & Crisp, 1963:34;
Stubbings, 1967:270; Sumner, 1911:128; Tarasov &
Zevina, 1957:174; Utinomi, 1966:36; Wells, 1966:84;
Weiss, 1947a:56 (tolerance to copper and mercury);
1947b:240 (attachment of cyprids); 1948:116 (abnormal
development); Weltner, 1897:266; 1898b:12; Wharton,
1948:180 (primary attachment); Visscher, 1928b:193
(fouling and resistance to fresh water); Zevina, 1963:73;
Zevina & Goryn, 1971:771; ZuUo, 1963b:ll.
Distribution: Endemic to western Atlantic. Boston to
Rio de Janeiro; introduced to Europe, Mediterranean,
Indian Ocean, Japan, Hawaii and other islands of
Pacific Oceania. Miocene, Haiti; Pleistocene, Panama.
Tertiary, Jugoslavia.

Balanus hophinsi Zullo. 1968:4
Distribution: Plio-Pleistocene. Iceland.

Balanus improvisus Darwin, 1854b:250
Synonymy/diagnosis: Henry, 1959:196.
References: Arbuzova, 1959:462 (permeabihty of basis)
Barnes & Barnes, 1961b:4 (sahnity and biometry); 1962:1
1965a:391 (variations in egg and nauphus size); 1968a
135 (regional variations in egg numbers); Barnes &
Healy, 1969:51 (biometrical studies); Barnes & Klepal,
1971:81 (pedicel of penis); Barnes et al, 1951:227 (orien-
tation); 1970:70 (impaction); 1971:188 (spermatozoa);
Bartha & Henriksson, 1971:7 (anti-fouhng); Bassindale,
1964:39; Belyaev, 1949:901 (osmoregulation); Bishop,
1947:501; 1951:531 (introduction to Australia); Bishop
et al, 1957:4; Blom, 1965:59; Blom & Nyholm, 1961:149
(setthng); Bocquet-Ve'drine, 1962:144; IBocquet-Vedrine
& Parent, 1972:239 (parasitism by Boschmaella): Book-
hout & Costlow, 1959a:212 (feeding, molting, growth);
Borradaile, 1916:132; Bousfield, 1954:120; 1955a:l (eco-
logical control in Miramichi estuary); Brattstrom,
1957:8; Broch, 1924a:81; 1924b:203; 1927c;25; 1935:2;
Bucholz, 1951:49 (larvae); Carlton & Zullo, 1969:1 (early
records on Pacific coast N. America); Ciurea et al,
1933:2; Costlow, 1956:359 (sheU development); 1959:177
(effect of inhibitors); Costlow & Bookhout, 1953:420
(molting and growth); 1957b:224 (body vs. shell growth);
Crisp. 1953:331 (changes in orientation); 1958:483; Crisp
& Southward, 1961:271 (cirral activity); Davadie, 1963:
61; Doochin, 1951:15 (morphology during metamorpho-
sis); Eloffson, 1952:47; Filatowa, 1902:379 (post-embry-
onic development); Fischer, 1872:432; Fischer-Piette &
Prenant, 1956:11; Foster. 1970:388 (acchmation to salin-
ity); Gordon, 1969:139 (influence of sahnity): Gordon et
al, 1970:461 (sodium/manganese content of shell); Gra-
ham & Gay, 1945:381 (attachment and growth); Gruvel,

64

1903b:136; 1905a:231; 1907a:105; 1912a;345; Henry.
1942:110; 1954:443 (distribution); 1973:976,992; Hiro,
1936a:61; Hoek, 1875:60; 1909:271.308; Holmes & Pryor,
1938:795; Jackson & Ross, 1971:188 (on snapping
turtle); Jones & Crisp, 1954:765 (larval stages); Kauri,
1962:131 (frontal filament and nauplius eye); 1966:115
(x-organ); Kawahara, 1961:65 (differences in fouling
communities); 1963b:301 (first record from Japan ■
Pacific side); Kolosvary, 1941a:43; 1942b:204 (as B. i
fossilis); 1943a:82; 1943c:129; 1951c:411; 1959:197
1961a:78; 1962b:206; 1963a:174; 1965a:271 (fouling)
1966b:143; 1967a:388; 1967b:392; Kriiger, 1927a:13
1927b:5; 1940:464; Kuhl, 1965:120; 1967b:965 (in Elbe
estuary); 1968:1 (metamorphosis); Lacombe & Monteiro,
1974:633; Luther, 1950:155; MacDonald, 1951:87; Mak-
simov et al, 1971:1090 (factors influencing population);
Mawatari, 1967:99 (fouling); Mawatari et al, 1968:24
(propagation by ships); McDermott, 1960:199 (preda-
tion); McDougaU, 1943:323; Moore, 1933:969 (orienta-
tion); Moore & Frue, 1959:421 (settlement and growth);
Miiller, 1868:393 ("hybrid", B. armatus); Neu, 1932:143;
1935b:92; Newman, 1967b:1041 (physiology and behav-
ior); Nilsson-CanteU, 1921:310; 1927b;91; 1928a:33;
1931a:110; Norris et al, 1951:444 (variabihty in larval
stages); O'Riordan, 1967:293; Pilsbry, 1916:84(= B.
improvisus var. assimilis Darwin, 1854b:250; = B.
improvisus var. gryphicus Miinter. 1878); Poulsen. 1935
18; Prenant & Teissier, 1923:176; Relini, 1969:173
Schafer, 1952:241 (settling); Schwarz, 1932:437 (influ
ence of Ught); SneU, 1972:1; Southward, 1957:325
Southward & Crisp, 1963:33; Stubbings, 1967:270; Tara
sov & Zevina, 1957:168; Tengstrand, 1931:108 (larvae)
Tornava, 1948:3 (aUmentary canal); Utinomi, 1966:36
van Breeman, 1934:247 (biology); Visscher, 1928a:327
(attachment); 1928b: 193 (fouling); Visscher & Luce,
1928:336 (reaction to Ught); Weiss, 1947a:56 (tolerance
to copper and mercury); 1947b:240 (settlement); 1948:
116 (abnormal development); Wells, 1966:84; Weltner,
1895:289; 1897:266; 1898a:441; 1898b:5; Zevina. 1963:73;
Zevina & Goryn. 1971:771 (Sea of Japan); Zevina & Lit-
vinova, 1970:172; ZuUo, 1963b:12: 1966b:235.

Distribution: East Coast of the Americas, North Atlan-
tic; west coast of Africa to Cape of Good Hope; Mediter-
ranean; Black Sea; Red Sea; Northwestern coast of U.S.
from Washington to San Francisco; Ecuador; Japan;
Austraha; OUgocene. U.S.S.R.
Balanus maroccana Broch, 1927:21

References Beach, 1972:5; Stubbings, 1967:266.

Distribution: North Africa; 40-75m.
Balanus oppidieboraci Ross, 1964a:490

Distribution: Miocene, Virginia.
Balanus pallidas Darwin, 1854b:240

Synonymy: Utinomi, 1967:206.

Diagnosis: Stubbings, 1963b:15.

References: AuriviUius 1898a:31; Barnes & Healy, 1969:
51 (biometrical studies); Barnes & Klepal, 1971:83 (pedi-
cel of penis); Bassindale, 1961:485; Broch, 1924b:202;
1927c:26;' 1931:58; Darwin, 1854b:240 (as B. amphitrite
stutsburii and B. Candidas n. sp.); Davadie, 1963:44;
Gauld, 1957:10; Gruvel, 1903b:137,143 (as B. dybowski);
1905a:233,257; Harding, 1962:278,281; Henry, 1954:443;
1959:192 (= B. subalbidus Henry, 1973); Karande,
1967:1247 (as B. a. insignis); Kruger, 1914:437; 1927b:
13; LaCombe & Monteiro, 1974:633; Menesini, 1965:104
(as B. pallidas stutsburii); Moroni-Ruggieri, 1950:72;
Nilsson-CanteU, 1925:28 (as venustas); 1931a:124;
1938a:179; 1938b:38,41 (as B. a. insignis n. sp.); Sandi-
son, 1962:517 (populations on Guinea coast); 1966:363
(effect of saUnity fluctuations); 1967:161 (naupUar
stages); Sandison & HiU, 1966:235 (distribution in rela-
tion to saUnity); Stubbings, 1959:1282 (abnormal devel-
opment); 1961b:24; 1964b:338; 1965:887; 1967:277; Welt-
nar, 1897:256; 1922:83; Zevina & Litvinova, 1970:173;
ZuUo, 1963b:9.

Distribution: West coast of Africa; Northern Indian

Ocean; Gulf of Siam, southwestern AustraUa; Gulf of
Mexico, Caribbean; Argentina (Patagonia). PUocene,
Italy.
Balanus patellaris (Spengler), 1780:101

Synonymy: Utinomi. 1968b:174.

Diagnosis: NUsson-CanteU, 1929a:4.

References: Annandale, 1907:40; CaiUiaud, 1865:38;
Caziot, 1921:52; Darwin, 1854b:259; Gruvel, 1903b:139;
1905a:238; Hoek, 1913:152,158; NUsson-CanteU, 1921:
328; 1938b:46; Weltner, 1897:268.

Distribution: India to PhiUppines.
Balanus playagrandensis Weisbord, 1966:29

Distribution: PUocene, Venezuela.
Balanas poecilotheca Kruger, 1911a:48

Synonymy: Utinomi, 1958a:294.

Diagnosis: Hiro, 1937c:435.

References: Barnard, 1924:71, Broch, 1931:59 (as B.
amphitrite forma poecilosculpta); Hiro, 1938a:303;
1939e:263; Kruger, 1911b:460; Nilsson-CanteU, 1934a:61;
PUsbry, 1916:110; Tarasov & Zevina, 1957:188; Utinomi,
1949a:22; 1962:216.

Distribution: Japan; Formosa; Sulu Arch.; South Africa.
Balanus reticalatus Utinomi, 1967:216

Synonymy/diagnosis: Southward, 1975:11.

References: Broch, 1922:314; 1931:58 (as B. a. com-
munis); Darwin, 1854b:240 (as B. a. communis, in part);
Henry, 1973:968; Hiro, 1938a:301 (as B. a. communis);
Hoek, 1913:168 (as B. a. communis); Kolosvary, 1939b:
129 (as B. a. communis); 1962b:205 (fossU); Mawatari,
1967:99 (as B. a. tesselatus); RoseU, 1973b:79 (as B. a.
amphitrite); Southward & Crisp, 1963:43 (as B. a. amphi-
trite var.); Stubbings, 1961a:173; Utinomi, 1960:44;
1969b:51,52; 1970:356; Utinomi & Kikuchi, 1966:5 (as
B. variegatus tessalatus nom. nov.).

Distribution: Circumtropical fouUng form.
Balanus salaami NUsson-CanteU, 1932c:5

Distribution: Dar-es-Salaam.
Balanus subalbidus Henry, 1973:968

Distribution: Southeast U.S.; Gulf of Mexico and West
Indies; generaUy in brackish water.
Balanus suturaltus Henry, 1973:983

Distribution: West coast of Central America.
Balanus uliginosis Utinomi, 1967:202

Synonymy/diagnosis: Utinomi, 1957c:202.

References: Hiro. 1938a:305; 1939e:263; Kruger. 1911a:
51 (as Balanus amphitrite niveus); Mawatari, 1967:99
(distribution of fouling organisms); Mawatari et al, 1962:
93; 1968:24 (propagation by ships); RoseU, 1973b:86 (as
B. a. krugeri); Tarasov & Zevina, 1957:190; Utinomi
1949a:22; 1962:216; 1967:202 (new name for B. a. krugeri
NUsson-CanteU, 1932a:24); 1969b:52; 1970:356; Utinomi
& Kikuchi, 1966:5; Zevina & Tarasov, 1963:93.

