Antartica Cirripedia

ANTARCTIC
RESEARCH
SERIES
American Geophysical Union

ANTARCTIC
RESEARCH
SERIES
American Geophysical Vnion
Volume 1
BIOLOGY OF THE ANTARCTIC SEAS
Milton O. Lee, Editor
Volume 2
ANTARCTIC SNOW AND ICE STUDIES
Malcom Mellor, Editor
Volume 3
POLYCHAETA ERRANTIA OF ANTARCTICA
OlgaHartman
Volume 4
GEOMAGNETISM AND AERONOMY
A.H. Waynick, Editor
Volume 5
BIOLOGY OF THE ANTARCTIC SEAS II
GeorgeA. Llano, Editor
Volume 6
GEOLOGY AND PALEONTOLOGY OF THE ANTARCTIC
Jarvis B. Hadley, Editor
Volume 7
POLYCHAETA MYZOSTOMIDAE AND SEDENTARIA OF ANTARCTICA
OlgaHartman
Volume 8
ANTARCTIC SOILS AND SOIL FORMING PROCESSES
1. C. F. Tedrow, Editor
Volume 9
STUDIES IN ANTARCTIC METEOROLOGY
Morton 1.Rubin, Editor
Volume 10
ENTOMOLOGY OF ANTARCTICA
1. Linsley Gressitt, Editor
Volume 11
BIOLOGY OF THE ANTARCTIC SEAS III
Waldo L. Schmitt and GeorgeA. Llano, Editors
Volume 12
ANTARCTIC BIRD STUDIES
Oliver L. Austin, Jr., Editor
Volume 13
ANTARCTIC ASCIDIACEA
Patricia Kott
Volume 14
ANTARCTIC CIRRIPEDIA
WilliamA. Newman andArnoldRoss =
ANTARCTIC
Volume 14 RESEARCH
SERIES
Antarctic Cirripedia
Monographic account based on specimens

collected chiefly under the United States
Antarctic Research Program, 1962-1965
William A. Newman and Arnold Ross
Published with the aid of a grant from the National Science Foundation
PUBLISHER
AMERICAN GEOPHYSICAL UNION

O F THE
National Academy of Science-National Research Council

January 29, 1971
A N T A R C T I C
l ~ o l u m e14 R E sE A R cH
( S E R I E S
ANTARCTIC C I R R I P E D I A
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THE ANTARCTIC RESE14RCH SERIES
THE A\T-~RCTIL
RESEARCHSERIESis designed to provide a medium for presenting
authoritati~ereports on the extensile and detailed scientific research work being
carried out in Antarctica. The series lias been successful in eliciting contributions
from leading research scientists engaged in antarctic investigations; it seeks to main-
tain high scientific and publication standards. The scientific editor for each volume
is chosen from among recognized authorities in the discipline or theme it represents,
as are the reviewers on whom the editor relies for advice.
Eeginning uith the scientific investigations carried out during the International
Geophysical Year, reports of research results appearing in this series represent origi-
nal contributions too lengthy or other~+iseinappropriate for publication in the
standard journals. In some cases an entire volume is devoted to a monograph. The
material published is directed not only to specialists actively engaged in the ~ + o r but
k
to graduate students, to scientists in closely related fields, and to interested laymen
versed in the biological and the physical sciences. Many of the earlier volumes are
cohesive collections of papers grouped around a central theme. Future volumes may
concern themselves with regional as well as disciplinary aspects. or 1% ith a comparison
of antarctic phenomena ~ + i t hthose of other regions of the globe. But the central
theme of Antarctica nil1 dominate.
In a sense, the series continues the tradition dating from the earliest days of
geographic exploration and scientific expeditions-the tradition of the expeditionary
volumes which set forth in detail everything that Nas seen and studied. This tradi-
tion is not necessarily outmoded, but in much of the present scientific mork one
expedition blends into the next, and it is no longer scientifically meaningful to
separate them arbitrarily. Antarctic research has a large degree of coherence; it
deserves the modern counterpart of the expeditionary volumes of past decades and
centuries which the Antarctic Research Series provides.
With the aid of a grant from the Eational Science Foundation in 1962, the
American Geophysical Union initiated the Antarctic Research Series and appointed
a Board of Associate Editors to implement it. A supplemental grant received in 1966,
the income from the sale of volumes in the series, and income from reprints and
other sources have enabled the AGU to continue this series. The response of the
scientific community and the favorable comments of reviewers cause the Board to
look for^+-ard with optimism to the continued success of this endeavor.
To represent the broad scientific nature of the series, the members of the Board
were chosen from all fields of antarctic research. At the present time they include:
Eugene L. Boudette, representing geology and solid Earth geophysics; A. P. Crary,
seismology and glaciology; George A. Llano, botany and zoology; Martin A. Pome-
rantz, aeronomy and geomagnetism; Morton J. Rubin. meteorology; Waldo L. Schmitt,
marine biology and oceanography; and Laurence M. Gould, honorary chairman.
Fred G. Alberts, secretary to the U. S. Advisory Committee on Antarctic Names,
gives valuable assistance in verifying place names, locations, and maps. AGU staff
members responsible for the series are: Judith S. McCombs, managing editor. and
Jane Bruce, style editor.
MORTONJ. RUBIN
Chairman, Board of Associate Editors
Antarctic Research Series
PREFACE
THIS IS THE FIRST ATTEMPT to bring together what is known of the antarctic cirriped
fauna in one comprehensive treatise. It covers the systematics of the barnacles of
subantarctic and antarctic regions and their distribution, both recent and paleonto-
logical. A second part already in preparation will report on planktonic larvae.
Antarctic Cirripedia, the second monographic contribution in the Antarctic Re-
search Series, generally follows the pattern set by Kott7sAntarctic Ascidiacea (1969).
The study, as far as possible, covers all records made by previous expeditions to Ant-
arctic waters and culminates in this volume with the 1962-1965 cruises of USNS
Eltanin. The list of the principal vessels which have brought back cirriped materials
from antarctic regions is a notable one: Aurora, Belgica, Challenger, Discovery, En-
deavour, Ob', Pourquoi-Pas?, SGya, Terra Nova, Umitaka-Maru and William Scoresby.
Yet the number of benthic samplings of cirripeds made by Eltanin under the United
States Antarctic Research Program from south of the Antarctic Convergence exceeds
the total of those returned by all earlier expeditions.
In accordance with procedures established under the U.S. Antarctic Research
Program, biological materials from terrestrial and marine activities forwarded to the
Smithsonian Oceanographic Sorting Center are made available to qualified individuals
recommended by the Advisory Committees for distribution of specimens to specialists.
William A. Newman of the University of California, San Diego, was the specialist
entrusted with the Eltanin cirriped collections in 1966. He was joined by Arnold
Ross, Natural History Museum of San Diego, California, as junior author.
Altogether, Newman and Ross have examined 85 species, 29 genera, and 9 families
of cirripeds. Of these 20 species, 9 genera and 1 family are described as new. The
text is illustrated by 48 plates, 90 text-figures, 11 charts, and 3 tables. All the Cirri-
pedia from the Antarctic are figured and described with keys to each; charts are used
to show their present geographical distribution. Paleogeographical and biogeographi-
cal aspects of the antarctic cirriped fauna are discussed in context with related forms
throughout the world. The report includes the extraordinary discoveries of a new
gigantic, sessile barnacle, the deepest balanomorphan known, and the first deep-water
acrothoracican; the remarkably complex and uniquely constructed shell walls of the
former are illustrated in three color plates.
GEORGEA. LLANO
Member, Board of Associate Editors
ACKNOWLEDGMENTS
THE RESOIJRCES O F M A \ Y INSTITLTIOAS AND COLLEAGLES have been drar+n upon during
the course of this studj. Initial arrangements for the cirripeds collected by the LSYS
Eltanin were made ~ i t Dr.h R. B. Tibbj. Universit) of Southern California. under the
auspices of the United States Antarctic Research Program ( U S A R P ) . The Eltanzn
collections mere processed largely b j the Smithsonian Oceanographic Sorting Center.
Washington, D. C.. under the directicn of Dr. H. Adair Fehlman. ~ i t the h assistance
of Mrs. Beatrice Burch and her successor. Miss P. A. McLaughlin. The efficient and
enthusiastic services of the SOSC have been much appreciated.
We are especiallj indebted to Dr. Thomas E. Bouman. Smithsonian Institution, for
providing t j p e specimens and for critical selection of comparative materials housed
in the collections of the United States Kational Museum. Many thanks are due Miss
Elizabeth Pope of the Australian Museum for the loan of specimens collected by the
Aurora; Mrs. Patricia Lofthouse-Barker of the British Museum (Aatural History),
for the loan of specimens taken by the Challenger; Dr. C. A. Fleming of the h e w
Zealand Geological Survey. u h o made available the fossil specimens of Hexelasma
az~cklandicumupon ~ j h i c hthe late T. H. Withers based his descriptions; Dr. I. G.
Speden for providing specimens described in his ~ o r kon antarctic Pleistocene fossil
deposits. presently housed in the New Zealand Geological Survey collections; Dr. Jan
Stock, Zoological NIuseum, Amsterdam. for the loan of paratypes of Bathybalanus
Hoek; and Dr. Huzio Utinomi. Seto Marine Biological Laboratory. Japan, for making
arrangements with Dr. Jiro S&no.Tokyo University of Fisheries, for us to borrow the
JARE specimens he described. Special thanks are due Dr. A. J. Southward, The
Laboratory. Pljmouth, England. for the remarkable photographs of living and pre-
served specimens of Hexelasma hirsutum, and for the kindness of comparing this
species with type material of H. aucklandicum and H. corolliforme housed in the
British Museum (Natural History).
For the loan or donation of other specimens that materially aided this study. we
thank Dr. H. G. Stubbings. Admiralty Materials Laboratory, England; Dr. Brian
Foster, Lniversity of Auckland, New- Zealand; Dr. J. T. Tomlinson. San Francisco
State College. California; Dr. Victor A. Zullo, Marine Biological Laboratories, Woods
Hole; Dr. J. A. Bullivant, New Zealand Oceanographic Institute; Norman E. Weisbord,
Florida State University, Florida; Dr. Gilbert Rowe, Duke University, North Caro-
lina; and Dr. Torben Wolff, Universitetets Zoologiske Museum, Denmark.
The able bibliographic work by Mrs. Carol Kourtz-Platt, the inking of drawings by
Mrs. Sally Beer, drafting by Howard Shirley, portions of the photographic work by
Ronald Lam and Miss Nancy Ford and the stenographic assistance by Mrs. Cecelia
Ross during the initial phase of the work were much appreciated.
Support under a grant entitled Cooperative Systematic Studies in Antarctic Biology
(GA-1374) from the National Science Foundation through Dr. I. E. Wallen, Director,
Office of Environmental Sciences, for systematic studies of biological materials col-
lected under USARP made this work possible and is gratefully acknowledged.
WILLIAMA. NEWMAN ARNOLDROSS

