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Full text of "Memoirs of the San Diego Society of Natural History"

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 



Abel, O. 
1920. Lehrbuch der Palaozoologie, Gustav Fischer, Jena. 
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1964a. Some relations between habitat behaviour and 
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1975. Effects of local anesthetics and hemicholinium-3 
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1974. The effect of ecdysterone on cyprids of Balanus 
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1973. The biocenoses incrusting Eternit panels immersed 
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1974. On the nutrition and metabohsm of zooplankton. 
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1975. Tidally-produced internal bands in the shell of 
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longirostrum krusadaiensis, B. madrasensis, B. 
patellaris, B. roonwali, B. sinnurensis, B. tintin- 
nabulum tintinnabulum, B. t. occator, B. t. validus, 
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C hembeli, C. malayensis, C moro, C. steltatus, C. 
withersi, Creusia spinulosus euspinulosa, Octomeris 
angulosa, O. intermediia, Pyrgoma conjugatum, P. 
gonioporae, P. grande, P. projectum, Tetrachtha- 
malus oblitteratus, Tetraclita alba, T. coerulescens, 
T. purpurascens, T. rosea, T. squamosa communis, 
T. s. patellaris, T. s. rufotincta, T. s. serrata, T. s. 
viridis, T. vitiata, T. wireni africana) 



101 



Davis, C. W. and J. D. Costlow 

1974. Evidence for a molt inhibiting hormone in the 
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93(2);85-92. 

Devillez, E. J. 

1975. Observations on the proteolytic enzymes in the 
digestive fluid of the barnacle Balanus nubilus. 
Comp. Biochem. Physiol, A Comp. Physiol. 51(2): 
471-474. 

Dresdner, G. W., F. Ojeda and H. Hess-Ojeda 

1974. Microscopical structure of muscles from the 
operculum of the barnacle Balanus psittacus 
Molina. Zool. Anz. 192(1/2):15-21. 

Fyhn, U. E. H. 

1976. Holeuryhalinity and its mechanisms in a cirriped 
crustacean, Balanus improvisus. Comp. Biochem. 
Physiol. 53A: 19-30. 

Fyhn, U. E. H. and J. D. Costlow 

1975. Tissue culture of cirripeds. Biol Bull. 149(21:316-330. 
(Balanus amphitrite, B. ebumeus, B. improvisus) 

Ganapati, P. N. and D. R. K. Sastry 

1974. Record of Athanas indicus (Coutiere) (Decapoda: 
Alpheidael associated with Stomopneustes vario- 
laris (Lamarckl (Echinodermata: Echinoideal from 
Visakhapatnam Coast. Proc. Indian Nat. Sci. 
Acad., Part E., Biol. Sci. 38(5/61:367-372. (Balanus 
amphitrite amphitrite, B. trigonus) 

Gardner, D. and J. P. Riley 

1972. Seasonal variations in the component fatty acid 
distributions of the Upids of Balanus balanoides. 
J. Mar. Biol. Assoc. U. K. 52(41:839-845. 

Gorin, A. M. and A. M. Murakhveri 

1973. Seasonal dynamics of settling and growth of 
Balanus and Mytilus in the Peter the Great Bay. 
Ekologiia 4(2):86-89. (Balanus crenatus) 

Granier, J. 

1973. Le genre Balanus sur les cotes de Camargue et du 
Gard. Soc. Linn. Lyon Bull. Mens. 42(8):203-212. 
(Balanus amphitrite, B. crenatus, B. ebumeus, B. 
perforatus, B. trigonus, Megabalanus tintinnabulum) 

Henry, D. P. and P. A. McLaughlin 

1975. The barnacles of the Balanus amphitrite complex 
(Cirripedia, Thoracica). Zool. Verhandel. 141:1-254. 
(Balanus alatus, B. amphitrite albicostatus, B. a. 
aeratus, B. a. amphitrite, B. abeli, B. a. cirratus, B. 
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. 
a. poecilotheca, B. a. rafflesi, B. a. stutsburi, B. a. 
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. 
dybowskii, B. ebumeus, B. improvisus. B. i. 
assimitis, B. i. gryphicus, B. minutus, B. mirabilis, 
B. pacificus, B. pacificus brevicalcar, B. pallidas, 
B. pallidus krugeri, B. patellaris, B. patelliformis, 
B. aquila, B. reticulatus, B. subalbidus, B. sutural- 
tus, B. tintinnabulum maroccana, B. uliginosus, B. 
variegatus, B. v. cirratus. B. v. tesselatus, B. 
venustus, B. v. modestus, B. v. niveus, B. v. 
obscurus, B. violaceus, B. armatus) 

HUlaire-Marcel, C, G. Prichonnet and B. De Boutray 

1974. Marine Pleistocene facies from the hills at Oka, 
Quebec. Nat. Can. (Que.) 101(5):781-802. (Balanus 
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Hochstein, S., B. Minke and P. Hillman 
1973. Antagonistic components of the late receptor 



potential in the barnacle photoreceptor arising 

from different stages of the pigment process. J. 

Gen. Physiol. 62(1):105-128. (B. amphitrite, B. 

ebumeus) 
Holland, D. L. and P. J. Hannant 
1973. Addendum to a micro-analytical scheme of the 

biochemical analysis of marine invertebrate larvae. 