Distribution: Southern Japan; southern Korea; China;
Formosa.
Balanus variegatus variegatus Darwin, 1854b:241

Synonymy: Utinomi, 1968b:171 (includes Balanus amphi-
trite malayensis Hoek, 1913:172).

Diagnosis: "Harding, 1962:291.

References: Bhatt & Bal, 1960:439; Broch, 1916:5; 1931:
58; Daniel, 1955a:97 (gregariousness); 1955c:19; 1956:21
(influence of color); 1957a:305 (effect of iUumination);
1957b:866 (influence of tide); Darwin, 1854b:241 (as
Balanas amphitrite var. 8, variegatus); Foster, 1974:48
(as B. amphitrite malayensis); Gruvel, 1905a:233;
1907d:6; Hoek, 1913:172; Hutton, 1879:328; Karande,
1967:1245; 1974:229 (larval comparison with B. amphi-
trite); Karande & Palekar, 1966:143; Moore, 1944:333;
NUsson-CanteU, 1934a:60; 1934b:57; 1938b:39; Pope,
1945:362 (as Balanus amphitrite cirratus); Stubbings,
1963a:329; Tarasov & Zevina, 1957:183; Weltner, 1897:
266; 1900:305.

Distribution: New Zealand; AustraUa; Indonesia; Viet-
nam; Bay of Bengal.
Balanus variegatus cirratus Darwin, 1854b:241

Synonymy: Utinomi, 1967:214.

65

Diagnosis Harding, 1962:293.

References Darwin, 1854b:241 (as Balanus amphitrite
var. 9 cirratus); Davadie, 1963:44; Gruvel, 1903b:137
1905a:234; Hiro, 1938b;302; 1939e:262; Kolosvary
1961a:78; 1961c:150; 1962d:202; 1967b:392; Mawatari
1967:99 (distribution of fouling organisms); Nilsson
CanteU, 1921:316; 1931a;lll; 1932c:5; 1934a:61; 1934b
56; 1938b:40; Pope, 1945:362; Rosell, 1973:91; Skerman
1960:610; Stubbings, 1963a:331; Tarasov & Zevina
1957:182, 184 (as ? B. amphitrite vladivostokensis); Uti-
nomi, 1949a:22; 1962:216; 1970a:357; Utinonii & Kikuchi,
1966:6; Weltner, 1897:266; Wisely & BUck, 1964:164
(nauplii); Zevina & Tarasov, 1963:89.
Distribution: India, Indonesia, Australia, Philippines
north to Korea. Miocene, U.S.S.R.
Balanus venustus venustus Darwin, 1854b:240
Synonymy/diagnosis Stubbings, 1967:280.
References Annandale, 1906:138; Barnes & Klepal,
1971:81 (pedicel of penis); Broch, 1924b:202 et seq.;
Daniel, 1955c:21; Darwin, 1854b:240 (as B. amphitrite
var. 2, venustus); Gauld, 1957:10; Gruvel, 1903b:137;
1905a:233; 1912a:346; Harding, 1962:283; Henry, 1973:
976; Karande, 1967:1245; Karande & Palekar, 1966:145;
Kolosva'ry, 1967b:392; Nilsson-CanteU, 1925:28; 1931a:
110; 1938b:37; Stubbings, 1961b:29; 1961c:188; 1963b:
21; 1964a:109; 1965:887; Tarasov & Zevina, 1957:189;
Utinonii, 1960:46; 1969a:86; 1970:355; Weltner, 1897:265.
Distribution: Mediterranean; west coast of Africa; South
Africa; Persian Gulf; Bay of Bengal; Sea of Japan;
5-60ni.
Balanus venustus modestus Darwin, 1854b:240
Synonymy/diagnosis Harding, 1962:287.
References Darwin, 1854b:240 (as B. amphitrite var. 5,

modestus); Gruvel, 1905a:233; Weltner, 1897:266.
Distribution: West Indies; Gulf Coast from Florida to
Texas.
Balanus venustus niveus Darwin, 1854b:240
Synonymy: Zullo, 1966b:232.
Diagnosis Harding, 1962:286.

References Barnes & Klepal, 1971:81 (pedicel of penis);
Bernard & Lane, 1961:438 (absorption, excretion); 1962:
19 (early settlement and metamorphosis); Bousfield,
1954:122; Costlow & Bookhout, 1956:107; Darwin.
1854b:240 (as B. amphitrite var. 4, niveus); Dawson,
1957:1068 (fouling of shrimp); deOliveira. 1941:19 (= B.
citerosum Henry, 1973, in part); Doochin, 1951:15
(attachment and metamorphosis); Driscoll, 1968:27;
Eldred, 1962:203 (fouling of shrimp); Fowler, 1912:pl. 46;
Gruvel, 1903b:137; 1905a:224 (as Balanus armatus), 233;
1907d:6; 1909b:25; Henry, 1954:443 (distribution); 1959;
193; Hiro, 1939e:263 (see B. utiginosis); Kolosvary,
1943a:84; 1961a:78; 1961c:150; 1962d:202; 1967b:312;
Kruger, 1911a:51; 1911b:460; Lanchester, 1902:369;
Matsui et al, 1964:144; McDougaU, 1943:354; MuUer,
1867:329 (as Balanus armatus); Nilsson-Cantell, 1921:
318; 1925:31; 1928a:33; 1931a:lll; 1938b:39; 1939a:4;
1957:10; Pearse, 1947:326 (on Limulus); PUsbry, 1916:
92; 1953:25; Ross, 1962:14; Smith, 1946:51 (effect of
water currents); Stubbings, 1964b:340; 1967:268; Tara-
sov & Zevina, 1957:168; Utinomi, 1969a:87; Wells, 1966:
85; Weltner, 1895:289 (as Balanus armatus); 1897:265,
267; 1922:83; Zevina & Litvinova, 1970:174; Zullo,
1963b:12.
Distribution: Western Atlantic, Cape Cod to BrazU;
Mediterranean; west and south Africa; Madagascar;
Red Sea; Persian Gulf; to 55m. Miocene, U.S.S.R.; PUo.-
Pleistocene, Florida.
Balanus venustus obscurus Darwin, 1854b:241
Synonymy/diagnosis Harding, 1962:289.
References Barnard, 1924:70; Darwin, 1854b:241 (as B.
amphitrite var. 7, obscurus); Davadie, 1963:44; Gruvel,
1905a:233; Lanchester, 1902:369; Wells, 1966:85; Welt-
ner, 1897:266.
Distribution: Caribbean; South Africa.

Genus Tetrabalanus Cornwall, 1941

Tetrabalanus polygenus Cornwall, 1941:228
Synonymy/diagnosis ZuUo, 1969d:2.
References Henry, 1973:983,992.

Distribution: Ecuador; Costa Rica; prefers estuarine
conditions.

Group of Balanus trigonus

Balanus alatus Hoek, 1913:175
Reference: Pilsbry, 1916:110.
Distribution: Sulu Arch.; 50-564m.
Balanus calidus Pilsbry, 1916:118
Synonymy: Zullo, 1966b:235.
Diagnosis Pilsbry, 1916:118.

References Daniel, 1955c:21; Darwin, 1954b:225;
DePalma, 1963:19 (fouling); Henry, 1954:443; HuUngs,
1961:215; Karande, 1966:146; 1967:1245; Kolosvary,
1943a:87; 1962a:85; 1967b:391; Nilsson-Cantell, 1939a:6;
Ross et al, 1964:312; WeUs, 1966:83; WeUs & Richards,
1962:586; WeUs, WeUs & Gray, 1964:561.
Distribution: North CaroUna; Gulf of Mexico; West
Indies; 27-64m. OUgocene, Bulgaria. Pleistocene : north-
ern Columbia and Cape Hatteras. y
Balanus calidus nonstriatus Kolosvdry, 1941a:41

Distribution: Gulf of CaUfornia.
Balanus curvirostratus Menesini, 1968c:619

Distribution: PUocene, Italy.
Balanus darwinii Seguenza 1876:453
References Seguenza, 1876:455 (var. calabrus); Davadie,

1953:99; Davadie-Suaudeau, 1952:29; Withers, 1953:62.
Distribution: Tertiary, Italy.
Balanus kanakoffi ZuUo, 1969a:7

Distribution: PUocene, CaUfornia.
Balanus laevis Brugiere, 1789:164
Synonymy: Nilsson-CanteU, 1921:321.
Diagnosis Pilsbry, 1916:120.

References Barnes & Klepal, 1971:83 (pedicel of penis);
CaiUiaud, 1865:38; Darwin, 1854b:227; Davadie, 1963:36;
Gruvel, 1905a:228; Hoek, 1883:150; 1907:4; Kolosvary,
1941e:l (as B. laevis nonsulcatus n. sp.); 1943a:87; 1955:
184; 1959:198; 1960:590; Miers, 1881:79; Newman &
Ross, 1971:174; NUsson-CanteU, 1930c:254; 1931a:112;
1939b:237; 1957:18; Ortmann, 1902:254 (probably
includes B. apertus PhiUppi 1887:224); Weltner, 1895:
291; 1897:263; 1898b:5; 1900:305; Zevina & Kurshakova,
1973:183.
Distribution: Argentina to Tierra del Fuego; Falkland
Islands to Peru; tidal to 275m. Miocene of Europe and
North Africa; Pleistocene of South America.
Balanus laevis coquimbensis Sowerby {in Darwin), 1846:264
Synonymy/diagnosis PUsbry, 1916:122.
References Darwin, 1854b:227; 1897:623; Newman &

Ross, 1971:175; PhiUipi, 1887:224; Weltner, 1897:263.
Distribution: Straits of MageUan to Coquimbo, Chile.
Pleistocene, Coquimbo.
Balanus laevis fossilis Kolosvary, 1950b:3

Distribution: Miocene, Hungary.
Balanus laevis nitidus Darwin, 1854b:227
Synonymy/diagnosis PUsbry, 1916:122.
References Davadie, 1963:37; Davadie-Suaudeau, 1952
26; Gruvel, 1903b:136; 1905a:228; Kolosvary, 1940a:91
Newman & Ross, 1971:175; NUsson-CanteU, 1957:19
Weltner, 1887:101.
Distribution: Straits of MageUan to CaUao, Peru. Mio-
cene, Algeria.
Balanus laguairensis Weisbord, 1966:18

Distribution: PUocene, Venezuela.
Balanus leonensis Weisbord, 1966:43

Distribution: Miocene, Florida.
Balanus minutus Hoek, 1913:177
Synonymy/diagnosis Hoek, 1913:177.
References Broch, 1922:317; NUsson-CanteU, 1925:31

PUsbry, 1916:78; Utinomi, 1968b:173.
Distribution: Sulu Is.; Benin Is.; Singapore; 28-146in.