Scripps Institution of Oceanography Natural History Museum
La Jolla, California 92037 San Diego, California 92112
TABLE OF CONTENTS
The Antarctic Research Series . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v
Preface . . . . . . . . . . . ................................... vii
Ackno~~ledpments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Historical Account . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Biogeographical Account . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Contemporary Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Systematic Account . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Key to Recent Antarctic Cirripeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Order Ascothoracica Lacaze-Duthiers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Synagogidae Gruvel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Ascothorax Djakanov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Order Acrothoracica Gruvel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Suborder Pygophora Berndt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Family C r y p t ~ ~ h i a l i d aGerstaecker
e ...........................
Genus Cryptophialus Darwin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Key to the Austral Group of Cryptophialus . . . . . . . . . . . . . . . . . .
Family Zapfellidae Saint-Seine ................................
Genus Brachyzapfes Codez . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Order Thoracica Darwin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Suborder Lepadomorpha Pilsbry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Family Lepadidae Darwin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Key to Antarctic Lepadidae .................................
Genus Lepas
. ,
Linne ........................................
Subgenus Lepas s.s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Subgenus Dosima Gray . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Conclzoderma Olfers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family Scalpellidae Pilsbry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Key to Recent Antarctic Genera of Scalpellidae . . . . . . . . . . . . . . . .
Genus Smilium Leach . . . . . . . . . . . . . . . . . . . . . . . . . .
. ........
Genus Euscalpellum Hoek . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Genus Cretiscalpellum Withers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Zeugmatolepas Withers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Arcoscalpellum Hoek . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Key to Antarctic Species of Arcoscalpellum . . . . . . . . . . . . . . . . . .

xi
Key to Genera Allied to Neoscalpellum . . . . . . . . . . . . . . . . . . . . . . 93
Genus Neoscalpellum Pilsbry . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Genus A b a t h e ~ c a l ~ e l l u mnov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
Genus Gymnoscalpellurn nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Genus Litoscalpellurn nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
Genus Mesoscalpellum Hoek . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
Genus Annandaleum nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
Genus Scalpellum Leach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
Key to Species-Groups of the Genus Scalpellum . . . . . . . . . . . . . . 123
Genus Australscalpellurn nov . . . . . . . . . . . . . . . . . . . . . . . 130
GenusBrochianov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
Suborder Verrucomorpha Pilsbry . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Family Verrucidae Darwin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
Genus Verruca Schumacher . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Key to Subgenera of Verruca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Subgenus Altiverruca Pilsbry . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Suborder Balanomorpha Pilsbry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Key to Families of Balanomorpha . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Family Bathylasmatidae nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Families of the Balanomorpha: Comparative Aspects . . . . . . . . . . 139
Key to Genera of Bathylssmatidae . . . . . . . . . . . . . . . . . . . . 142
Genus Bathylasma nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Key to Species of Bathylasma . . . . . . . . . . . . . . . . . . . . . . . . . 143
Genus Tetrachaelasma nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
Genus Tessarelasma Withers . . . . . . . . . . . . . . . . . . . . . . . . . 155
Genus Hexelasma Hoek . . . . . . . . . . . . . . . . . . . . . . . . . . 155
Key to Species of Hexelasma . . . . . . . . . . . . . . . . . . . . . . . . . 135
Genus Aaptolasma nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Key to Species of Aaptolasma . . . . . . . . . . . . . . . . . . . . . . . . 158
Family Balanidae Leach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
Genus Balanus DaCosta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
Subgenus Austrobalanus Pilsbry . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
Subgenus Chirona Gray . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
Subgenus Bathybalanus Hoek . . . . . . . . . . . . . . . . . . . ..
. . . ..... 173
Subgenus Balanus DaCosta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
Subgenus Megabalanus Hoek . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
Genus Coronula Lamarck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
Genus Xenobalanus Steenstrup . . . . . . . . . . . . . . . . . . . . . . . . . 180
Order Rhizocephala Miiller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Suborder Kentrogonida Delage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123
Family Peltogastridae Lilljeborg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Genus Briarosaccus Boschma . . . . . . . . . . . . . . . . . . . . . . . . . . ..
.
. 183
xii
Appendices
1. Localities arid Bathjmetry of Previously Known Antarctic Cirripedia
2. Summary of Stations Yielding Specimens Covered in This Study
3. Eltaniri Cirripedia Listed According to Station
4. Eltanin Cirripedia Listed According to Depth
5. Rathymetric Distribution for Sampling Devices llsed during El~anzn
Expeditions Covered in This Repor t
References
Glossary . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . .
Plates .................... ...... . . . . . . . . . . . . . . . . . . . . . . . . ...
Plate Legends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Indices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Systematic . . . . . . . . . . .. . . . . . .. . . . . . . . . . . . . . . ... . . . .
Subject
Antarctic Research Series Antarctic Cirripedia Vol. 14
INTRODUCTION
This report is concerned primarily ~ i t the
h Cirripedia rounding oceans, forming the northern portion of the
of the antarctic region. Cirripeds are benthonic and West Wind Drift, and this zone forms the Antarctic
pelagic crustaceans occurring in all seas, and those of Convergence. The water along the Convergence under-
the oceans surrounding Antarctica are poorly k n o n. ~ goes downwelling that has been said to form the
The antarctic region is remarkably different from the sharpest boundary of any known faunal province
Arctic in centering upon land rather than an ocean. (Broch, 1961). Surface waters continue to move into
This polar continent is generally considered to support the Convergence while relatively warm midwaters
some 90% of the world's ice, which, if melted, would move south to higher latitudes before being drawn to
be sufficient to raise the level of the world's oceans the surface to replace them. The Convergence, al-
some 8.0 meters. The ice of course has a profound though not fixed as indicated in the charts utilized
effect on the biology of the land. This effect is equally herein, tends to remain around 60°S in the Pacific-
profound along the shores to depths of at least 100 Antarctic sector and around 5 0 " s in the Atlantic and
meters. The chilling of the waters there causes them to Indian Ocean sectors.
descend and creep out into the depths of the surround- While this report is concerned primarily with the
ing seas. The antarctic continent has not always been cirriped fauna south of the Antarctic Convergence,
so severely glaciated, nor is it believed to have always many species found there also occur in the oceans to
been separated from adjacent continents. Coal seams, the north, and of necessity they must be taken into
reputed to have been formed some 150 million years consideration if an attempt to understand the past, as
ago, are ample proof of milder climates, and the con- well as the present, is to be made.
cept of Gondwanaland and its fragmentation by con- Antarctica presently has no land connections with
tinental drift and sea-floor spreading, beginning some other continents. If there were connections more than
250 million years ago, is currently gaining acceptance. 100 million years ago, they would have little bearing
The hydrography of the cold temperate and antarctic on the content of this study, since its historical aspects
regions has been thoroughly discussed by Deacon and began in the upper Cretaceous. While Antarctica pres-
Knox (in Pantin, 1960). Antarctic coastal waters are ently has no land connections to the north, it is con-
generally near freezing (-1.86OC). Seasonal freezing nected by several submarine oceanic rises and ridges
precludes a sedentary shore fauna (Knox, op. cit.). that extend, in several cases, well above the deep-sea
Furthermore, it apparently accounts historically for a floor (chart 1 ) . At least two of these appear signifi-
depauperate sublittoral cirriped fauna to depths of cant to the material covered herein. One actually con-
about 100 meters, and freezing of the benthos to depths sists of two routes from South America to Antarctica:
of 33 meters presently occurs seasonally in McMurdo the Scotia Ridge running from Cape Horn eastward
Sound (Dayton et al., 1969). The fauna then is for the before looping south to connect with the Antarctic
most part pelagic, or benthonic in relatively to ex- Peninsula, which forms the western boundary of the
tremely deep water. Weddell Sea; and the Albatross Cordillera with its
The general near-shore currents around the con- various avenues to the Americas connecting with An-
tinent are counter-clockwise and constitute the East tarctica via the Macquarie Rise, relatively near and
Wind Drift. On the other hand, the greater part of more or less an extension of the western rim of the
the off-shore surface waters are driven clockwise Ross Sea. The second connection not only joins An-
around the continent as the West Wind Drift. The tarctica to the Americas but also to the Indo-Pacific
antarctic portion of the West Wind Drift tends north- faunal region, via Australia, and to a lesser degree,
ward to mingle with subantarctic waters of the sur- New Zealand. Oceanic rises apparently serve as sub-
Copyright American Geophysical Union