J. Mar. Biol. Assoc. U. K. 53(4):833-838. (Balanus 

balanoides, B. hameri, Elminius modestus) 
Houk, J. L. and J. M. Duffy 

1972. Two new sea-urchin-acorn barnacle associations. 
Calif. Fish and Game 58(4):321-323. (Balanus 
concavus pacificus, B. nubilus) 

Hoyle, G.. P. A. McNeill and A. I. Selverston 

1973. Ultrastructure of barnacle giant muscle fibers. J. 
CeU Biol. 56(1):74-91. (Balanus nubilus) 

Hughes, G. R. 

1974. The sea turtles of South-east Africa, II. Oceanogr. 
Res. Inst. (South Africa), Inves. Rept. 36:1-96. 
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Hurley. A. C. 

1975. The establishment of populations of Balanus 
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532. 

Ireland, M. P. 
1974. Variation in the zinc, copper, manganese and lead 
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Jocque, R. and D. Van Damme 

1972. Introduction to the ecological study of intertidal 
clay and peat banks at Raversijde Belgium. 
Biologisch Jaarboek, Belgium 39:157-190. (Balanus 
balanoides, B. crenatus, Elminius modestus) 

Kasymov, A. G., R. M. Bagirov and G. M. Fihppov 
1974. Benthos of the southeastern Caspian Sea coast. 
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Kidson, C. and R. Wood 

1974. The Pleistocene stratigraphy of Barnstable Bay. 
Proc. Geol. Assoc. 85:223-237. (Balanus balanoides) 

Krischer, C. C. 
1971. The photo-electric efficiency of the median and the 
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Kuznetsova, I. A. 

1973. Assimilation of some food kinds of cirriped crusta- 
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noides, B. ebumeus, B. improvisus) 

LaCombe, D. 

1973. Cria^ao de Balanideos em laboratorio. Trab. V 
Congr. Latinoam. Zool. Montevideo, 1:168-174. 
(Balanus amphitrite albicostatus, B. a. denticulata, 
B. a. hawaiiensis, B. tintinnabulum tintinnabulum, 
Chelonibia patula, Chthamalus stellatus) 

Larman, V. N. and P. A. Gabbott 

1975. Settlement of cyprid larvae of Balanus balanoides 
and Elminius modestus induced by extracts of 
adult barnacles and other marine animals. J. Mar. 
Biol. Assoc. U. K. 55:183-190. 

Long, E. R. 

1974. Marine fouling studies off Oahu, Hawaii, USA. 
Vehger 17(l):23-36. (Balanus amphitrite, B. crena- 
tus, B. ebumeus, B. tintinnabulum, B. trigonus) 

Magre, E. J. 

1974a. Population density of Balanus balanoides in rela- 
tion to tide pool water level. (Cirripedia Thoracica). 
Crustaceana 26(2):139-142. 

1974b. Ulva lactuca L. negatively affects Balanus 
balanoides (L.) (Cirripedia Thoracica) in tidepools. 
Crustaceana 27(3):231-234. 
Maurer, D. and L. Watling 

1973. Studies on the oyster community in Delaware: the 
effects of the estuarine environment on the associ- 



102 



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1974. DDT and the rugophilic response of settling 
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1975. Competitive relationship between Balanus amphi- 
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special reference to their larval settlement. Hydro- 
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Moore, H. B., H. B. Albertson and S. M. MiUer 

1974. Long-term changes in the settlement of barnacles 

in the Miami area. Bull. Mar. Sci. 24(1|:86-100. 

IBalanus amphitrite amphitrite. B. ebumeus. B. 

improvisus. B. reticulatus. B. trigonus) 
Nielsen, R. 

1972. A study of the shell-boring marine algae around 

the Danish Island Laeso. Botanisk Tiddsskrift 

67(31:245-269. (Balanus balanoides) 
Paine, R. T. 

1974. Intertidal community structure: experimental 

studies on the relationship between a dominant 

competitor and its principal predator. Oecologia 

(Berl.) 15(21:93-120. IBalanus cariosus. B. glandula, 

Chthamalus fissusi 
Partaly, E. M. 

1974. Seasonal changes of epibiotic communities on 

Balanus improvisus on overgrowth biocenosis. Zh. 

Obshchei Biol. 35(31:454-459. 
PiUai, N. K. and Balakrishnan Nair. 

1974. Observations on the incidence and seasonal fluc- 
tuations of certain crustacean larvae in the plank- 
ton of the southwest coast of India. Hydrobiologia 
43(3/4):443-461. IBalanus amphitrite communis) 

PoUock, L. W. 

1975. Observations on marine Heterotardigrada, includ- 
ing a new genus from the western Atlantic Ocean. 
Cah. Biol. Mar. 16(1):121-132. (Balanus balanoides) 

Pratt, D. M. 

1974. Attraction to prey and stimulus to attack in the 
predatory gastropod Urosalpinx cinerea. Mar. Biol. 
27(l):37-45. IBalanus balanoides. B. ebumeus) 
Rao, D. G. V. and P. N. Ganapati 
1972. Respiration in relation to salinity variation in inter- 
tidal barnacles. Proc. Indian Nat. Acad., Part B, 
Biol. Sci. 38(5/6):425-429. IBalanus amphitrite 
amphitrite. B. tintinnabulum tintinnabulum) 
Rogers, F. L. 

1948. Description of a new species of barnacle from 



Panama. Bull. Southern California Acad. Sci. 47(3): 
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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145