66

Balanus ochlockoneensis Weisbord, 1966:46

Distribution: Miocene, Florida.
Balanus parkeri Zullo, 1967c:l

Distribution: Gulf of California; 25-36m.
Balanus poecilus Darwin, 1854b:246
Synonymy/diagnosis: Henry, 1960:142.
References: Gruvel, 1905a:229; Nilsson-Cantell, 1957:3;
Pilsbry. 1916:110; Weltner, 1895:289; 1897:266; 1898b:9.
Distribution: Gulf of California and western coast of
South America.
Balanus provisoricus Kolosv^y, 1961:101

Distribution: Miocene, SSSR.
Balanus spongicola Brown, 1844:121
Synonymy/diagnosis: Stubbings, 1963b:22.
References: Barnard, 1924:69; Broch, 1927c:23 (as Bala-
nus doUfusi n. sp.); Crisp & Southward, 1961:271 (cirral
activity); Darwin, 1854a:16; 1854b:225; Davadie, 196
49; deAlessandri, 1895:275; 1906:290; 1907b:277; Gru>
1903b:136; 1905a:225; 1907b;l64; 1909b:25; 1920..
Hoek. 1875:59; 1909:271; Kolosv^ry, 1943a:87; 1947a:65
1951c:412; KrUger, 1940:464; Menesini, 1965:106; 1966
115; 1967b:220; 1972:40; Nilsson-Cantell, 1927a:784:
1938a:180; 1939c:93; ORiordan, 1967:294; Pilsbry, 1916
115; Relini. 1969:171; Seguenza, 1876:288; Southward &
Crisp, 1963:30; Stabbing, 1910:568; Stubbings. 1961b:
32; 1961c:188; 1964b:327 (as B. dollfusi Broch); Weltner.
1897:263; Withers, 1953:61; ZuUo, 1966b:235.
Distribution: Southwestern England; Portugal; Madeira;
Azores; West and South Africa; Indian Ocean. Oligo-
cene to Pleistocene, Mediterranean Basin; PUocene,
England.
Balanus spongicola pliocenicus Seguenza, 1876:443

Distribution: Tertiary, Italy.
Balanus trigonus Darwin, 1854b:223
Synonymy/diagnosis: Pilsbry, 1916:111 (includes B. arma-

tus MuUer, 1868:393).

References: Barnard, 1924:68; Barnes & Klepal, 1971:83

(pedicel of penis); Broch, 1922:320; 1924b:202; 1931:60;

1935:1; 1947:6; Chilton, 1920:53; Cornwall, 1928:11;

1958:81; Cornwall, in Steinbeck & Ricketts, 1941:431.

433; Davadie, 1963:58; Dawydoff, 1952:128; Day &

Morgans. 1956:303; deOUviera, 1941:15; Foster. 1967a:

82; 1967b:33 (early stages); Freiberger & Cologer. 1966;

881 (laboratory rearing); Gordon, 1970:86; Gruvel,

1903b: 136; 1905a:223; 1907a:105; 1907b;164; 1909b:25;

1912a:345,350; GuUer, 1952:20; Henry, 1941:104; 1942:

127; 1943:369; 1954:443; 1960:139; Hirano, 1953:139

(rearing and metamorphosis); Hirano & Okushi, 1952

639 (attachment and growth rates); Hiro, 1932a:551

1937c:439; 1938b:473 (on Macrocheira kaempfen); 1939e:

263; 1939f:210; Hoek, 1883:149; 1913:152; Hutton. 1879

330; Jennings, 1918:61; Kawahara, 1961:65; 1962:27

1963a:391; 1965:319 (fouling); Kolosvary, 1941d;210

1943a:86; 1947a:65; 1951c:411; 1955:184; 1959:197

1963a:173; 1963b;175; 1967b:392; KrUger, 1911a:49

1911b:460; 1940:468; Lacombe & Monteiro, 1974:633

Luckens, 1970c;510; Matsuda, 1973:41; Mawatari, 1967

99 (distribution of fouhng organisms); Mawatari et al

1962:93 (water conduit fouhng); Millard, 1950:266

Moore & McPherson, 1963:418; Moore, 1944:333

NUsson-CanteU, 1921:319; 1927a:784; 1928a;34; 1931a

111; 1938a:180; 1938b:13; 1939a:5; 1939c:93; 1957:10:

Ortmann, 1902:252; Pilsbry, 1909:70; 1916:111; Pope

1945:361; ReUni. 1962:1; 1964:405; 1966:179; 1968a:219:

1968b:186; 1969:173; Rehni & Giordano, 1969:251 (set

tlement); Resig, 1969:20; Ritz & Foster, 1968:551 (tem

perature responses); Ross, 1962:22; 1964a:490; 1964b

271; Ross et al, 1964:313; Sandison. 1954:81; Skerman

1960:610 (predation of); Stubbings, 1936:41; 1940:390

1961b:31; 1963c:188; 1963b:21; 1964a:109; 1964b;341

1965:890; 1967:267; Tarasov & Zevina, 1957:166; UU

nomi, 1949a:22; 1950:63; 1958a:294; 1962:216; 1968b

173; 1969a:88; 1969b:52; 1970:357; Utinomi & Kikuchi

1966:6; Weisbord, 1966:20 (cf. trigonus); WeUs, 1966:83

WeUs et al, 1964:567; Weltner, 1897:262; 1900:307; 1922:
85; Werner, 1967:64 (distribution and ecology); Wisely &
BUck. 1964:164 (larvae); Withers, 1924:33; 1953:74 et
seq.; Zevina & Litvinova, 1970:174; ZuUo, 1963a:122
{B. aethiops Philippi, 1887:224 probably B. trigonus).
Distribution; Cosmopolitan in warm seas; distribution
for the most part natural. Miocene; Europe, Africa and
North America; Pliocene, Italy and Red Sea; Pleistocene
Hawaii.

Group of Balanus perforatus

Balanus hystrix Hoek, 1913:218
Reference: Pilsbry, 1916:78.
Distribution: Sunda I.; 40in.
Balanus obtiquus Ross, 1964a:486
Distributio.n: Miocene, Virginia.
Balanus pacificus Pilsbry, 1916:104 (= Balanus concauus
pacificus)
Synonymy: Ross, 1962:16; 1964a:489.
Diagnosis; Pilsbry, 1916:104.

References: Boolootian, 1964:185 (on Dendraster excen-
tricus): Cornwall, in Steinbeck & Ricketts, 1941:432;
CornwaU, 1951:328; 1956:647; 1958:84; 1959:406; 1962:
625; Darwin, 1854b:235 (in part, figs. 4a-c); Davadie.
1963:52; Giltay, 1934:1 (on Dendraster): Henry, 1942:
104; 1943:367; 1959:200; 1960:146; Hertlein, 1934:61;
Kolosvary, 1955:185; Merrill & Hobson, 1970:595 (on
Dendraster excentricus): NUsson-CanteU, 1957:6; Orcutt,
1921:24; PUsbry, 1907d:199 (as B. concavus - recent,
Point Loma); 1909:67 (as B. concavus - fossU, Peru);
Weltner, 1895:291 and 1897:261 (as Balanus tintinnabu-
lum occator): ZuUo, 1969a: 10.
Distribution: South of San Francisco to ChUe. PUo-
Pleistocene of CaUfomia; Pleistocene of Magdalena Is.;
fossil, Peru.
Balanus pacificus brevicalcar Ross, 1964a:488

Reference: PUsbry. 1916:107,337 (as Balanus concavus

pacificus forma brevicalcar); Ross, 1964a:488.
Distribution: Newport, CaUfornia.
Balanus pacificus prebrevicalcar Ross, 1964a:488

Distribution: Miocene, Virginia.
Balanus perforatus Brugiere, 1789:167
Synonymy/diagnosis: PUsbry, 1916:123.
References: Austin et al, 1958:497 (chromosome num-
bers); Barnes & Barnes, 1965a:391 (variation in egg and
naupUus size); 1966a:83 (ecological and zoogeographical
observations); 1968a:146 (variation in egg production);
1974:197 (embryonic development and saUnity); Barnes
& Crisp, 1956:636 (self-fertiUzation); Barnes & Klepal,
1971:83 (pedicel of penis); Barnes et al, 1970:70 (behav-
ior on impaction); 1971:173 (spermatozoa); 1972:191;
Bassindale. 1964:37; Bishop et al, 1957:9; Bocquet-
Vedrine & Pochon-Masson, 1969:595 (spermiogenesis);
CaiUiaud, 1865:38; Caziot, 1921:52; Ciurea et al, 1933:7,
16; Crisp, 1964a: 181. et seq. (effects of severe winter);
Crisp & Patel, 1958:1078 (relationship between breeding
and ecdysis); Crisp & Southward, 1961:271 (cirral activ-
ity); Daniel, 1955c:22; Darwin, 1954b:231; Davadie,
1963:38; Davadie-Suaudeau. 1952:20; deAlessandri,
1895:279; 1907b:278; Ephrusi, 1922:141 (spermatozoa);
Fischer, 1872:432; Fischer- Piette & Prenant, 1956:16;
Grasse & Tuzet, 1928:1543 (spermatozoa); 1932:9 (sper-
matozoa); Groom, 1894a;119 (early development); 1894b:
81 (hfe history); Groom & Loeb, 1890:160 (nauphar be-
havior); Gruvel, 1905a:230; 1907d:6; 1912a:345; Hoek,
1909:271,283; 1913:158; Knight-Jones, 1953:585 (gregar-
iousness); Kolosvary, 1943a:88; 1944:33; 1947a:14;
1947d:425; 1951b:292; 1951c:411; 1955:184; 1960a:591;
1963a:173,175; 1967b:392; Kruger, 1940:464; LeReste,
1965:64 (larva); Lochhead, 1936:429 (feeding mechanism
of naupUus); Menesini. 1965:95; 1967b:217; Moore, 1936:
703; Moyse, 1960:120; Munn & Barnes, 1970b:261 (fine
structure of spermatozoa); NUsson-CanteU, 1931a:112;
Norris & Crisp, 1953:393 (distribution and planktonic

67

stages); Norris et al, 1951:444 (variability in larval
stages); ORiordan, 1967:292; Patel & Crisp, 1960b:104
(rates of development of embryos); Prenant & Teissier,
1923:173; Pochon-Masson, et al 1969-1970:205; Relini,
1964:404; 1966:179 (fouling); 1968b:185; 1969:171; ReUni
& Giordano, 1969:251 (vertical distribution); Riedl, 1963:
258; Seguenza, 1876:293; Southward, 1955a:1124 (feed-
ing); 1955b:403 (cirral activity and temperature); 1963:
798 (hemoglobin); Southward & Crisp, 1963:29; Stub-
bings, 1963b:30; 1964b:342; 1967:268; Tarasov & Zevina,
1957:193; Taylor, 1970:211 (frontolateral horns and
glands); Weltner, 1898b:12; Withers, 1953:57 et seq.;
Zevina, 1963:72.

Distribution: Great Britain; France; Spain; Mediterra-
nean; Black Sea; northwestern coast of Africa. Oligocene-
Pleistocene, Europe and Africa.
Batanus perforatus altavellensis Seguenza, 1876:446

Distribution: Tertiary, Italy.
Balanus perforatus angustus (Gmehn), 1789

Synonymy: Darwin, 1845b:231.

Diagnosis: Davadie, 1963:39.

References: Broch, 1924b:204; 1927b:22; 1935:2; Gruvel,
1903b:136; 1905a:230; Kolosvary, 1942d:149; Nilsson-
Cantell, 1931a:112; 1938a:180.

Distribution: Great Britain; France; Spain; Mediterra-
coast of Africa; Indian Ocean.
Balanus perforatus chordatus Menesini, 1966:113

Distribution: Miocene, Italy.
Balanus perforatus cranchii (Leach), 1818:pl. 57

Synonymy: Darwin, 1854b:231.

Diagnosis: Davadie, 1963:39.

References: Brown, 1844:121; Gruvel, 1905a:230; Mene-
sini, 1965:101; Pilsbry, 1916:125; Weltner, 1897:264.

Distribution: Pleistocene, Italy.
Balanus perforatus fistulosus (Poli), 1791:22

Synonymy: Darwin, 1854b;231.