2 ANTARCTIC CIRRIPEDIA
marine causeways for midwater migrations, and their Although the Eltanin has worked primarily in ant-
presence or absence should affect faunal composition arctic waters, cirripeds from earlier cruises to the
somewhat independently of surface conditions. No Arctic, and to the trenches bordering the Pacific coast
doubt this has been the case in Antarctica. of South America, are included herein. Moreover, ap-
While knowledge of the cirriped fauna of Antarctica propriate collections from elsewhere have been utilized
began with Darwin (1851), because of its deep-water for comparative purposes in an attempt to understand
nature the first important contributions were not made better the relationship of the antarctic fauna to other
until 1883. They then appeared in Hoek's report on faunas of the world (chart 2 ) . These collections con-
the Cirripedia of the Challenger Expedition. Subse- sist, notably, of specimens taken by the U. S. Fisheries
quent explorations and contributions are discussed in steamer Albatross, the Japanese Antarctic Research
the next section, bringing us to the materials covered Expedition vessels S6ya and Umitaka-Maru, the Duke
in the present report, primarily taken by the USNS University RV Eastward, the New Zeland Oceano-
Eltanin. The role of the USNS Eltanin in antarctic graphic Institute vessel Endeavour under the New
waters has been summarized by Sandved (1966), and Zealand Antarctic Research Programme. the Woods
the scientific work carried out along the Pacific coast Hole Oceanographic Institute vessels Chain and Atlan-
of South America by Menzies (1963). Scientific re- tis 11, the Tui Expedition, the HMS Challenger, the
ports published as an outgrowth of the Eltanin cruises British Antarctic Expedition vessel Terra ATova, and
are cited in Sandved's paper and the Antarctic Bibliog- the Dutch man-of-war Siboga.
raphy prepared by the Library of Congress, published Institutional and expeditional abbreviations used
by the U. S. Government Printing Office. throughout this study are as follou s :
1 Scotia Ridge 7 Albatross Cordillera 12 Mid-Indian Ridge

2 Albatross Cordillera 8 Macquarie Rise 13 Atlantic - Indian Ridge
3 Cocos Ridge 9 South Tasmania Rise 14 Walvis Ridge
4 Cornegie Ridge 10 Southeast Indian Rise 15 Mid-Atlantic Ridge
5 Solo y Gornez-Nazca Ridge 11 Kerguelen Plateau 16 Southeast P o c i f ~ cBasin
6 Chile Rise
Chart 1. Oceanic ridges and rises.

Chart 2. 1,ocalitics for cirriped specimens talcen by the Eltunin (hcnthir, closed circ:lcs; sea surfa(,c, open circle?),
and for other materials (triangles) utilized in this rcport.
Auckland University AU Ncw Zraland Antarctic Rcscarch Expedition NZARP

Australian Antarctic Expedition AAE New Zealand Geologic,al Survry NZO S
British-Australian-New Zealand Antarctic New Zealand Oceanographic: Tnstituic &ZOI
Expedition BANZARE Scripps hlsiitution ol Oceanography NO
I3ritisli Museurn (Natural History) EM ( N H ) United States Antarctic Research Program USARP
1)ukc University Rlarinc Riology Laboratory UIJMnT, United States Eurcau of Cnmmercial I'isl~eries USRCF
.lapanese Antarclie Rrscarch Exlredition JilRE IJnited Statcs National blusrurn USNh2
hlarine Giological Association Unitcd Kingdom hlGAUK Woods Holc Oceanographic: Tnsiilution WkIOI
During the course of this study, Me were hampered We have attempted to produce descriptions at least as
to a certain degree by what can fairly be called an comprehensive as contemporary standards require, but
overly conservative and antiquated classificatiori of no description can ever be truly complete. In an
thoracicarl Cirripedia, and we found it necessary to attempt to obviate this difficulty, pertinent materials
revise certain groups. Materials have not permitted Irawe been illustratrd as well as described, and keys to
the revisions to be as extensive as desired, hut the areas all species known in antarctic waters provided. This
of difiiculty have become more clearly delimited and monograph includes G5 species, 29 genera, and 9 fami-
will be the u o r k of further studies. Many of the lies, of which 20 species, 9 genera, and one family
problems are the result of very inadequate descriptions.

HISTORICAL ACCOUNT
The earliest sightings and mapping of Antarctica lack The cirripeds collected during the British Antarctic
documentation, but indirect evidence suggests there Expedition aboard the Terra Nova formed the basis of
was knowledge of this great southern continent prior Borradaile's study on this unusual fauna (1916). Of
to the year 1513 (Hapgood, 1962). Documented sight- the several species he discovered, subfossil material
i n g ~took place early in the 19th century, and to date from a glacier, Hexelasma antarcticum, proved to be
more than 150 voyages returning with scientific infor- one of the most perplexing encountered in developing
mation have taken place in antarctic waters (Caras. the present report.
1962). Of the many subsequent expeditions to the Ross Sea
Studies on the cirriped fauna of the South Atlantic, sector, only one resulted in a report devoted to the
South Pacific, and Indian Oceans begin with the voy- Cirripedia of the region (Bage, 1938). The specimens
age of Charles Darwin, naturalist aboard HMS Beagle. described were collected by the Aurora from Common-
Although the Beagle did not enter waters south of the wealth Bay, and the report included two previously
Antarctic Convergence, Darwin discovered at least two known and one new species.
species which extend south of this boundary. For Carl August Nilsson-Cantell stands among the prin-
more than 11 decades Darwin's studies (1851, 1854) cipal contributors to our knowledge of the antarctic
have served as a basis for all biogeographical, ana- cirriped fauna. His first report was concerned with
tomical, and evolutionary work on cirripeds. specimens collected by S. Vallin in the Ross Sea and
The Challenger ventured south of the Antarctic Con- was published in 1926. This was followed by a note on
vergence, but only for a short while. The cirripeds the fauna of Stewart Island and South Georgia
collected during the entire expedition were studied by (1930a). The Thoracica collected by the RRS Discov-
the Dutch scientist, P. P. C. Hoek, and the results ery I and Discovery 11, as well as the RRS William
appeared in two separate reports in the series issued Scoresby, provided the material for Nilsson-Cantell's
by the Challenger Office (1883, 1884). Six of Hoek's major contributions to the knowledge of the antarctic
antarctic species are recognized today. and southern faunas. To the previously known fauna
The cirripeds dredged by the SY Belgica also fell he added four scalpellids, extended the ranges of many
into the capable hands of Hoek (1907a). The material species, and provided detailed descriptions and illus-
included one antarctic species, the rest having been trations (293013, 1939).
collected around the southern tip of South America. The only known representatives of the Rhizocephala
The noted French cirripedologist, A. Gruvel, de- and Ascothoracica in antarctic waters were also taken
voted many studies to the cirripeds brought back by during Discovery cruises, but these were not identified
several expeditions to Antarctica-particularly the until recently. The rhizocephalan, parasitic on lithodid
Ross Sea sector otherwise made famous by the ex- crabs, was described by Boschma (1962). The asco-
plorers Scott, Shackelton, and Amundsen. The first thoracican was discovered by T. Mortensen during his
in his series of papers, published in 1906, noted the work on antarctic ophiuroids in 1936 and was de-
barnacles collected by HMS Discovery under the cap- scribed by Heegaard (1951).
taincy of Robert Falcon Scott. The species Gruvel de- The Discovery reports summarized a series of faunal
scribed in this study and the additional ones added to surveys taken in waters associated with the great whal-
the fauna subsequently in reports of the National Ant- ing industry based in the Falkland Islands during the
arctic Expedition (1907a), L'Expedition Antarctique early part of this century. It is largely as an outgrowth
Allemande du Gauss (1907b), the Deutsche Siidpolar- of the economic interests centering on the whaling in-
Expedition (1910), and finally the results of the ex- dustry that something is known of the cetacean epi-
plorer Jean Charcot's voyage on the Pourquoi-Pas? fauna (R. Clarke, 1966).
(1911a) appreciably advanced our knowledge of the Japanese interests in Antarctica began with explora-
antarctic fauna. tions of the Kainan Maru during the years 1911-1912.

More recently. the Japanese Antarctic Research Expe- highly illuminating as regards the history of he re-
dition aboard the S6ya and Umitaka-Maru brought gion. The oldest are from beds of Cretaceous age, and
back materials studied by Huzio Utinomi (1965). the youngest from Quaternary deposits (the localities
During the International Geophysical Year 1957- and bathymetry of all previously described species are
1958 marly nations undertook scientific surveys in and given in appendix 1).
around Antarctica. Of theee, the Russian research Weisbord (1965, 1967) has already reported on two
vessel Ob' collected numerous benthic samples, and samples collected hy the Eltanir~during the course of
some results on barnacles were published by Zevina sedimentological studies off South America. The pres-
(196.1). No new species were described, and the re- ent study is also concerned with materials taken by the
port probably covers but a fraction of the cirripeds Eltanin from 67 stations. Materials from numerous
collected. Results of Zevina's (196E) study were add- oiher expeditions are also covered in this report (see
ed in proof. appendix 2 ) . Knowledge of the composition and dis-
The search for fossils on the antarctic continent as tribution of antarctic and deep-water barnacles from
well as on islands within the limits of the Antarctic other parts of the world has reached a point where it
Convergence has proven fruitful, and even though only is now reasonable to attempt the unraveling of the
a few barnacle species have been found, the results are region's biogeography.

BIOGEOGRAPHICAL ACCOUNT
Torberi Wolff's biogeographical remarks (1962, p. arc acanthopod representatives of thv pelagic co~npl(ax,
300 j primarily concern abyssal faunas, yet the prin- hut thry apparently d o not cross the Antarctic Conver-
ciples stated are fundamental to biogeographical prob- gence. As a general statement then, antarctic cirripetls
lrms in general: "It is possible that the differencrs (in are microphagus forms, hut specdic diets or prefer-
distribution patterns between groups) can to a certain ences are unknown.
degree be explained by divergent opinions on the spe- Sub~tratum~ecjuireme~its are interesting but com-
cies concept, but on the whole there is no doubt that plex. The acrotlioracicaris require a solid calcareous
the differences of distribution are primarily due to one substrate in nhich to bore and are capable of burrow-
o r more of the following factors: geological age, abil- ing into virtually all known types. Therefore, their dis-
ity to adapt to changes of environment (e.g., at the tribution is obviously limited by the availability of
onset of the Glacial Age), time of migration . . . and such materials and this, in connection with their long
mearis of dispersal once there, present-day rate of geological history, is undoubtedly responsible for the
speciation, dependency on type of substratum, choice present diversity of the order in coral seas. As far as
of food, and type of reproduction." the Antarctic is concerned, present distribution (of the
Jn the present report, an attempt will be made to single known species) is apparently that of Bathylasma
look at the biogeographical problems surrounding the shells and El rirza skeletons.
cirripeds, keeping these points in mind. From the In the Thoracica, the problem of substratum is by
point of view of geological history arid adaptations no mearis as straightforward. As a generalization, the
to changes of environment, it appears that certain sig- Lepadomorpha are better adapted to supporti~igheir
nificant correlations can be deduced. However, there bodies from a relatively small attachment point and
is very little information on feeding habits, substratum therefore do not require broad, solid substrata, as do
requirements, and reproductive cycles of deep-sea cir- the Verrucomorpha and Balanomorpha. Uut diversity
ripeds-too little information, really, to be brought of form in all three suborders found about the world
directly to bear in a significant manner on the bio- is great; this, then, carlnot be stated as a rule. How-
geographical problems at hand. Some of these will be ever, the benthic repre=entatives in the Antarctic ale
touched on in the systematic section of this report, but relatively stereotype-the lepadoinorphans being typcs
before that is taken up, and before beginning the bio- that can attach to minute as well as large objects on
geographical account, a few general comments shoulcl the sea floor (pls. I, 11), and the ve~rnc~oinurphar~s
be made. (pl. XIV) and balanomorphans (pl. XV) requiring
Thoracican and acrothoracican cirripeds are basi- substantial (echinoid .pines and barnacle shells) to
cally setose filter-feeders and obtain food actively by extensive hard bottom.
drawing the cirral net through the water, or, when liv- The verrucomorphans are not and apparently have
ing in currents, passively by allowing the water to pass not been an important antarctic element. The bala-
through the cirral net. The cirri take on three gen- nomorphans, although represented by a single spe-
eral forms: ctenopod (setae arranged in rows along cies today, were quite diverse during the late Cenozoic.
each ramus 1 , lasiopod (setae arranged in brush-like The subsequent reduction is in part related to bathy-
clumps), and acanthopod (setae spine- or claw-like and metry of the habitats available, as will be taken up
arranged in a manner so as to grasp and rasp) (fig. below, but it is also related to the type of substratum
3 ) . All of the benthic species covered in this report are available within the vertical range inhabitable by
either ctenopod or hypolasiopod forms and are there- members of the group. The Pleistocene and Recent
fore capable of capturing particulate matter and micro- history of the continent has been one of considerable
plankton. This is also true of the dominant circum- glacial activity, and what little shelf region there is
antarctic pelagic species Lepas australis. Lepas fasci- must have been subjected to the cyclic raining down
cularis arid Conchotlesma v i ~ g a t u m on
, the other hand, of a variety of sediments. The sole surviving balano-

by widely flaring the apertural end in the manner so

characteristic of the species (pl. XV A-C). Tetra-
chaelasma, found at great depths immediately north
of the Antarctic Convergence, does not take on the
corolliform growth pattern and requires broad sur-
faces on which to grow (pl. XXVI). Its North Atlantic
counterpart, Bathylasma hirsuturn, likewise limited by
the nature of the substratum, has a comparable growth
form (pl. XXIII) .
The basic life cycle of acrothoracican and thoraci-
can cirripeds includes eggs brooded in the mantle
cavity of the adult, six planktonic naupliar stages, and
the cyprid larva. Species in which the naupliar stages
are passed through in the egg brooded in the mantle
cavity are much more limited in their ability to dis-
perse than species with planktonic nauplii. The cyprid
stage selects the settling site and attaches, and under-
goes metamorphosis into the juvenile; however, it is
a weak swimmer and cannot feed. Thus it is not gen-
erally involved in dispersal.
The antarctic acrothoracican Cryptophialus occurs
in relatively shallow water (to 600 meters) and re-
leases eyed, virtually nonswimming cyprid larvae,
comparable to those known for the New Zealand spe-
cies (Batham and Tomlinson, 1965). This, no doubt,
does much in explaining the restricted distributions of
species of this genus, hut not necessarily other gen-
era of the order.
As far as can be determined, all scalpellids in the
Antarctic have separate sexes (females accompanied
by dwarf males), brood their young, and release ad-
vanced nonfeeding cyprids-as would be expected of
deep-sea or high latitude species. The release of a non-
feeding larva of limited dispersal potential and the
past and present oceanic conditions surrounding the
Antarctic probably account for the high degree of en-
demism to be taken up shortly. While there is pres-
+ loose shells ently insufficient data to indicate whether there is sea-
sonality in the release of cyprids, they were frequently
living specimens encountered in the brood chambers of the adults.
The Balanomorpha, on the other hand, are gener-
-
ally cross-fertilizing hermaphrodites and usually pass
Fig. 1. Bathymetric distribution of deep-water through the complete larval sequence including plank-
antarctic balanomorphs.
tonic naupliar stages, exceptions in both cases being
morphan, Bathylasma corolliforme, is, in its mariner known only in one or two species in the entire world
of growth, well adapted to such conditions. It is a (Nilsson-Cantell, 1921 ; Henry and McLaughlin, 1967).
large species and thus less easily smothered, and un- No specimens of Bathylasma or Tetrachaelasma taken
like most, has the ability to settle on relatively small from antarctic waters have been found with developing
objects, such as the isolated plates of its kind resting eggs or brooding larvae. They were all collected clur-
on the bottom. Such an established individual could ing summer months. This is not only remarkable but
not enlarge much basally; hence, large size is achieved it leaves us without knowing which of the two life

BIOGEOGRAPHICAL ACCOUNT 9
cycles might be taking place. The occurrence of free ticum (Withers 1951), and Zeugmntolepas georgiensis
naupliar stages is so general to the Balanomorpha, (Withers, 1947).
were these not deep-water forms, one would ex- Acrothoracica inhabit a variety of calcareous sub-
pect them to produce free nauplii. From certain strata, and fossil genera and species are inferred from
lines of evidence one can infer that they do not the form of their characteristic burrows. Brachyzapfes
conform to the general rule that deep- or cold-water from the Lower Cretaceous of Alexander Island in-
invertebrates tend to brood their young; unfortunately, habited belemnite shells and, according to Taylor
documentation will have to wait until work is com- (1965), differs from B. elliptica Codez of Europe
pleted on the New Zealand Oceanographic Institute's mainly in its relatively large size. There are no Re-
material, taken in the vicinity of the Ross Sea by the cent species attributed to the genus in the world today.
Endeavour. In essence, the situation looks much like The Recent antarctic acrothoracican, Cryptophialus,
the following: It can only be inferred, but it seems clearly had a long history, but it forms a burrow that
that Bathylasma corolliforme releases larvae as nauplii. could not easily be confused with those of other genera
There are four principal reasons for assuming this: of the order and is probably not closely related to
virtually all balanomorphans do so, it is a bathyal Brachyzap fes.
rather than abyssal species, it appears too widely dis- The three lepadomorphans are also known from the
tributed to utilize more limited means of dispersal, and Cretaceous of Europe. The extinct genus Cretiscal-
embryonic development apparently takes place during pellum is known only from the Cretaceous of Britain
winter months. The nauplii, which require plankton and the Antarctic (Alexander Island). The remaining
for food, would not be released until the ensuing two genera were much more widely distributed during
spring, when antarctic production takes a dramatic the Cretaceous than Cretiscalpellurn, but only one,
upswing and can support extended free-living larval Euscalpellum, survives today.
forms. On the other hand, while reproductive and Euscalpellum is represented by some six Recent spe-
growing periods for antarctic scalpellids are un- cies confined to the Indo-West Pacific and West Afro-
doubtedly geared to the south polar productivity cycle, Caribbean, one being within 15 to 100 meters of the
the larvae, as cyprids, do not feed and are not known surface, but most at several hundred meters, and one
to extend far nor remain long among the plankton. down some 3475 meters, well into the abyss. The
genus is well represented in the Eocene of Europe,
PALEOGEOGRAPHY England, USA, Tierra del Fuego, and the Miocene of
"No other large faunal region in the world can match Cuba and Australia (Withers, 1953), but the oldest
the Antarctic in the sharpness of its boundaries" record is for E. antarcticum, in the Upper Cretaceous
(Ekman, 1953). "The Antarctic Convergence - is of of Antarctica.
course the reason why the limit between the pan- Zeugmatolepas is known from Jurassic remains in
Antarctic and antiboreal regions is more obvious . . . Europe, the Lower Cretaceous in Denmark, England,
as compared to the Arctic" (Broch, 1961). It is there- Western Australia, and South America (Peru, U. Se-
fore interesting, although probably not an unsuspected nonian: Withers, 1953; Pilsbry and Olsson, 1951),
fact, that during the Cretaceous Antarctica had a cir- and Zeugmatolepas georgiensis Withers, 1947, from
riped fauna quite similar to that of the rest of the South Georgia, but the genus did not survive into the
world. This fauna followed much the same lines of Tertiary.
extinction as generally marked the close of the Meso- There are only two records of balanomorphans from
zoic. The Cenozoic again was a time of invasion, at the antarctic late Cenozoic and both are considered no
least for shallow-water balanomorphans. But this pe- older than Miocene. Balanus (Austrobalanus), a rela-
riod too was followed by extinction, apparently before tively generalized or primitive group, is at present con-
or during the Pleistocene. Today the shallow-water fined to Australia, New Zealand, and South America.
elements, those within 100 meters of the sea surface, A Recent species in New Zealand, Balanus ( A . ) vesti-
constitute but 5% of the cirriped fauna. This is essen- tus, has inhabited the region at least since the Lower
tially the reverse of other regions of the world. Oligocene (Withers, 1953), and a South American
The Cretaceous fauna is known by four species: the species, B. ( A . ) flosculus Darwin, considered most
acrothoracican Brachyzapfes elliptica gigantea (Tay- closely related to B. ( A . ) imperator Darwin from Aus-
lor, 1965), the lepadomorphans Cretiscalpellum aptien- tralia, was found in the ?Miocene of Kerguelen Islands
sis antarcticum (Taylor, 1965), Euscalpellum antarc- (Fletcher, 1938). It is not known whether Austro-