Diagnosis: Gruvel, 1905a:230.

References: Broch, 1927c:23; Nilsson-Cantell, 1931a:112;
Vivi, 1938:111 (digestive tract); Weltner, 1897:264.

Distribution: Denmark; Morocco; Canary Is.
Balanus perforatus mirabilis Darwin, 1854b:232

Synonymy/diagnosis: Darwin, 1854b:231.

References: Gruvel, 1905a:230; Pilsbry, 1916:125; Welt-
ner, 1897:254.

Distribution: RocheUe, France.

GeTwis Megabalanus Hoek, 1913

Megahalanus ajax (Darwin), 1854b:214

Synonymy/diagnosis: Nilsson-Cantell, 1938b:34.

References: Fischer, 1884:357; Gruvel, 1903b:126; 1905a
214; 1907b:164; 1909b:25; 1912a:350; Hoek, 1913:151
Kolosvary, 1956:189; 1959:197; Kriiger, 1940:464
Pilsbry, 1916:74; Weltner, 1897:262.

Distribution: Indian Ocean; Philippines; Solomon Is.;
New Caledonia; Japan. Miocene, Hungary.
Megabalanus atgicola (Pilsbry), 1916:72

Synonymy: Utinomi, 1968b:170.

Diagnosis: Pilsbry, 1916:72.

Reference.S: Barnard, 1924:67 (includes var. costatus);
Barnes &. Barnes, 1965a:391 (variation in egg and naup-
hus size); Barnes & Klepal, 1971:81 (pedicel of penis);
Dakin et al, 1948:176; Kolosvary, 1941a:43 (as B. algi-
cola algicola, S. Africa; as B. algicola japonica. n. subsp.
Japan); 1943a:80; 1947c:424 (as fi. algicola forma typica.
Pacific; as B. algicola forma novarae n.f.. Pacific); Krii-
ger, 1940:466; Millard, 1950:266; Nilsson-CanteU, 1939b:
236; Ritz & Foster, 1968:553 (temperature response);
Sandison, 1954:80 (nauplii).

Distribution: South Africa; found elsewhere on ships
(AUen, 1953).
Megabalanus antillensis (Pilsbry), 1916:63

Synonymy/diagnosis: Pilsbry, 1916:63.

References: DePalma, 1963:15 (fouling); de Oliveira,
1941:14; Kriiger, 1940:471; Lacombe & Monteiro, 1974;

633; Nilsson-Cantel, 1928a:31; 1931a:109; 1939a:3;
Pilsbry, 1927:38; 1953:24; Ross, 1968:18; Weisbord,
1966:13; WeUs, WeUs & Gray, 1964:567.

Distribution: North Carolina to Rio de Janeiro.
Megabalanus azoricus (Pilsbry), 1916:62

Reference: Stubbings, 1967:265.

Distribution: Azores.
Megabalanus califomicus (Pilsbry), 1916:65

Synonymy: Ross, 1962:10.

Diagnosis: Henry. 1942:118.

Reference: Aleem. 1957:51; Barnes & Klepal, 1971:79
(pedicel of penis); Boolootian, 1958:91; Broch, 1922:310:
Bruff, 1946:234; Coe, 1932:63; Coe & AUen, 1937:126;
CornwaU, 1951:324; 1959:405; Graham & Gay. 1945:382;
Henry, 1943:367; 1960:138; Hewatt, 1946:194; Hughes,
1914:212; Johnson & Snook, 1927:264; Kanakoff &
Emerson, 1959:20; Merrill & Hobson, 1970:613; Ras-
mussen in Shelford, 1935:306; WiUett, 1937:383; ZuUo,
1968:1.

Distribution: Monterey Bay to Cape San Lucas, Baja
California; Guaymas, Mexico. Plio-Pleistocene of Cali-
fornia and Baja California.
Megabalanus campbelli (Filhol), 1885:487

Synonymy; Foster. 1967a:82.

Diagnosis: Broch. 1922:310.

References: Chilton, 1909:607; Gruvel, 1903b:128; 1905a:
214; Kruger, 1940:464; Linzey, 1942b:3; Pilsbry, 1916:
54; Weltner, 1897:276; 1900:305; Withers, 1924:27.

Distribution: Campbell I.; Otago Peninsula, New
Zealand.
Megabalanus clippertonensis (Zullo), 1969c:501

Distribution: CUpperton I.
Megabalanus coccopoma (Darwin), 1854b:196

Synonymy: Ross, 1962:9.

Diagnosis: Henry, 1942:120.

References: Broch, 1922:310; Davadie, 1963:26; Gruvel,
1903b:126; 1905a:212; Henry, 1941:102; 1973:983:
Jordan & Hertlein, 1926:420; Kolosvary, 1943a:79
Kruger, 1940:472; Lacombe & Monteiro, 1974:633
Nilsson-CanteU, 1931a:109; Pilsbry, 1916:68; Weltner
1897:260.

Distribution: Mazatlan, Mexico to Panama; Rio de Jan-
eiro; Mauritius; China; New Caledonia. Pliocene, Baja
CaUfornia.
Megabalanus concinnus (Darwin), 1854b:196

Synonymy/diagnosis: Pilsbry. 1916:69.

References: Barnes & Klepal, 1971:81 (pedicel of penis);
Broch, 1931:56; Foster, 1967a:81; Gruvel, 1903b:126;
1905a:213; Hiro. 1936a:60 (commensalism); Jennings,
1918:61: Kolosvary. 1943a:79; Moore. 1944:333; Nilsson-
CanteU. 1957:7; Stubbings, 1967:265; Weltner, 1897:260.

Distribution: West coast of South America.
Megabalanus costatus (Hoek). 1913:165

Distribution: HuU of "Siboga."
Megabalanus crispatus (Schrbter). Darwin, 1854b:195

Synonymy/diagnosis: PUsbry, 1916:60.

References: Barnes & Klepal, 1971:81 (pedicel of penis);
Gruvel, 1903b:212; Stubbings, 1967:265; Weltner,
1897:261.

Distribution: La RocheUe, Senegal; East Indies; on ships.
Megabalanus cylindricus (Gmehn), 1780:3213

Synonymy: Holthuis & Sivertsen, 1967:44 (includes B.
capensis EUis, 1758 and B. maxillaris Gronovius, 1763.).

Diagnosis: Darwin, 1854b:209.

References: Barnard, 1924:67; Davadie, 1963:33; Gruvel,
1903b:129; 1905a:218; Kolosvary, 1943a:90; 1943b:121;
Kruger, 1940:466; Nilsson-CanteU, 1925:28; 1930c:254;
1939b:237; 1939c:93; Pilsbry, 1916:77; Ritz & Foster,
1968:533 (temperature response); Sandison, 1954:90
(naupUi); Stebbing, 1910:568; Stubbings, 1967:267;
Weltner, 1887:101; 1897:261.

Distribution: South Africa.
Megabalanus decorus (Darwin), 1854b:212

Synonymy/diagnosis: Newman & Ross, 1971:176.

68

References Barnes & Klepal. 1971:81 (pedicel of penis)
Broch, 1931:57; Chilton, 1909:607; 1911:311; CornwaU
1959:401 (as Balanus concavus paciftcus); 1960:831
FUhol. 1885:486; Foster, 1967a:81; Hutton, 1879:328:
Gruvel, 1903b:126; 1905a:214; Jennings, 1918:60
Kriiger, 1940:464; Linzey, 1942a:279; 1942b:l (append-
ages); Monod & Dollfus. 1932:71; Moore, 1944:333;
Nilsson-Cantell, 1927a:784; Pilsbry, 1916:77; Skerman,
1958:224 (fouling); Weltner, 1897:261; 1899a:443; 1900:
307; Withers, 1924:25.
Distribution: New Zealand, including Kermadec Is.,
Chatham I., Auckland Is.; subHttoral to 51m. Mio-
cene and Pliocene, New Zealand.
Megabatanus dollfusii (de Alessandri), 1907b:275

Distribution: Upper Miocene, France.
Megabatanus dorbignii (Chenu), 1843
Synonymy/diagnosis: Darwin, 1854b:196.
References: Gruvel, 1903b:126; 1905a:213; PUsbry, 1916:

71; Weltner, 1897:261.
Distribution: On ship from Java.
Megabatanus gatapaganus (Pilsbry), 1916:70
Reference: Hedgpeth, 1969:11 (as S. tintinnabutum).
Distribution: Galapagos Is.
Megabatanus giganteum (Kolosv^ry), 1949:190

Distribution: Miocene, Hungary.
Megabatanus honti (Kolosvary), 1950b:l

Distribution: Miocene, Hungary.
Megabatanus hungaricus (Kolosvary), 1941:282

Distribution: ]\4iocene, Hungary.
Megabaianus intermedius (Darwin), 1854b:196
Synonymy/diagnosis: Darwin, 1854b:196.
References: Gruvel, 1905a:213; Pilsbry, 1916:71; Welt-
ner, 1897:261.
Distribution: ?Peru (Weltner).
Megabatanus isotde (Holthius & Sivertsen), 1967:41

Reference: Nilsson-Cantell, 1939b:237 (as B. maxiltaris).
Distribution: Tristan da Cunha.
Megabatanus javanicus (Withers), 1923:282

Distribution: Miocene, Java.
Megabaianus krakatauensis (Nilsson-Cantell), 1934b:53
Reference: Kriiger, 1940:464.
Distribution: Krakatau, Sunda Strait.
Megabatanus teganyii (Kolosvary), 1950:2

Distribution: Miocene, Hungary.
Megabatanus muttiseptatus (Ross), 1964a:485

Distribution: Miocene, Virginia.
Megabatanus nigrescens (Lamarck), 1818:391
Synonymy: Darwin, 1854b:210.
Diagnosis: Pope, 1945:361.

References: Barnes & Klepal, 1971:84 (pedicel of penis)
CornwaU, 1960:829; Dakin et al, 1948:176; Davadie
1963:32; Endean et al, 1956:88 (ecology and distribu
tion); Gruvel, 1903b:129; 1905a:218; Kolosvary, 1943a
81; KrUger, 1914:429; 1927a:13; 1940:464; Stubbings
1967:266; Weltner, 1897:241; Womersley & Edmonds
1958:232 (ecology).
Distribution: Australia; elsewhere on ships.
Megabaianus occator (Darwin), 1854b:196
Synonymy: Hiro, 1939e:260.
Diagnosis: Kolosvary, 1950a:290.

References: Borradaile, 1900:799; Foster, 1974:46;
Gruvel, 1905a:213; Kolosvary, 1943a:78; Kruger, 1940:
471; NUsson-Cantell, 1938b:34; 1957:6; Nomura, 1938:87;
Pilsbry, 1916:59; Utinomi, 1949a:25; 1954:22; Weltner,
1895:291; 1897:261; Zevina & Tarasov, 1963:88.
Distribution: Indian Ocean; Indonesia; Fiji; Philippines;
Formosa; Bonin Is. Pliocene, Ryukyu Is.
Megabaianus peninsularis (Pilsbry), 1916:66
Synonymy/diagnosis: Pilsbry, 1916:66.
References: Henry, 1941:102; 1942:127; 1943:367; 1960:
146; Kolosvary, 1943a:78; Nilsson-Cantell, 1927a:783
(= M. volcano).
Distribution: Cape San Lucas, Baja California; Acapulco,
Mexico.

Megabatanus plicatus (Hoek), 1913:165

Distribution: Hull of "Siboga."
Megabatanus psittacus (Molina), 1782

SvNONY.MY.DlAGNOSis: Pilsbry, 1916:75.