10 ANTARCTIC CIRKIPEDIA
balanus ha? penetrated any farther than this into the teresting, tells us nothing of the biogeography of the
Antalctic. region in a historical sense (chart 4 ) . Likewise, the
The other record is given by Hennig (1911), who whale barnacles (Conchoderma auritum, Coronula and
reported Balar~us sp. occurring on scallops in the Xcnohalanus; appendix 1, pls. 111, XLVIII ) have the
Pleistocene of Cockburn Island, off Graham Land. same distribution as their migratory hosts and are
However, only l~hotc~graphswere given; subgeneric therefore not strictly of the antarctic fauna.
determinations were not made. One of his specimens The benthic cirripeds car1 be divided into five bio-
(pl. XLVI 6 ) loohs superficially like Bathyla,rna, but geographical groups. Some reprcscntatives of each are
it can I-)e ohservcd to have well-developed radii and is fairly well known, hut others, especially among the
therefore piobahly more assignable to Balanus than to scalpellid Lepadomorpha, can be placed only tenta-
Batkylnsrna. The olher specimen (pl. XLVIl C ) , tively. The genus Arcoscalpel(um is particularly diffi-
although clearly a balanornorphan, is iemarkahle in cult; of the 100 or so specirs only a Few are known
appearing LO have more than six platcs, ill which case well enough to he l~iogeographicall useful. The
it could not be a Balanrrs as presently dcfined. Two of groups . ~ n dincluded taxa are as follows:
the calcareous hases on the scallop shells (pl. XT,VII
A ) Spccics encountered in the Antarctic, but widely
A, 13) appear to be lhose of Balanus, but the third
distributed elsewhere :
(111.XLVI C ) does not. This material is much in need
Briartrsaccus callosus Uosc.hrna, 1930
of stud), but under arly circumstance it is evident that
AT-coscalpellurnvitr cum (Hoek, 1883)
there were at least t ~ ifo not more ~ p x i e sp l e ~ e n t .
Arcosca/pellurn lo?n ~ turn
o ( Hoek, 1907h)
The age is in questionpalthough pssi1,ly Pliocene,
Vcrruca gibbosa Hoek, l:K3
u ~ ~ p Miocerre
er oi interglacial Pleistocene cannot be ex-
cluded (Fleming, 1057, according to Speden, 1002, p. U ) Bipolar forms :
7 0 0 ) . In any event these have no apparent affinities Ascothorax spp.
nilh Uathylasrr~a,and there ale no surviving balanids Bathylasrna spp.
in Antarctica today. C) Circumpolar antarctic ant1 suhantarctic forms:
l'leistocerle deposits are exceedingly r ara, hut thr Cryptophialus spp. (austral group)
occurrence of thc living species, Rathylasma iorolli- Lcpas australis Darwin, 1Z51
lorme ( Hoek ) ( =IIr~xrlasma arztarclicurn Horra- Arcoscalpellr~rnweltnrri (Gruvel, 1907b)
daile) , in the S(.allop Hill Formation of the McMur do Scalpellz~rnannlroflcrti Gruvel, 1907b
Sound resion is certain (Spedcn, 1962). Closc- rela- Uathylasrna corollsforrnr (Hoek, 1383)
tives, R. aucklar~tliczrm(Hector) from the Miocene and Balnnus (Austrobalanuc) spp.
perhaps ?Hc,xelasrna sp. (Withers, 1055) from the Ralanus (IMegabalanus) spp. (austral group)
Lower Oligocene. are known from New Lealand.
D) Forms with signific,arrt antarctic or suhantarctic
Uathylasrrtn rorolli/orn~e is the sole balanomorl~han
clistrihutions. hut not circumpolar:
surviving south of the Antarctic Convergence. Tts bio-
Cryptophial~rsI ~ I ~ L I Z I I Ssp.
O I L1101.
~
geographical afinities will be taken up shortly.
Scalpellurn grbbc~rurn Aurivillius. 1391.
Clcarly, great changes occurred in the antarctic cir-
Litoscalpellun~.tl~scowryi( Gruvel, 1906)
ripctl fauna during the Cretaceous ancl Tertiar, anti
Arcosca'pclEzim cornpnc turn (Borradaile, 1916)
no genera or spec.irs known by fossils have burvi\c.d.
Ar coscalpellurr~brevccnrinatum iHoek, lU83 )
Yet the liair~gfauna must he rclatively long stantling
Tctrachar~asrrtacotrthu~ardigen. ct sp. nov.
since 27 (75' : ) of the 36 living species are cndemic
Rulanus (Balanus Inevis Rruguikre, 1'789
to the ar~t,l~ctic region. Endemism will he returnled to
Balanus (Au,~robalanus) flosculus Darwin,
irr the di~cu5sior1 of the modern fauna.
1854
( ‘ O N ' r L hI I ' O K A R Y BIOGEO(~1iAI'kiY El Endemic forms from south of the Antarctic
The Recent antarctic and subantarctic cirriped fauna Convergence :
,,an he divided into three ecological groups : benthic Ascothora.t- hulhosus Heegaard, 1951
specics, pelagic species, and those occurring on whales. Cryptophialus tornlinsoni sp. nov.
Only the fi 1st will he considered here to an apprrciahle Arcoscalpellum acicularum sp. nov.
extent. 'l'his is because the only truly antarctic pelagic A. angulare (Nilsson-Cantell, 1930)
species, l,epns ausllalis D a r ~ i n while
, ecologically in- A . unlarcticurn (Hoek. 1E83)

A. berrulti (Gruvel, 1907b) Antarctic faunal communications and ascothoracicans

A. bouveti (Nilsson-Cantell, 1939) simply may not have been available to draw upon in
A. bouvieri (Gruvel, 1906) either direction.
A. compactum (Borradaile, 1916) ACROTHORACICA: The discovery of a living acrotho-
A. gaussi (Gruvel, 1907b) racican of the genus Cryptophialus in the Antarctic is
A. latusculum sp. nov. extremely interesting. As mentioned in the section on
A. liberum (Nilsson-Cantell, 1930) paleogeography, the Cretaceous form, Brachyzapfes,
A . magnaecarinae (Nilsson-Cantell, 1930) was recognized in the Antarctic by its characteristic
A . multicostatum sp. nov. burrow (Taylor, 1965). The burrow of Cryptophialus
A. recurvirostrum (Hoek, 1883) is unique and could not be confused with that of
A . weltneri (Gruvel, 1907b) Brachyzapfes, because the apertural end of the animal
Neoscalpellum schizoplacinum sp. nov. is constricted into a relatively long cylinder, and con-
Gymnoscalpellum tarasoui gen. et sp. nov. sequently the burrow is likewise formed (cf. fig. 5K,
Litoscalpellum fissicarinatum gen. et sp. nov.
L) .
L. simplex sp. nov. Until recently Cryptophialus s.1. was known only
L. walleni sp. nov. from the southern hemisphere, but now species are
L. discoveryi (Gruvel, 1906) known to occur north of the equator in both the At-
L. convexum (Nilsson-Cantell, 1921) lantic and the Pacific (Tomlinson, pers. comm.) . The
L. aurorae (Bage, 1938) genus can be divided into two groups on morphologi-
Scalpellum vanhoffeni Gruvel, 1907b cal grounds, and the more generalized of the two, rep-
Australscalpellum schizomatoplacinum gen. et resented by a species in New Zealand and in South
sp. nov. Africa, is here distinguished as the "austral group of
Bathylasma corolliforme (Koek, lSX3) Crypt~phialus."~The new species from the Antarctic,
RHIZOCEPHALA: Briarosaccus callosus Boschma is the having a known range from the Ross Sea to the Falk-
only member of the order known from south of the land Islands, belongs to this section, and knowledge of
Antarctic Convergence. It is widely distributed else- its existence and distribution significantly decreases
where, being found north of the equator in the Atlantic the gap between the previously known species (chart
Ocean and in the Bering Sea. Rhizocephalans are para- 3 ) . As far as is known, members of the genus pass
sitic primarily on decapod crustaceans and are more their pelagic naupliar stages in the egg and emerge as
diverse in cold than in tropical waters. Broch (1961) weak or nonswimming cyprids (Batham and Tomlin-
and Yaldwyn (1965) have pointed out that the Ant- son, 1965). It might be assumed then that dispersal
arctic has a very depauperate decapod fauna and the has been accomplished through the adults being trans-
virtual absence of rhizocephalans is surely concomitant ported in the shells of other organisms, such as gastro-
with this fact (Hedgpeth, pers. comm.) . pod molluscs clinging to seaweeds. The species are
ASCOTHORACICA: These cirripeds are parasitic on coe- very distinct, however, so either dispersal by such
lenterates and echinoderms, and the latter are par- means must be relatively infrequent or conditions have
changed significantly and it no longer occurs.
ticularly diverse in the Antarctic. Yet, but one asco-
thoracican, Ascothorax bulbosus Heegaard, is known THORACICA: This order constitutes the most diverse
from the region. The lack of hosts cannot be respon- group of cirripeds in the Antarctic, as one would ex-
sible for the situation as it may be for the rhizocepha- pect from knowledge of other regions of the world.
lans. The problem is manifold, partly because the Endemism is very high, 75%, but this also would be
group is poorly known. However, it is curious that expected from what is known of other antarctic groups
while there is a great diversity of ascothoracican genera (fishes, echinoderms, mollusca, etc. ; Ekman, 1953) .
and species in the Arctic, North Pacific, Indo-West The remarkable aspect of the fauna is the extremely
Pacific and Mediterranean, they are extremely rare in high ratio of lepadomorphans to balanomorphans; ap-
the Atlantic (Wagin, 1964). This curiosity is height- proximately 32 : 1. By contrast, an isolated oceanic
ened when it is noted that Ascothorax is bipolar with archipelago like Hawaii has a ratio on the order of
no species known in between. The lack of forms in the
Atlantic may be part of the explanation since histori- After this paper went to press, this group was formally
cally there have been strong northeastern Atlantic- separated as a new genus, Aust~nlophialus (Tomlinson, 1969).

TABLE 1. Benthic Cirriped Genera, and Numbers of Species, Known from South of the Antarctic Convergence. Arranged
Systematically
--
Number of
Previously New Species Recent and
Known and Exten- Endemic ( )
Fossil Recent sions ( into Taken by Antarctic
Species Species Antarctic Eltanin Species
Ascothoracica 0 1 0 0 1 (1)
Rhizocephala 0 1 0 0 1 (0)
Acrothoracica t1 0 1 1 1 (1)
Thoracica
Lepadomorpha
Scalpellidae
Euscalpellum
tZeugmatolepas
tCretiscalpellum
Arcoscalpellum
Neoscalpellum
Gymnoscalpellurn
Litoscalpellum
Scalpellum
"Australscalpellum
Verrucomorpha
Verruca
Ralanomorpha
Bathylasmatidae
Bathylasma
Balanidae
Balanus (Austrobalanus)
tBalanus spp.
Totals 8 25 lO(4) 21 36 (27)
t Extinct genera and species.

* Endemic genus.
1.33 : 1 (Newman, in press) and continental margins tolepas and Cretiscalpellum) are extinct, and the third
like west Mexico and Japan have ratios of 0.2:l and (Euscalpellum) has representatives living at low lati-
0.9 :1, respectively (Ross, 1962; Utinomi, 1958) . The tudes. There are, then, some 18 Recent lepadomorphan
reason for this marked difference is that the Balano- species south of the Antarctic Convergence, and prior
morpha holds its greatest diversity in the littoral re- to this report these were mostly included in Scalpellurn
gion, within the first 100 meters or so. Very few are s.l., a genus consisting of more than 200 species occur-
found at 1000 meters and only two of the approxi- ring throughout the deep waters of the world. This
mately 250 species in the world reach depths of 2000 is a rather large and unmanageable assemblage. How-
meters (e.g., Bathylasma hirsutum (Hoek) in the ever, it is composed of a number of fairly distinct
North Atlantic and Tetrachaelasma southwardi gen. species-groups. Various authors in the past, particu-
et sp. nov. in the subantarctic). In the lepadomorph- larly Hoek (1883, 1907b), and Pilsbry (1907b, 1908),
ans much the reverse is true since most species occur have attempted to organize and define them. Of the
at depths in excess of 100 meters or so and many at numerous groups proposed, Mesoscalpellum and Arco-
depths exceeding 4000 meters. Thus, the lepadomorph- scalpellum have been recognized in the literature as
ans would not have been affected as much by glaci- genera or subgenera by some but not all workers. We
ation, which must account for the difference. consider these to be genera and propose that the re-
The three lepadomorphan genera known from the mainder of Scalpellurn, with the exception of Scalpel-
Antarctic by their fossil remains were discussed under lum s . ~ . ,be further divided into six genera. These,
the section on paleogeography. Two of these (Zeugma- plus two new genera erected herein, bring the group

BIOGEOGRAPIIICAL ACCOUNT 13
TABLE 2. Bathymrtric Distribution of Bcnthic Cirripeds Known from South of the Antarctic Convergence.
Nunlhcrs of species cncuuntcred within each depth interval for each genus arc given in two colurrrns under such genus. Thc
left c:olumn of each pair is for he riumher of tinlrs species wcre enoounteretl within each drpth interval prior to the present
report; the right columrr, for the Eltanin and othcr expeditions covered in this report. Totals represent number of species
encounters, not total numbcr of specie?.
Depth
(meters) Totals
4 0
21 21
12 4
1 5
0 0
0 1
1 4
1 1
0 0
0 4
2 2
Totals 18 19 11 4 0 1 0 5 3 2 0 3 1 1 9 4 0 3 42 42
up to 10 genera. So divided, a formerly very unman- (chart 5). Nothing can be said biogeographically at
ageable group becomes more manageable. Undoubtedly this point, but the bathymetry of the genus in the Ant-
there will be further revision, particularly in the larg- arctic (table 2) is most interesting and will be re-
est genus Arcoscalpellum. turned to later.
The scalpellid genera falling under this revision can The next in importance is Litoscalpellum, consisting
be divided into three biogeographical groups: (1) of nine species, at least six of which are endemic to
those with no antarctic representatives; ( 2 ) those with the Antarctic. The remaining species have an Indo-
antarctic representatives; and ( 3 ) those endemic to West Pacific-austral distribution (chart 7). The re-
the Antarctic or subantarctic. maining genera, Scalpellum (chart g ) , Neoscalpellum,
Three genera fall into the first category: Annan- and Gymnoscalpellum (chart 6 ) , each have one ant-
daleum is strictly Indo-West Pacific in distribution arctic representative and in each case it is endemic.
(chart 8 ) . Mesoscalpellum, on the other hand, has a Scalpellum vanhofeni is known from two localities,
curious distribution, depicted in chart 7. The lines en- one on either side of the Antarctic Continent, in rela-
closing the localities of the included species are con- tively shallow water. Therefore, although apparently
nected across Central America, rather than across the rare, it may be circumpolar. Neoscalpellum schizo-
Arctic Ocean, because of the record for the widely dis- placinum, on the other hand, is known from one local-
tributed Atlantic species, M. imperfectum, from the ity at little-explored depths in excess of 4800 meters
Galapagos Islands. Monotypic Abathescalpellurn from (appendix 4 ) . The remaining species in the genus are
Korea (chart 6) inhabits remarkably shallow water also from deep waters of the Indo-West Pacific, East
for a scalpellid with reduced armament and this may Pacific and North Atlantic. The remaining genus in
account for its lack of diversification. The same situ- this group, Gymnoscalpellum, is known from three
ation is encountered in the antarctic endemic, Aus- widely separated localities in the world. Gymnoscal-
tralscalpellum. pellum tarasovi, endemic to the Antarctic, has a bathy-
Of the five widely distributed genera having repre- metric distribution that is especially interesting because
sentatives in the Antarctic, Arcoscalpellum is the best it tends to fill the hiatus left by Arcoscalpellum be-
represented. Of the 21 species, 14 are endemic, being tween 1200 and 2500 meters. Whether this is due to
known only from south of the Antarctic Convergence sampling deficiencies cannot be readily determined, but

14. ANTARCTIC CIRRIPEDIA
as indicated in appendix 5, 40 samples totaling 27.11 direct affinity with the aforementioned genera, it is
hr of bottom time were taken between 1000 and 2500 probably with Aaptolasma.
meters by the Eltanin. Of the 19 samples totaling Members of these genera are neither strictly balanids
12.62 hr between 1000 and 1499 meters, 6 yielded cir- nor chthamalids, and are grouped together in the new
ripeds, while 21 samples for 14.49 hr between 1500 family, Bathylasmatidae. The generalized members of
and 2500 meters ~ i e l d e dnone. Nor have previous ex- the family, Bathylasma and Tetrachaelasma, occur in
peditions obtained Arcoscalpellum between 1000 and the North Atlantic and antarctic region below depths
2500 meters (table 2 ) . In fact, no barnacles of any inhabited by other Balanomorpha. Aaptolasma and
type have been taken between 1500 and 1999 meters Hexelasma, while living below the optimum balanid
south of the Antarctic Convergence, so the hiatus in zone, occur in the Indo-West Pacific where balano-
the data at least is real. What significance it has with morphan diversity is extremely great. The apparently
regard to the antarctic fauna remains to be explained. different distributions, North Atlantic-Antarctica and
The only genus endemic to the region south of the western Atlantic-western Pacific, indicate that this is
Antarctic Convergence is the monotypic genus Aus- an old group as far as balanomorphans are concerned,
tralscalpellum. Brochia (chart 9 ) , also monotypic, is and the fossil record takes them back to the Lower
a subantarctic endemic. Interestingly, like monotypic Oligocene in New Zealand. The suggestion is. then,
Abathescalpellum from the northwest Pacific, both are that Bathylasma and Tetrachaelasma are austral in
from relatively shallow water. origin and can be considered deep-water refugial
forms.
VERRUCOMORPHA: The suborder appeared and flour-
Three faunal similarities between Antarctica and the
ished in shallow seas during the Upper Cretaceous,
North Atlantic or Europe have been noted: the Cre-
but most species are restricted to fairly deep water
taceous acrothoracican, the Cretaceous scalpellids, and
today. Verruca gibbosa, the only species known from
the Lower Oligocene bathylasmids. Among the balan-
the Antarctic, is so widely distributed elsewhere in the
ids, we are confronted with another example, Balanus
world as to be considered cosmopolitan (chart 9 ) .
(Balunus) luevis Darwin. The Recent population ex-
Thus, the antarctic v e r r ~ c o m o r ~ h afauna
n is remark-
tends around South America, from 2 0 " s to 20°S, the
ably depauperate, a distinction it shares with the East
southern extent terminating just north of the Antarctic
Pacific in the northern hemisphere where no species
Convergence (chart 11). The fossil record indicates
are found.
this species was in Europe and North Africa during
BALANOMORPHA: Members of the suborder were dis- the Lower Miocene (Davadie, 1963) and has been in
cussed briefly in the section on paleogeography, where South America since the Pleistocene (Darwin, 1854;
it was pointed out that while several species were Richards and Craig, 1963).
known to have occurred south of the Antarctic Con- In conclusion, can any generalizations be made from
vergence in the Neogene, only one is known today. the fragmentary data available? We have already dis-
This species, Bathylasma corolliforme, found living cussed the marked faunal similarity between Antarctica
uithin the Antarctic Convergence between 100 and and northern Europe that existed during the Creta-
1500 meters, is circumpolar in distribution (chart 1 0 ) . ceous, there having been species as well as genera in
Only one other living species in the genus, B. hirsutum, common. This is not entirely unexpected, since faunas
is recognized, and it occurs in the North Atlantic to were generally similar about the world at that time.
depths of 2000 meters. A closely related species, Further, the occurrence of a Mediterranean Miocene
Tetrachaelasma southwardi, also occurs to depths of species in the Recent subantarctic of South America
2000 meters, but in the Southeast Pacific Basin to indicates a Tertiary exchange between the two regions.
around the southern end of South America. Bathy- We have also noted extensions of contemporary faunas
lasma is related to Hexelasma of the Indo-West Pac- of the northern hemisphere into antarctic waters. Lito-
cific, and the two occur together in deposits of Miocene scalpellum and the family Bathylasmatidae, while pri-
age in New Zealand. Another genus, Aaptolusma, more marily Indo-Pacific today, apparently made their u7ay
closely related to Hexelasma than to Bathylasma, also in one direction or the other, via the New Zealand re-
occurs in the Indo-West Pacific, but, curiously, is rep- gion. The direction z re sum ably would have been away
resented as well by a species from the western Atlantic from the apparent center of diversity, the Indo-Pacific,
(chart 101. A Miocene fossil from Burma, Tessare- at least for Litoscalpellum. The Bathylasmatidae, in
lasma, is somewhat enigmatic, but if it actually has a having two widely divergent genera common to both