References: Bahamonde, 1958:214; Chapman, 1914:53
67; Darwin, 1854b:207; Gruvel, 1903b:129; 1904:103:
1905a:217: 1905b:328; 1906a:270; 1907d:l: Henry, 1960
138; Kolosvary, 1941a:41; 1942c:139; 1943a:80;" 1943b
121; 1955:185; 1967b:393; Lacombe, 1970:164 (cement
glands); Menesini, 1967a:47; Nilsson-Cantell, 1929b:489
(mouthparts); 1931a:109; 1957:7; Ortmann, 1902:249
Phillipi, 1887:223; Pilsbry, 1909:66; Tournouer, 1903
471; Vayssiere, 1905:161; Weltner, 1895:291; 1897:261:
1898b:5; 1900:305; Zevina & Kurshakova, 1973:183.

Distribution: Chile and Peru; Juan Fernandez Is.;
Straits of Magellan; Southern Argentina. Plio-Pleisto-
cene, Chile.
Megabatanus psittacus chilensis (Menesini), 1967:47 [nomen

nudum)
Megabatanus rosa (Pilsbry), 1916:61

Synonymy/diagnosis: Yamaguchi, 1973:130,

References: Broch, 1931:56; Hirano, 1953:139 (rearing
and metamorphosis); Hiro, 1932a:549: 1937c:431; 1939f:
208; Kawahara (marine fouling communities), 1962:27
1963a:395; 1965:319; Kolosviiry, 1943a:79; Kruger, 1940
471; Mawatari, 1967:99 (distribution of fouling organ-
isms); Mawatari et al, 1962:93 (fouling); 1963:101
(growth rate, fouUng); Nilsson-Cantell, 1931a:109
1932b:16; Tarasov & Zevina, 1957:164; Utinomi, 1949a
21; 1950:63; 1958a:294; 1962:215; 1969b:51; 1970:349:
Utinomi & Kikuchi, 1966:5; Yamaguchi, 1971:124.

Distribution: Japan, Formosa. Pleistocene, Japan.
Megabaianus seguenzai (de Alessandri), 1895:277

Distribution; PUocene, Italy.
Megabaianus spinosus (Gmelin), 1791:3213

Synonymy: Stubbings, 1967:265.

Diagnosis; Stubbings, 1961c:184.

References: Darwin, 1854b:196; Gruvel, 1903b:126;
1905a:212; Kolosvary, 1943a:78; Lacombe & Monteiro,
1974:633; Nilsson-CanteU, 1931a:109; 1938b:13; Pilsbry,
1916:58; Weltner, 1897:260.

Distribution: Islands in the South Atlantic: St. Helena,
Sao Tome, Principe, Annobon; Rio de Janeiro.
Megabatanus stultus (Darwin), 1854b:216

Synonymy/diagnosis: Ross, 1968:14.

References: Gruvel. 1905a:221; Henry, 1954:443; Kolo-
svary, 1966:69 (as Batanus stultus forma morycowae);
1967b:393; Nilsson-CanteU, 1929a:l; 1939a:3; Pilsbry,
1916:235: 1927:38 (as Tetractita radiata); 1953:25; Welt-
ner, 1897:262.

Distribution: Florida and Caribbean; on Millipora.
Megabatanus tanagrae (Pilsbry), 1928:311

Reference: Gordon, 1971:83.

Distribution: Hawaiian Is.
Megabatanus tintinnabutum (Linneaus), 1758:668

Synonymy: Darwin, 1854b:194 (includes pre-Darwin
references).

Diagnosis: Pilsbry, 1916:55.

References: Annandale, 1906:147; 1911:1170 (growth
rate); Barnard, 1924:66; Barnes & Klepal, 1971:79 (pedi-
cel of penis); Boolootian, 1958:91 (attached to echinoid);
Borradaile, 1903:441; Brocchi, 1814:597; Broch, 1924b:
203; 1927c;20; 1927d:133; 1931:56; Bruntz, 1902:987
(excretion); CaiUiaud. 1865:36; Caziot, 1921:51; Chilton,
1911:132; Cole & Addison, 1931:72 (stimulation by alco-
hols); Cole, 1932b:143 (sensitivity of cirri); Daniel, 1952:
261 (respiratory mechanism); 1955a:99 (gregarious
attraction); 1955c:17; 1956:21 (influence of color on
settlement); 1957a:305 (effect of illumination on settle-
ment); Daniel, 1957b:866 (influence of stage of tide)
Darwin, 1854a:13; Davadie, 1952:26; 1963:26; Dawydoff,
1952:128; de Alessandri, 1895:270; 1906:285; 1907b;270
de Oliveira, 1941:11; 1947:720; Foster, 1967a:81; Gauld
1957:10; Gruvel, 1893a:405 (sheU growth and structure)

69

1903b:125; 1905a:211; 1909b:25: 1912a:345,350; GwiU-
iam, 1965:244 (photoreceptor response); Hart, 1967:1
(chromosomes); Hiro. 1937b:51; 1939a:128; 1939e:258
Hoek, 1883:147; Karande, 1967:1245; Karande & Pale-
kar, 1966:142; Kolosvary. 1943a:77; 1947a:12; 1947c
424; 1947d:425; 1951b:291; 1951c:411; 1959:197; 1960:
590; 1961c:149; 1967b:393; Kriiger, 1911a:47; 1911b;460
1940:464; Lacombe, 1966:1 (cement glands); 1967:1
1968:1; Lacombe & Ligouri, 1969:170; Lacombe & Mon
teiro, 1974:633; Menesini, 1966:104; Moore, 1944:333
Morch, 1852:67; Nilsson-CanteU, 1931a:119; 1938a:179
1938b:33; 1939c:92; 1957:10; O'Riordan, 1967:291; Rao
& Ganapati, 1969:193; ReUni. 1969:170; Riedl, 1963:258:
Seguenza, 1876:438; Stubbings. 1910:567; Stubbings
1936:40; 1961b:20; 1961c:183; 1963b:13; 1964a:108
1964b:336; 1965:885; 1967:263; Tarasov & Zevina, 1957
163; Visscher, 1928b: 193 (fouhng); Withers, 1924:24
Weltner, 1887:101; 1895:291; 1897:260; 1898b:6; 1900
305; 1910:528; Zevina, 1963:72; Zevina & Tarasov,
1963:87.
Distribution: Localities specifically for Balanus tintin-
nabulum tintinnabulum or Balanus tintinnabulum com-
munis: Western coast of Africa from Mediterranean to
Cape of Good Hope; Eastern Mediterranean; Madagas-
car, Arabian Sea; Bay of Bengal; Thailand; Formosa;
Sagami Bay, Japan; New Zealand; Rio de Janeiro; Peru.
Ohgocene and Miocene of Europe; Pho- Pleistocene,
Venezuela.
Megabalanus transsylvanicus (Kolosvary), 1950:3

Distribution: Miocene, Hungary.
Megabalanus transuersostriatus (Beurlen), 1958:3
References: Brito, 1972:2.
Distribution: Para, Brazil.
Megabalanus tubulatus (Withers), 1924:28

Distribution: Phocene, New Zealand (Withers, 1953:80).
Megabalanus tulipiformis (EUis), 1758:851
Synonymy: Utinomi, 1959a:382.
Diagnosis: Darwin, 1854b:204.

References: Crisp & Southward, 1961:271 (cirral activ-
ity); Davadie, 1952:27; 1963:30; de Alessandri, 1895:272
1906:287; Gauld, 1957:10; Gruvel, 1903b:128; 1905a:216
1909b:25; 1912a:350; 1920:53; Hoek, 1875:59; Kolosvary
1943a:81; 1951c:411; Kriiger, 1940:464; Menesini, 1965
92; 1966:107; 1967b:218; NUsson-CanteU, 1921:308:
1931a:108; ReUni, 1969:169; Seguenza, 1876:283; South
ward & Crisp, 1963:28; Stubbings, 1961b:21; 1961c:187
1963b:14; 1964a:108; 1964b:337; 1965:886; Visscher,
1928b:193 (fouhng); Withers, 1953:60,63.
Distribution: Mediterranean; France; Spain; Portugal;
Africa; Madeira, Canary and Cape Verde Is.; 25-250m.
Miocene-Pleistocene, Europe and North Africa.
Megabalanus tulipiformis arenarius (Seguenza), 1876:439
Reference: Davadie, 1963:30.
Distribution: Tertiary, Mediterranean Basin.
Megabalanus tulipiformis etruscus (Menesini), 1966:109

Distribution: Miocene, Italy.
Megabalanus validus (Darwin), 1854b: 195
Synonymy/diagnosis: Hoek, 1913:164,166.
References: Broch, 1931:56; Gruvel, 1903b:126; 1905a:
212; Kriiger, 1914:429; 1940:471; Nilsson-CanteU, 1938b:
12; Weltner, 1897:260.
Distribution: Hull of "Siboga"; southwest Australia;
Taiwan.
Megabalanus venezuelensis (Weisbord), 1966:17

Distribution: Phocene, Venezuela.
Megabalanus vesiculosus (Darwin), 1854b:195

References: Gruvel, 1905a:211; Weltner, 1897:260.
Megabalanus vinaceus (Darwin), 1854b:213
Synonymy/diagnosis: Darwin, 1854b:213.
References: Gruvel. 1905a:215; Kriiger, 1940:466;
NUsson-CanteU, 1957:3; Weltner, 1895:289; 1897:261;
1898b:9.
Distribution: West coast of South America.
Megabalanus volcano (Pilsbry), 1916:60

Synonymy/diagnosis: Yamaguchi, 1973:133.

References: Hiro, 1937c:430; 1938c:1848 (resistance to
saUnity and exposure); 1939:208; Kriiger, 1940:471
Mawatari et al, 1962:93 (fouhng); Nilsson-Cantell, 1927a
783 (as Balanus tintinnabulum peninsularis); 1938b:34
Tarasov & Zevina, 1957:165; Utinomi, 1949a:21; 1958a
293; 1958b:51; 1969b:51; 1970:350; Utinomi & Kikuchi,
1966:5.

Distribution: Southern Japan; Okinawa.
Megabalanus wilsoni (ZuUo), 1969a: 10

Distribution: Phocene, CaHfornia.
Megabalanus zebra (Darwin), 1854b:195

Synonymy: Stubbings, 1967:264.

Diagnosis: Pilsbry, 1916:57.

References: Barnard, 1924:66; Barnes & Klepal, 1971:81
(pedicel of penis); Davadie, 1963:26; Gruvel, 1903b: 126;
1905a:212; 1909a:214; 1912a:350; Hiro. 1939e:259;
Karande, 1967:1245; Karande & Palekar, 1966:143;
Kolosvary, 1943a:78; Menesini, 1966:106; Stubbings,
1961b:21; 1964a:108; Utinomi, 1968b:170; Weltner,
1897:260.

Distribution: West Africa; Cape Verde Is. to Walvis
Bay; Formosa; Phihpines.

Incertae Sedis

Chthamalus revilei Locard, 1878:17

Distribution: Neogene, France

Remarks: Absence of opercular parts, and size of shell
(basal dia. 27mm, height 15mm) precludes assignment
to Chthamalus ss.
Balanus borsodensis Kolosvary. 1952:410

Distribution: Miocene, Hungary.
Balanus chisletianus Sowerby, 1859

Reference: Withers. 1953:39.

Distribution: Eocene(?), England.
Balanus echinicola Hoek, 1912:408

Distribution: Malay Arch.; 216m.

Remarks: Apparently never described, hence nomen
nudum.
Balanus ecuadoricus Pilsbry & Olson, 1951:200

Distribution: Ohgocene of Ecuador.

Remarks: Authors suggest relationship with B. nubilus
but opercular parts appear close to crenatus.
Balanus flosculoidus Kolosvary, 1941e:9

Distribution: Japan.
Balanus gizellae Kolosvary. 1962c:195

Reference: Kolosvary. 1967b:392.

Distribution: Tonga I.
Balanus hohmanni Philippi. 1887:225

Distribution: Tertiary, Chile.
Balanus irregularis Broch, 1931:61

Distribution: Banda Sea; 290m.

Remarks; Mouth parts wrong for B. crenatus; form is
that of Solidobalanus, but Brock placed in his Eubal-
nus (porous wall), which for present precludes its
assignment.
Balanus humilis Conrad, 1846:400

Reference: Ross, 1967:173 (internal cast).

Distribution: Miocene, Florida.
Balanus mirabilis Kriiger, 1912:11

Reference: Pilsbry, 1916:79.

Distribution: Japan.

Remarks: Figures suggest it may belong to the group of
B. amphitrite.
Balanus microstomas Phihppi, 1887:225

Distribution: Tertiary, Chile.
Balanus pannonicus Kolosvdry, 1952:233

Distribution: Miocene, Hungary.
Balanus sauntonensis Parfitt, 1871:210

Distribution: Fossil, North Devon, England.
Balanus shilohensis Pilsbry, 1930:431

Distribution: Miocene, New Jersey.

Remarks: Too incompletely known to be placed in a

70

group. Pilsbry compares it to B. concavus and Semi-
balanus.
Balanus similis Weltner, 1922:83
Distribution: Off South Africa; 638ni.
Remarks Porous wall precludes placing in Solidobatanus;
figure suggests wall of 8 plates.
Balanus tuboperforatus Kolosvary, 1962c:197
Reference: Kolosvary, 1967b:392.
Distribution: Tonga I.
Balanus tumorifer Kolosvdry, 1962c:195
Reference: Kolosvdry, 1967b:392.

Distribution: Tonga I.
Balanus veneticensis Seguenza, 1876:303

Reference: Withers. 1953:62.

Distribution: Tertiary, Italy.
Balanus violaceus Gruvel, 1903b:133

Distribution: Unknown.

Remarks: Author compares with nubilus; appears to us to
be closer to group of B. amphitrite. Lamy and Andre
(1932:218, footnote) proposed specific name of abeli to
replace violaceus which was preoccupied.

71

LITERATURE CITED

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1920. Lehrbuch der Palaozoologie, Gustav Fischer, Jena.
500 pp.

1926. An der kalifornischen Kuste. In, Amerikafahrt,
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1927. Paracreusia Trolli n.g. n. sp., eine auf Stockkorallen
schmarotzende Balane aus dem miozanen Medi-
terranmeer. Verh. Zool.Bot. Ges. 77:101-103.

1928. Parasitische Balanen auf Stockkorallen aus dem
mediterranen Miozanmeer. Palaeobiolofrica 1:13-38.

1935. Vorzeithche Lebensspuren. Gustav Fischer, Jena.
644 p.
Achituv, Y.

1972. The zonation of Tetracht hamulus oblitteratus
Newman and Tetraclita squamosa rufotincta
Pilsbry in the Gulf of Elat, Red Sea. J. Exp. Mar.
Biol. Ecol. 8:73-81.
Addicott. W.O.
1966. Late Pleistocene marine paleoecology and zooge-
graphv in central California. U.S. Geol. Surv. Prof.
Pap. 523-C: C1-C21.
Aleem, A. A.
1957. Succession of marine fouling organisms on test
panels immersed in deep water at La Jolla,
California. Hydrobiologica 11:40-58.
AUen, F. E.

1953. Distribution of marine invertebrates by ships.
Austral. J. Mar. Freshwater Res. 4(21:307-316.
Allison, LB. and W.H. Cole

1935. Behaviour of the barnacle, BaJanus balanoides, as
correlated with the planktonic content of the sea
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gonioporae, P. grande, P. projectum, Tetrachtha-
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1975. Observations on the proteolytic enzymes in the
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1975. Tissue culture of cirripeds. Biol Bull. 149(21:316-330.
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Ganapati, P. N. and D. R. K. Sastry

1974. Record of Athanas indicus (Coutiere) (Decapoda:
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Gardner, D. and J. P. Riley

1972. Seasonal variations in the component fatty acid
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Gorin, A. M. and A. M. Murakhveri

1973. Seasonal dynamics of settling and growth of
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Granier, J.

1973. Le genre Balanus sur les cotes de Camargue et du
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Henry, D. P. and P. A. McLaughlin

1975. The barnacles of the Balanus amphitrite complex
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a. cochinensis, B. a. columnaris, B. a. communis,
B. a. denticulata, B. a. fluminensis, B. a. form-
osanus, B. a. franciscanus, B. a. hawaiiensis, B. a.
herzi, B. a. inexpectatus, B. a. insignis, B. a. kon-
dakovi, B. a. krugeri, B. a. matayensis, B. a.
modestus, B. a. niveus, B. a. obscurus, B. a.
pallidus, B. a. peruvianas, B. a. poecilosculpta, B.
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tesselatus, B. a. variegatus, B. a. venustus, B. a.
vladivostokensis, B. a. saltonensis, B. carenatus,
B. citerosum, B. concavus indicus, B. c. mexicanus,
B. c. pacificus, B. c. p. brevicalcar, B. aquila regalis,
B. c. sinensis, B. democraticus, B. dentivarians, B.
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B. pacificus, B. pacificus brevicalcar, B. pallidas,
B. pallidus krugeri, B. patellaris, B. patelliformis,
B. aquila, B. reticulatus, B. subalbidus, B. sutural-
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1972. Two new sea-urchin-acorn barnacle associations.
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1972. Introduction to the ecological study of intertidal
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1973. Cria^ao de Balanideos em laboratorio. Trab. V
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1974. Marine fouling studies off Oahu, Hawaii, USA.
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1974a. Population density of Balanus balanoides in rela-
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95-99. {Balanus panamensis: a senior synonym of
Balanus eyerdami Henry, according to D. P.
Henry, pers. comm.)

Roth, V. D. and W. L. Brown.
1975. A new genus of Mexican intertidal zone spider
(Desidael with biological and behavioral notes. Am.
Mus. Novit. 2568:1-7. ITetraclita squamosa)

Sergy, G. A. and J. W. Evans

1975. The settlement and distribution of marine organ-
isms fouUng a seawater pipe system. Veliger
18(11:87-92. IBalanus balanoides)

Shikami, T.

1973. MoUuscan assemblages of the basal part of the
Zushi Formation in the Miura Peninsula. Sci. Rep.
Tohuku Univ., Sec. Ser, (Geol.l, Spec. 6:179-204.
IBalanus aff. amphicostatus l=albicostatusl )

Southward, A. J.
MS. A reconsideration of the taxonomic status and
distribution of Chthamalus stellatus (Cirripedial in
the N. E. Atlantic region.

Stickle, W. B.

1973. The reproductive physiology of the intertidal
prosobranch Thais lamellosa (Gmelinl. 1. Seasonal
changes in the rate of oxygen consumption and
body component indexes. Biol. Bull. 144(3):51 1-524.
IBalanus cariosus, B. glandula)

Thomas, M. L. H., D. R. Grant and M. de Grace

1973. A new late Pleistocene marine shell deposit at
Shippegan New Brunswick. Can. J. Earth Sci.
10(81:1329-1332. IBalanus crenatus, B. hameri, B.
improvisus)

Wagh, A. B. and D. V. Bal.

1974. Observations on systematics of sessile barnacles
from the west coast of India: I. J. Bombay Nat.
Hist. Soc. 71(11:109-123. IBalanus am'aryllis
euamaryllis. B. amphitrite communis. B. a. hawaii-
ensis, B. a. stutsburi. B. a. venustus, B. tintin-
nabulum tintinnabulum. Chelonibia patula, C.
testudinaria, Chthamalus malayensis, C. withersi,
Tetraclita ITetraclitella) purpurascens)

Walker, G., P. S. Rainbow, P. Foster and D. J. Crisp

1975. Barnacles: possible indicators of zinc pollution?
Mar. Biol. 30(11:57-66.

Walker, G., P. S. Rainbow, P. Foster and D. L. HoUand
1975. Zinc phosphate granules in tissue surrounding the
midgut of the barnacle Balanus balanoides. Mar.
Biol. 33(21:161-166.

103

SYSTEMATIC INDEX

(Only italicized page numbers lead directly to valid species in the Catalog)