the Atlantic and the Indo-Pacific, perhaps had a Cre- Probably the most remarkable aspect of the antarctic
taceous-Paleocene u orld-u ide distribution that u as sub- cirriped fauna is the very high proportion of lepado-
sequently restricted, primarily to the Indo-Pacific. morphans to balanomorphans, and the concomitant
Neoscalpellurn, containing some of the deepest known lack of diversity between 100 meters and the shore.
scalpellids, is primarily northern Atlantic and Pacific, Glaciation and cyclic freezing to appreciable depths
but nith representatives along the nest coasts of North probably accounts for this, through extinction of lit-
and South America which suggest extensions into Ant- toral Tertiary elements and prevention of recoloniza-
arctica via this route. Then too, the bipolar distribution. Virtually no attempt has been made to compare
tion of Ascothornx further emphasizes exchange be- these findings and inferences with those proposed for
tween the t~ o hemispheres. Clearly, while the contem- other groups of organisms about the antarctic region.
porary antarctic cirriped fauna is highly provincial, For this we apologize, but with the belief that such
there have been avenues of communication for both should eventually be handled by a reviewer having a
shallow- and deep-nater forms during several Tertiary synthetic mind uncluttered by details and probably not
periods. burdened by the same personal biases.

MATERIALS AND METHODS

DISSECTIONS ing it to the sheath of the wall, allowed the entire mass
The Acrothoracica are burrowing forms, able to to be forced out through the base. Once removed, the
penetrate virtually any calcareous substratum, par- body was dissected free of the opercular parts, under
ticularly mollusk shells and skeletons of living and water. When the mantle cavity contained ovigerous
dead corals. ? l e y are less often found in echinoid lamellae, developing embryos, or larvae, they were
and balanoid shells, but are frequently encountered saved for study.
in limestone. Usually the only visible sign is the The opercular parts, as well as the wall plates, were
small slit-like opening to the burrow, generally a cleaned and disarticulared by soaking in sodiunl hypo-
millimeter or so in length and shaped like a straight chlorite (commercial bleach such as Clorox) .
or slightly curved apostrophe (fig. 6; pl. IV A ) . This Once the body was removed, it was studied under
small aperture opens into a commodious oval chamber water, note being taken of the attachment and arrange-
excavated and inhabited by a female, usually accom- ment of cirri, the presence or absence of filamentary
panied by one or more dwarf males. The chamber may appendages (Lepadomorpha) , or branchiae (Balano-
extend straight or somewhat obliquely down from the morpha) , etc. After these observations, specimens were
surface and can be studied by successively grinding the dissected, the cirri and mouth parts being mounted on
substratum, gradually exposing the outline (fig. 5L). one or two slides. For mounting extremely large speci-
Females were removed by chipping the substratum away, mens, glass slides measuring 50 X 75 mm were neces-
but usually not without some obvious damage because sary. Glycerine jelly, or permanent synthetic resins
they are cemented at a point to the wall of the burrow. and media such as Turtox CMC and Euparal were
Alternatively, the calcareous material was dissolved used. However, stock CMC-S stains much too intensely,
away with acid. Specimens were then studied exter- and is best diluted by 10 to 20 parts of CMC. CMC
nally and dissected in much the same manner as Lepa- uTas found to be quite acidic and could not be used
domorpha (below). Other specimens were treated with when calcareous parts were to be preserved, especially
dilute sodium hydroxide, in the manner described by plates of dwarf males.
Tomlinson (1960), until most of the soft tissues were The body of the barnacle in a dish of water usually
corroded away. lies on one side. It was convenient to place the animal
Preparation of adult thoracicans was relatively sim- on its left side and to dissect away the right cirri one
ple (figs. 2, 3 ) . Lepadomorphans were removed from by one, starting with the first. It was found preferable
the mantle by plucking a hole in one side, withdrawing to leave the caudal appendages attached to the base of
the body, and then freeing it completely by teasing the pedicel of each sixth cirrus, because their height,
away the attachments of the adductor muscle. The relative to that of the pedicel of cirrus VI, is of taxo-
body of armored lepadomorphans was usually removed nomic value. As each cirrus of the right side was re-
through the aperture by grasping the attachment area moved, it was placed in mounting medium on the slide
at the adductor muscle lying between the scuta and outside surface up, in order from left to right. The
gently withdrawing the body. Some teasing of the cirri of the left side were placed in a like manner so
membranes between the scuta was necessary in order that their inner surfaces were up. Outer surfaces of
to enlarge the aperture. right and inner surfaces of left appendages are gen-
In the Balanomorpha the body is too large, relative erally illustrated in the text-figures.
to the size of the aperture, to be removed in this man- The appendageless thorax and the prosoma support-
ner. But it can be removed, along with the scuta and ing the buccal mass now can be separated. The former
terga, through the base, leaving the wall intact. The can be discarded, provided it does not support filamen-
ease with which these parts were removed depended tary appendages. In dissecting out the mouth parts,
on the structure and condition of the barnacle. Usu- the first maxillae and mandibles were removed before
ally, inward pressure applied to the operculum, and attempting to remove the second maxillae, because the
teasing or cutting of the arthrodial membrane connect- second maxillae are rather delicate and fused together

C C
-=I
X-sect of balon~dwall
01
Lepos Sm111um Sca/pellum s I.
V
LEPADOMORPHA
VERRUCOMORPHA BALANOMORPHA
Bolanus
af B*
SP
scuturn tergum
Bolan~dopercular plates
Fig. 2. External morphology of Thoracica: a, ala; af, articular furrow; ad, adductor muscle pit;
ant, articular margin ; ar, adductor muscle ridge; arg, ar~icularridge ; ax, apex; bnz, basal margin ;
r, carina; ra, capitulum; cl, carinolatus; cm, carinal margin; fs, fixed scutum; It, fixed tergum;
if, intcrlaminate figure; il, inner lamina; iml, inframedian latus; 1, latus or lateral plate; lc,
longitudinal canal; Id, l a t ~ r a ldepressor muscle pit or crests; Is, longitudinal septum; nzs, movable
scutum; mt, movable tergurn; oc, occludent margin; 01, outer lamina; p, parics; pe, pcduncle; ps,
peduricular scales; r, rostrum; ra, radius; r(1, rostra1 drpressor muscle pit or crests; rl, rostrolatus;
s, scutum; sc, subcarina; sp, spur; t , tcrgum; ul, upper latus.
basally. The numerous muscles and adhering shreds of cement was melted on the slide, over a hot plate. Pres-
cuticle were removed from the labrum, and one or both sure was applied to the specimen before cooling, so
of the palps were usually removed and mounted sepa- that it came into even contact with the slide surface.
rately, thus completing the preparation. The melted cement is hot (145OC o r more) so that a
piece of sponge proved useful in applying the needed
T H I N SECTIONS O F THE SHELL pressure. A piece of thin aluminum-coated wrapping
lnterlaminate figures, tliscovered by de Alessar~dri paper placed betueen the sponge and the specimen
(1895), are found in balanomorphans having a wall prevented sticking.
permeated hy longitudinal tubrs. Cornwall (1962 and Rough cutting required abrasives of 100- to 200-grit;
earlier) and Ilavddie (1963) have used ir~terlamirlate finishing cuts and polishing, 400- to 600-grit. The slide
figures in the shell determination of balanomorphan holder consisted of two accurately ground metal blocks,
species from fragmentary remains. Figures are oh- held in alignment by rods running through them, and
served in the transverse section of the parietes and grooved to hold a standard slide by tightening a thumb
represent crystalline organization about an organic. screw, which draws the blocks together. The outer
matrix developetl during shell formation (Newm,ln. surface of the slide is held slightly below the surface
Zullo, arid Wainwright, 1967). of the blocks so that a specimen on the slide will be
A variety of techniclues and equipment are available ground to a thickness equaling the difference in sur-
for cutting thin sections. The simplest is to mourlt faces when the surface of the block comes into com-
a piece of shell to be sectioned on a glass microscope plete contact with the grinding plate at the end of the
slide and grind the exposetl surface to a thickness of cut. A properly ground specimen was dead flat and
about 1 mm. ?'he specimen is then removed, turned of uniform thickness, provided flat slides were selected.
over, and reaffixed with the freshly ground flat side Slide holders, such as those produced hy Buehler
toward the slide. The newly exposed surface is then and Co., Ltd., are designed to hold petrographic slides,
g~ourrdflat, and the section ground to 20-30 p m . which, after having been ground to a standard thick-
The barnacle shell to be sectioned was affixed to ness in another specially designed holder, will clear
the slide with petrographic. thermoplastic cement. The the grinding surface by approximately 50 p m , thereby