Aaptolasma 46,20-22,31,33

abeli (see violaceus). 70, 101

Abundantus 62

Acasta 53-54,49,23,28,34

Actinobalanus 49,23,24

actinomorphus 49

aculeata 53

acuta, -us, Conopea 54

acuta, -um, Cantellius 56

acutus, Balanus 62

aeneas 51

aeratus 62, 101

aethiops 66

aestuarii 40,41,31

africana 47, 100

ajojc 67,100

alaskensis 61

alatus 65,101

alba, Acasta 53

alba, Tetraclita 47, 100

albus, Chirona 50

albicostatus 62, 101,102

formosanus 62

alcyonicola 53

algicola 67

costatus 67

japonica 67

novarae 67

typica 67

allium 49

truncatus 49

alloplax 61

altissimus 61

altavellensis 67

amakusana 53

americanum 46,31

amaryllis 50

euamaryllis 50,100,102

dissimilis 50

laevis 50

nivea 50

amphitrite 62,64,70,33,34,

100,101,102

abundantus 62

acutus 62

aeratus 62, 101

albicostatus 62,101,102

archi-inexpectatus 62

cirratus 64,65,100,101

cochinensis.- 62, 101

columnarius 62, 101

communis 62,64,100,101.102

fluminensis 62, 101

helenae 62

hungaricus 62

inexpectatus 62-63,34,101

insignis 63,64,101

karakumiensis 63

kondakovi 63, 101

krugeri 64,101

litoralis 63

malayensis 64,101

merklini 63

obscurus 65,101

peruvianas 63, 101

peocilosculpta 63,64,101

rafflesi 63,101

stutsburii 64,101,102

tesselatus 64,101

tongaensis 63

variegatus 64,100,101

venustus 65,101,102

vladivostokensis 63,65,101

amphitrite, group of Balanus 62-

65,69,13,23,24,34

anchoris 54

Andromacheia 59

anglicum, -a 59, 58

angulosa 40, 100

angusticalcar 53

angustiradiata 58

angustiterga, Creusia 58

angustitergum, Chthamalus 41

angustus 67

anisopoma 41

annandalei 58

antarcticum 46

antennatus 4i,42,19

antillensis 67

antipathidis 53

antiqua, -um, Coronula 45

antiquus, Chthamalus 42,50

aotea 44,45

aperta, Acasta 53

apertus, Balanus 61,65

apertus, Balanus rostratus 61

appelloefi 40

aquila 6i, 100,101

arafurae 46

Archaeobalanidae, -inae 49-56,

11,23,24,38

Archaeobalanus 49,22-24

archi-inexpectatus 62

arcuatus, Balanus 49

arcuatum, Cantellius 57,34

arenarius 69

artica 59

armata, Acasta 53

Armatobalanus 49,50,23,24,

28,30,31

A. (Armatobalanus) 49,50,23,34

A. (Hexacreusia) 50,23

armatus, Balanus 64,65,66,101

assimilis 64,101

astacophilus 50

aucklandicum 45-46

aurantiacum 40

auricoma 50

Austrobalaninae 46,11,21,38

Austrobalanus 46,49,21,31

azoricus 67

balaena 45

balaenaris 45

Balanidae 59-69,11-16,23,39

Balanoidea 49-69,9,12,15,

22-24,38,30,31

balanoides 55-56,22,25,28,

100,101,102

calcaratus 56

Balanoidomorphoidea 43,2,20-22,

36,30,31
Balanomorpha 9-24,36,26,27,

28,29,31
Balanus 59-69, 14,23,28,30,

31,33,34

balanus 59-60, 100

pugetensis 59,60

balanus, group of Balanus 59-60,

23

barbadensis .58

barbara 45

basicupula 53

(Bathybalanus) 52,23

Bathybalanus 52,22,23

Bathylasma 45,46,15,20-22,31.33

Bathylasmatidae, -inae 45,46,11

13,21,37

belyaevi 41

bifida 45

bimae 50

bimanicus 50

biscayensis 45

bisexlobata 44

bisinuatus 43

bisulcatus 49

plicatus 49

bloxhamensis 61

borsodensis 69

Boscia 5923,28

Boscinae 59,11,23,24,39

brachialis 52

Brachylepadomorpha 11.12,15,16

brevicalcar 66, 101

breviscutum 46,33

brevitergm 57

brintoni 46

brunnea, Chamaesipho 43

brunnea, Octomeris 40

caboblanquensis 63

calabrus 65

calcaratus 56

calcareobasis 41

calceolus, -a 54

calidus 65

nonstriatus 65

califomica, Diadema 45

califomicus, Megabalanus 67,30

callistoderma 46

calvertensis 49

campbelli 67

cancetlorum 53

cancellata 58

candidum, Coronula 45

candidus, Balanus 64

CanteUius 56-57,23,34

capellini 43

capensis 67

carenatus 62,101

caretta 43

caribensis 63

cariosus 56,30,102

Catomerus 40,14,17,18,31

Catophragmidae 40,11,19,36

Catophragmus 40,14,17,18,29

caudatus, -a 41

cepa 49

Ceratoconcha 58-59,23,24,28

Ceratoconchinae 58,11,23,24,39

Cetolepas 45,21

Cetopirus 45,21

challengeri 4i, 42,100

krakatauensis 41

nipponensis 41

104

Chamaeosipho 43,17,18

Chelonibia 43,20-22,29,33

Chelonibiinae 43,44,11,21,37

cheltrypetes 43

chesapeakensis 61

chilensis 68

chinense, Pachylasma 40

chinensis, Tetraclitella 46

Chionelasmus 40,4,17,18,

19,31,32

Chirona. 50

C. (Chirona) 50,23

C. (Striata balanus) 50,23

chisletianus 69

chordatus 67

Chthamalidae, -inae, -oidea 40,41-

43,11-16,17-20,36,29.30,31

Chthamalus 40,13,17,18,31,32

ciliatus 50

circe 49

cirratus, Chthamalus 4i,40

cirratus. Balanus ... 64-65,63,100,101

citerosum 6?,65,101

cladangiae 59

ctavatus 60

clippertonensis 67

coccopoma 67

cochinensis 62, 101

coerutescens 47, 100

columna 43

columnaris 62, 101

communis, Balanus amphitrite . . . . 62,

100,101,102

communis, Chthamalus stellatus. . . 42

communis, Megabatanus 68

communis, Tetraclita 48,100

complanatus, -a 45

compressus 51

concavus 6i, 66,68,70

alloplax 61

chesapeakensis 61

coosensis 61

dallonii 61

eseptatus 61

finchii 61

glyptopoma 61

indicus 61,101

mexicanus 61, 101

oligoseptatus 61

pacificus 66,68,101

proteus 61

raphanoides 61

rariseptatus 61

rubescens 61

scrutorum 61

sinensis 67, 101

concavus, group of Balanus 61-

62,23,28,31

concinnus 67

confinis 48

conica 5,3,34

conicocystata 58

conjugatum 58, 100

connelli 60

Conopea 54-55,23,28,30

coosensis 61

coquimbensis 65

coriobasis 53

comutus, Chthamalus 43

comutus, -a, Conopea 55

comwalli 51

corolliforme. -is 46,31,33,100

Coronula 44,45,11,21

Coronulidae, -inae 43-45,11,20,

21,24,37

corrugatus 49

costata, -um, Ceratoconcha. 58,59

costata, Tetraclitella 46

digita 46

costatus, Megabalanus 67

cranchii 67

crassa, Acasta 53

crassa, Octomeris 40

crenatibasis 43

crenatiformis 57

crenatum, Savignium 57

crenatus, Balanus 60,59,69,30,

100,101,102

curviscutum 60

delicatus 60

cretaceum 40

Creusia 57-58,56,59,13,23

creusioides 58

crinoidophilum 40

crispatus 67

cristallinus 52

Cryptolepas 45,21

ctenodentia. 53

cuneiformis 52

curvirostratus, Balanus 65

curviscutum 60

cuspidatus 53

cyathus 53

cybosyrinx 44

cylindricus 67

Cylindrolepas 44, 21

cymbiformis 55

dalli, Balanus 61

dalli. Chthamalus 42

dalloni 61

darwini, Balanus (B.) 65

calabrus 65

darwini, Cetopirus 45

darwini, Chionelasmus . . . 40, 4, 31, 32

darwini, Coronula 45

darwini, Tetraclitella 47

darwiniana, Ceratoconcha 58

darwiniana, Cylindrolepas 44

darwinianum, Pachylasma 40

decima 57

declivis 53

decorata 44

decorus 67-68

delicatus 60

democraticus (see ebumeus) 101

dentata, Chelonibia 44

dentatum, Savignium 57

dentatus, Chthamalus 42, 100

denticulata, Acasta 53

denticulata, Balanus 62, 101

dentifer 55

dentivarians 63, 101

depressa, Chelonibia 43

depressa, Tetraclitella 47

depressa, Tetraclita 47, 48

depressus, -a, Euraphia 41, 40

devonica, Paleocreusia 58

Diadema 45

diadema. 45

digita 46

diploconus, -a 58

dissimitis 50

divisa 47, 31

dotfleini 53

dollfusi, Balanus 66

dollfusii, Megabalanus 68

dolosus 49

domingensis 58

dorbignii 68

dormitor 45

dumortieri 47

duploconus 58

durhami 50

duvergieri 49

dybowskii 64, 101

ebumeus 63, 100, 101, 102

ecaudatum 40

echinata 53

echinicola 69

echinoplacis 51

ecuadoricus 69

elegans, Stomatolepas 44

elegans, Tetraclita 48

elizabethae 51

Elminius 52, 46, 16, 23

elongatum 57, 55

Emersonius, -inae ... 44, 11, 13, 21, 37

emkweniensis 51

engbergi 51

Eobalanus 60

Eoceratoconcha 55, 23, 28

Eoverruca 11

Epopella 46, 21, 22, 33

eseptatus 61

estrellanus, Balanus 61

etruscus 69

euamaryllis 50, 100, 102

Eubalanus 69

Euraphia, -inae 40-41, 11, 17-20,

36, 27, 31, 32

euspinulosa, -um 57, 100

evermanni 50

eyerdami 61, 102

fallax 51

fenestrata 53

ficarazzensis 45

filigranus 49

finchii 61

fischeri 53

fissus 42, 102

fistulosus 67

flexuosa 53

floridana. Ceratoconcha 58

floridana, Tetraclita 48

flosculus 52

sordidus 52

flosculoidus 69

flos 61

fluminensis 62, 101

folliculus 55

foraminifera 53

formae 53

formosana, Tetraclita 48

formosanus, Balanus 62, 101

fossata 53

fossilis, Balanus improvisus 64

fossilis, Balanus laevis 65

fosteri 46

fragilis, Chthamalus 42, 31

fragilis, Conopea 55

franciscanus 62, 101

fuchsi 59

funiculorum 49

fujiyama 49

fujiyamaformis 49

galapaganus SS

galeatus, -a. 55, 30

georgiana 49

giganteum, Megabalanus 68

105

giganteum. Pachylasma 40

gilmorei 43

gizellae, Balanus 69

glaber. 41

glandula 60, 28, 30, 102

glans 53

globicipitis 45

glyptopoma 61

gonioporae (see orbicetlae) 100

grandis. -e 5S, 31, 100

granulatus, -a 55

gregarea, -ius, Cantellius 57

gregaria, Acasta 53

gregarius, Radiolites, Tamiosoma . . 61

gregarius, Balanus 61

gryphicus 64, 101

halomitrae 58

hameri 50. 100, 101, 102

hammeri (=hameri) 50

hantkeni 49

hawaiensis, Solidobalanus 51

hawaiiensis. Balanus 62, 100,

101, 102

helenae 62

hembeli 4i, 13, 19, 31, 100

hemisphaerica 43

hentscheli 47

hertleini 45

herzi 62, 101

(Hespenbalanus). 51-52, 23

hesperius 51

laevidomiformis 51

laevidomus 51, 30

nipponensis 51

heteropus 61

hexastytos 44

ichthyophila 44

IHexacreusia) 49, 50, 23

Hexelasma 46, 40, 50, 1 1,

14-17, 20. 21

Hexelasminae 46, 21, 37

Hiroa 57, 23

hirsuta 53

hirsutum 46

Hoekia 58, 23, 24, 34

hoekianus, -um 50

hohmanni 69

honti 68

hopkinsi, Balanus 63

humilis, Balanus 69

hungaricus, (Balanus) 62

hungaricus, Megabalanus 68

hyastina 47

hystrix 66

ichthyophila. 44

idiopoma 53

imbricatus 40

imperator 46, 20, 31

imperatrix 42

improvisus 63, 62, 100, 101, 102

assimilis 64, 101

fossitis 64

gryphicus 64, 101

inclusus 49

indicum, Creusia 57-58

indicum, Pyrgoma 57-58

merulinae 58

symphylliae 58

indicus, Balanus 61, 101

indicus, Platylepas 44

inexpectatus 62-63. 101

insignis 63, 64, 101

integrirostrum 40

intermedia 40, 100

intermedius 68

intertextus, -a 41, 27, 32

investitus, -a 55

irregularis 69

Isolde 68

isseti 47

iwayama 57

japonica, Acasta 53

japonica, Diadema 45

japonica. Megabalanus 67

japonica, Pyrgoma 58

japonica, Tetraclita 48

japonicum, Pachylasma 40

javanicus 68

jedani 55

Jehlius 43,18

jungi 59

kanakoffi 65

karakumiensis 63

karandei 47

Kathpalmeria 49,23

kingii 52

kleinii 45

kojumdgievae 58

komaii 54

kondakovi 63, 101

krakatauensis, Chthamalus 4i,42

krakatauensis, Megabalanus 68

krambergeri 59

krugeri, Balanus 64,101

krugeri, Chirona 50

krugeri, Platylepas 44

krusadaiensis 53,100

kugleri 55

kuri 58

laevidomiformis 51

laevidomus 5i,30

laevigata 54, 58

laevis, Balanus 65,28,34

coquimbensis 65

fossilis 65

nitidus 65

nonsulcatus 65

laevis, Chioma 50

laevis, Eliminius 52

laguairensis 65

latum 59

leganyii 68

leonensis 65

leptoderma 46, 33

libera 54

ligusticus 42

litoralis 63

lobatobasis 43

longibasis 55

longirostrum 53

krusadaiensis 53,100

macsotayi 45

maculatus 50

madrasensis 56, 100

madreporicola, Acasta 49

madreporum, Cantellius 57

madreporarae, Boscia 59

major 45

malayensis, Balanus 64.101

malayensis, Chthamalus 42,41,

100,102

maldivensis 5J, 50

manati 43

crenatibasis 43

lobatobasis 43

maroccana 64, 101

mastignotus 51

maxillaris 67,68

maxima 41

Megabalanus 67-69,13,23,28,

29,30.31,34

Megatrema. 59

membranacea 54

Membranobalanus 52,23

merklini 63

merrilli 55

merulinae 58

Metabalanus 50

mexicanus 61, 101

microforamina 54

microstomus 69

microtretus 42

milensis 51

milleporum, Savignium 57

milleporosa, Tetraclita 48

minuta, Ceratoconcha 59

minutus, Balanus 65, 101

miocaenica, Ceratoconcho 59

miocenicus, Actinobalanus 49

mirabilis, Balanus 69, 101

mirabilis, Balanus perforatus 67

mitra, Tetraclita 48

modestus, Balanus 65, 101

modestus, Elminius 52, 100,101

laevis 52

mojbergi 55

molluscorum 52

monticutariae 58,34

moro 41,42,100

moravica 58

morycowae 68

multicostata, Tetraclitella 47

multicostatum, Pyrgoma 59

multidecorata 44

multiseptatus 68

murata 45

muricata, Acasta 54

muricata, Stephanolepas 44

mylensis 51

nascanus 51

natalensis 45

navicula 55

nebrias 53

nefrens 49.30.31

neogenica 59

neuseelandicus 41

Newmanella 47,21

nigrescens, Megabalanus 68

nigrescens, Tetraclita 48

nipponensis, Chthamalus 41

nipponensis, Solidobalanus 51

nipponensis, Tetraclitella 46

nitida, Acasta 54,34

nitidus, Balanus 65

nivea, Chirona 50

niveus, Balanus.. . 65,64,62,31,100,101

Nobia 58,23,31

nonstriatus 65

nonsulcatus 65

noszkyi 59

Notobalanus 52,10,23

nubilus 60-6;,69,70,34,100,101

nubilus, group of Balanus 60-61,

69,70,23

obscurus 65. 101

obliquus 66

oblitteratus 43,32,100

occator 68,66,100

occidentatis 51

106

ochlockoneensis 66

octavus 57

Octomeris 40,17-19,31

oligoseptatus 61

ophiophilus 44

oppidieboraci 64

orbicellae 58

orcutti 53

orcuttiformis 53

oryza 49

oulastreae 59

Pachydiadema 40,17

Pachylasma, -inae 40,11,14,16-19,

22,29,31

pacified, Tesseropora 47,33

pacificus, Balanus 66,68,101

brevicatcar 66, 101

prebrevicalcar 66, 101

Paleocreusia 58

palaoensis 49

pallidas, Cantellius 57

pallidus, Balanus 64, 101

krugeri (see kondakoui) 101

stutsburii 64

panamensis, Balanus 102

panamensis, Chthamalus 42

panamensis, Tetraclita 48

pannonicus 69

pantanelli 49

parahesperius 51

parkeri 66

patellans, Balanus 64,100,101

patellaris, Tetraclita 48,100

patelliformis (see B. patellaris) . . . . 101

patula 45,44,32,101,102

dentata 44

pectinipes 54

peninsularis 68,69

pentacrini 52,34

perfecta, Tetraclita 48

perforatus, Balanus 66-67,100,101

altavellensis 67

angustus 67

chordatus 67

cranchii 67

fistulosus 67

mirabilis 67

perforatus, group of Balanus 66-

67,23

permitini 42

peruvianas 63,101

phineus 51

pictus 62

pilsbryi, Euraphia 41

typica 41

neuseelandicus 41

pilsbryi, Catophragmus 40

pilsbryi, Tessarelasma 46

pilsbryi, Tetraclitella 47

Platylepas 44,21

Platylepadinae 44-48,49,11,21

playagrandensis 64

plicatus, Actinobalanus 49

plicatus, Epopella 46

plicatus, Megabalanus 68

pliocenicus 66

poecilosculpta 63,64,101

poecilotheca 64, 101

poecilus 66

Pollicipes 17

polygenus 65

Polylepas 45

polymerus 40,31

polyporus 61

porata 54

porcatus 59

porosa 48

communis 48

nigrescens 48

viridis 48

praegustator 44

praespinulosa 58,59

prebrevicalcar 66

prefloridana 59

proinus 51

projectum 58, 100

proteus 61

Protobalanus 60

proripiens 55

Proverruca 11

provisoricus 66

pseudauricoma 51

Pseudoacasta 54,23

pseudopallidum 57

psittacus 68,31,34,101

chilensis 68

pugetensis 59,60

purpurascens 47,46,100,102

darwini 47

nipponensis 46

purpurata 54

Pycnolepas 11

pygmaeus, -a 55

Pyrgoma 58,55,57,59,23

Pyrgomatidae, -inae 58,11,13,23,

24,28,39,31

Pyrgomina 59

Pyrgopsella 58,23

Pyrgopsis 58

quadrivittatus 49

quadratoradiata 59

quarta 59

quinquevittatus 49

quintus 57

radiata 47,68

wagneri 47

radicifer. 50

Radiolites 61

rafflesi 63,101

ramosa 44

rangi 59

latum 59

raphanoides 61

rariseptatus 61

regalis 61, 101

reginae 45

remi 55

reticulatus 64,101,102

revilei 69

rhachianecti 45

rhizophorae 47,40

roonwali 53,100

rosa, Megabalanus 68

rosea, Chirona 50

rosea, Tesseropora 47, 100

rostratus 6i, 100

alaskensis 61

apertus 61

heteropus 61

dalli 61

rubescens, Balanus 61

rubescens, Tetraclita. 48

rufotincta. 48,31,100

rugosus 46

salaami 64

saltonensis 62,101

sanctacrucensis 59

sauntonensis, Balanus 69

sarda 54

Savignium 57,23

scabrosus 42

scandens 55

schafferi 54

Scillaelepas 17

scrutorum 61

sculptura 54

scutelliformis 43

scuticosta 54

scutistriata 40,31

secundus 57

seguenzai, Megabalanus 68

seguenzai, Boscia 59

Semibalanus, -inae 55-56,70,

11,22-24,38,25,28,30

semicanaliculatus 49

semota 54

Septimus 57

serrata, Acasta 54

serrata, Tetraclita 47, 100

sextus 57

shilohensis 69-70

similus 70

simplex 46

sinensis 61, 101

sinnurensis 56, 100

sinuatus 52

snelliusi 57

socialis 51

solida, Chelonibia 44

solidus, Solidobalanus 51

Solidobalanus 50-51,69,70,23,30

S. (Bathybalanus) 52,23,34

S. (Hesperibalanus) 51-52,23

S. (Solidobalanus) 50-51,23

sookensis 51

sordidus 52

southwardi 46, 108

spinifera, Acasta 54

spiniferus, Balanus 61

spinitergum 54

spinosa, Acasta 54

spinosus, Megabalanus 68

spinulosa 58,56,57,59,100

spongicola 66

pliocenicus 66

spongites 54, 100

sporillus 54

squamosa 48,47,102

depressa 48

formosana 48

japonica 48

milleporosa 48

panamensis 48

patellaris 48, 100

perfecta 48

rubescens 48

elegans 48

rufotincta 48, 100

viridis 48, 100

stalactifera 48

con finis 48

floridana 48

milleporosa 48

stellaris 49

miocenicus 49

stellula 58

stellatus. . . 42,40,41,43,13,100,101,102

bisinuatus 43

comutus 43

107

thompsoni 43

stenonotus 51

Stephanolepas 44,21

stokesii 59

Stomatotepas 44,21

straeleni 50

striata, Acasta 54

striata, Tubicinella 45

(Striatobalanus) 50

stubbingsi 57

stuchburii 40

stultus 68

morycowae 68

sturi 59

stutsburii 64,101,102

subalbidus 64,101

sublaevis 50

subquadrata, -us 47

sulcata, Acasta 54

anchoris 54

spinosa 54

sulcata, Octomeris 40.31

sumbawae 57

suturalis 61

suturaltus 64,101

symphylliae 58

taiwanensis 50

talquinensis 61

tamiamiensis 61

Tamiosoma 61

tanagrae 68

tantillus 51

tenuis 50

terebratus 49-50,34

radicifer 50

tenuivalvata 54

Tessaretasma 46,17,21

tesselatus 64,101

Tesseroplax 47,21

Tesseropora 47,21,22,33

testudinaria 44, 102

solida 44

Tetrabalanus 65,23

Tetrachthamalus 43,18,32,100

Tetraclita 46-48,11,13,

20,21.28,29,31
Tetraclitella, -inae. . . 46,47,11,21,38,31

Tetraclitidae. -inae 47,48,11,13

19-21,24,37,38

Tetrachaelasma 46,21.22,108

thompsoni, Chthamalus 43

thompsoni, Solidobalanus 51

tintinnabulum SS-69,61,66

100.101,102

communis 68

coosensis 61

occator 66,100

peninsularis 69

tongaensis 6,3

trachealis 45

transversa 44

transversalis 57

transversostriatus 69

transsylvanicus 69

tredecimus 57

tridacophylliae 57

triderma 46

trigonus 6630,100,101,102

trigonus, group of Balanus. . . 65-66,23

trolli 59

truncatus 49

tuberculatus 50

Tubicinella 45,21

tuboperforatus 70

tubulatus 69

tulipa 54

tulipiformis 69

arenarius 69

etruscus 69

tumorifer 70

typica, Euraphia 41

typica, Megabalanus 67

typica, Pyrgoma 57

uliginosis 64,65,101

umitosaka 54

unguiformis 50

undulata 54

unisemita 47

vadaszi 62

validus 69, 100

variegatus 64, 100,101

cirratus 64-65,63,101

tesselatus 64,101

varians, Balanus 41

varians, Chirona 50

varians, Solidobalanus 52

velutinum 46

veneticensis 70

venezuelensis 69

venustus 65,64.101,102

modestus 65, 101

niveus 65, 101

obscurus 65, 101

Verruca, Verrucomorpha 11,

12,15,16

vesiculosus 69

vestitus 52

vialovi 52

vinaceus 69

violaceus 70,62,101

viridis 48.100

vitiata 48, 100

vladivostokensis 63,65,101

volcano 6968.100

vulgaris 45

wagneri 47

wilsoni, Megabalanus 69

wilsoni, Platylepas 44

withersi, Balanus 60

withersi, Chthamalus 40,100,102

withersi, Euraphia 41, 19,100

wireni 47

africana 47, 100

pacifica 47

Xenobalanus 45,21,31

zealandicus 50

zebra 39

zuiho 54

108

Figure 17. The remains of Tetrachaelasma sp., blanket the sea floor at a depth of nearly
2000m on the flanks of a seamount off Madagascar (26°29'S, 46°07'E). The relatively
primitive balanomorphoid Tetrachaelasma southwardi was first discovered by the R/V
Eltanin in the Antarctic Basin, off southern Chile and off Cape Horn at comparable depths
(Newman and Ross, 1971). It is the only balanomorphan known to occur in the abyss. The
calcareous deposits depicted here, composed of more than 90% calcitic barnacle remains,
including rostra up to 10 cm in length, represent the remains of animals that once Uved on
the seamount and were subsequently concentrated in the valleys and gorges around its
flanks. Other accumulations of comparable barnacle content occur in the fossil record, but
these developed in situ in shallow water. Photo courtesy of Robert L. Fisher, Scripps
Institution of Oceanography.

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ACME
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NOV 28 1984

100 CAM8r;10G£ STREET
CH^.RLESTOWN, MASS.

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