MATERIALS AND METHODS
Fig. 3. Internal anatomy and larval morphology of Thoracica. A-C, cirral types: A, ctenopod; B, lasiopod; C, aranthopod.
D-H, trophi: D-E, unnotchrd and notched labra; F, mandible; G, first maxilla; H, second maxilla (arrow indicates papilla
supporting maxillary gland opening). I, antenniform cirral ramus (cf. A-C). J-L, caudal appendages: J, unsegmented; K,
uniarticulate; L, multiarticulate. M-Q, types of cirral setae: M, simple; N, plumose; 0, pinnate; P, bipectinate (card) ; Q,
card with guards. R, schematic generalized adult: 1, first antennae; 3, mandible; 4, first maxilla; 5, second maxilla; 6, lab-
rum; 7, mandibular palp (attached to mandible rather than margins of Iabrum in Acrothoracica) ; 8-13, cirri I through VI,
respectively; 14, caudal appendage; 15, ~robosiformpenis (arrow indicating basidorsal point, found only in balanids) ; 16,
testes; 17, female genital aperture; 18, ovaries; 19-21, filamentary appendages, at base of first cirrus, on prosoma, on dorsal
margin of thorax; 22, male cyprid in mantle margin pocket of scutum; 23, scutoral adductor muscle; 24, mouth; 25, stomach;
26, digestive gland; 27, anus; 35, developing embryos in mantle cavity. S, second of six naupliar stages: 1, 3 and 6 as in R ;
2, second antenna; 28, caudal spine (arrow, caudal process) ; 29, frontolateral horn; 30, frontal filament; 31, nauplius eve.
T, bivalved cyprid larva: 31, as in S ; 32, compound eye; 33, 6 pairs thoracic natatory appendages (cirri in adult) ; 34. caudal
furca (caudal appendages in adult).
yielding sections too thick for our purposes. Ordinary the slide and backing face of the holder. When the last
microscope slides are considerably thinner than petro- polishing cut was to be made, it was found best not to
graphic slides, so that when used without backing rely on estimates made with the micrometer of the
shims, the first surface grinding results in a section number of shims necessary to bring the section to final
more than 100 p m thick. Such a section was strong thinness, but rather to use a 0.001-inch shim (approxi-
enough to be turned over and pressed flat against the mately 26 p m ) as a feeler gauge inserted between the
slide, in order to cut the second surface. The thickness surface of the slide and a straight edge (the long edge
of the section was estimated by subtracting the thick- of another slide), held across the runners. Sections
ness of the slide adjacent to the specimen from thick- were occasionally lost without taking this precaution.
ness of the slide and the specimen, as measured with a While we have found it convenient to use ordinary
micrometer before beginning the second cut. The microscopic, rather than petrographic slides, slides of
amount of material to be removed was then estimated a given lot, although parallel-sided, usually vary in
and an appropriate number of shims placed between thickness from one end to another and from side to

side, some excessively. A box was checked over quickly wear in feeding or damage, or in need of regeneration
by measuring the thickness of individual slides at each can be renewed by moulting without exposing the ani-
end, and at each side near the middle. Those varying mal.
only a few micra, especially from side to side, were In many respects, thoracic cirripeds are much like
used for grinding thin sections. The micrometer, cali- bivalved molluscs because the soft tissues and organs
brated in lOOths of a millimeter (10 p m ) and read- reside within a cavity formed by a permanent shell.
able to within a few micra by interpolation, was satis- They differ in having the filter-feeding mechanism
factory for this purpose, and for estimating the thick- formed from modified limbs rather than ciliated gills.
ness of the section. While the external form has been molded by natural
A sheet of plate glass, about 9 x 12 inches or larger, selection to a wide variety of habitats, the basic feed-
mounted on a pair of wooden supports arranged to ing mechanism has generally undergone very little
bridge a sink, formed a convenient grinding surface. change. The very obvious differences in the external
The surface should be wet, and when using sandpaper armament, in contrast to the relatively subtle differ-
running water is necessary. A rubber tube from the ences in trophic structures, account for the relatiire
cold water faucet provided a slow stream of water, the importance of the former in classification as well as in
runoff spilling into the sink. distinguishing between populations.
Ordinary waterproof sandpaper, especially in coarser The acrothoracicans, on the other hand, in having
grits, was satisfactory. The relatively new abrasive- gained protection by burrowing into the substratum,
coated Mylar films sold under the trade name Flex-i- have undergone relatively little evolution in gross form.
grit are more durable, especially in the finer grits, but The principal external characteristics of taxonomic
generally are not locally available. value are found associated with the opercular bars that
Preparation of blanks of the material to be sectioned guard the aperture of the burrow when the feeding
was accomplished by power grinding. Small, relatively structures are withdrawn, and variations in the funda-
inexpensive, diamond-impregnated discs or wheels mental theme are relatively limited as compared to the
available through dental supply houses, driven and thoracicans. Being setose filter-feeders, the same rela-
manipulated by hand-held motors such as those avail- tive limitations as seen in the Thoracica prevail, and
able under the trade name Mototool, are highly recom- the form of the appendages is, if anything, more stereo-
mended. With them, relatively thin, flat-sided portions typed, in general differing more in number than in
of a specimen can be removed, often leaving the bulk kind. Therefore, the basic classification from its incep-
of the specimen intact. Although manufacturers recom- tion has depended both on internal and on external
mend that their wheels be run wet, satisfactory cut- characteristics.
ting or microslabbing was accomplished with the The degree to which various taxonomic characteris-
wheels dry. tics are used in this study depends to a large extent
on the foundations built by previous workers, and
CHARACTERISTICS OF TAXONOMIC therefore varies from group to group. For example,
VALUE UTILIZED IN CLASSIFYING while the day when the classification of Balanomorpha
C I R R I P E D S
can depend largely on gross structure has passed, de-
Cirripeds are crustaceans and crustaceans are funda- termination of supraspecific categories among the
mentally active, free-living arthropods. As such they Lepadomorpha by external morphology is still to be
generally moult periodically throughout life, and dur-
worked out, especially in the Scalpellidae. In the
ing the moulting process they are particularly vulner-
Balanomorpha we have had to rely heavily on details
able to depredation and mechanical injury. Motile
of the appendages and the fine structure of the shell,
forms can seek shelter and are somewhat reclusive
while in the revision of Scalpellurn s.l., macroscopic,
during this critical period, but sedentary or attached
forms become very much disadvantaged, be it only externally visible differences in form have been, and
temporarily. The plankton-feeding cirripeds have should continue to be, highly instructive as well.
overcome the problem of being attached and moulting It has already been noted that Scalpellurn s.1. is a
in two principal ways; either by burrowing into the large and unmanageable group of more than 200 spe-
substratum (Acrothoracica) or by developing an ex- cies and that in the present work it was found advis-
ternal armament that is retained throughout life able to divide it into 10 genera. It would be well at
(Thoracica) . So protected, the appendages subject to this point to state the philosophy behind this approach,

MATERIALS AND METHODS 21
and the method by which it was carried out. A good not only useful in classification but also in unraveling
example is probably found in the arcoscalpellids, an the biogeography of the family.
assemblage of similar forms having a cosmopolitan External characteristics are generally less useful in
distribution. A preliminary survey of the literature classifying Balanomorpha mainly because their organ-
revealed that while there were few gross differences in ization is the result of reduction of the lepadomorphan
the appendages between species, there were differences plan, and there are fewer readily visible characters to
in the form of the ~ l a t e scovering the capitulum. A go by. The manner in which the wall plates are articu-
catalogue of illustrations was made utilizing the Xerox lated, and the gross
- internal structure of the plates and
copying process. The illustrations were then sorted opercular parts, are useful mainly in advanced forms.
into groups simply by inspection of gross form without But in the simpler, more primitive groups, critical
attempting to analyze what the differences and similari- analyses of the appendages and mouth parts have been
ties were. Six or seven groups were initially segregated found more revealing (Darwin, 1854; Pilsbry, 1916 ;
in this way. These were re-examined and re-sorted to Nilsson-Cantell, 1921). More recently, thin sections
some extent, ending up with five principal groups. have been used in attempts to distinguish between spe-
It was possible to arrange these groups, not by an cies (Cornwall, 1962), and to some extent between
intuitive process, but by inspection. The question then higher categories (Davadie, 1963). Thin sections have
became, what was in the gross form that allowed one proved extremely valuable in the present study in dis-
to make the separation? Capitular plates were com- tinguishing between both species and genera, as can
pared between groups for five categories, including 10 be seen in the systematic section on Balanomorpha.
characteristics. For example, the first category con- Their full value is yet to be realized.
sidered the general form of the individual plates: un- We have already alluded to the relative importance
modified, forked, quadrangular, triangular, etc. It of appendages and mouth parts in classification, and
immediately became apparent that species grouped they have been used extensively in this work. The
under what is now known as Litoscalpellum differ from methods of dissection and microscopical preparation
Neoscalpellurn by having the basal margin of the scu- have been described; thus, only a technical note on
tum entire rather than forked, the tergum and upper cirral counts needs to be made here. Cirri are nor-
latus normal rather than "V" shaped, and the rostra1 mally biramous, the exopod (outer) and endopod
latus wider rather than higher. Once the groups could (inner) being supported by a two-segment protopod,
pedicel, or peduncle. While in some cirripeds the rami
be defined morphologically in this manner, distribu-
are clearly "inner and outer" there is a tendency to-
tions were plotted and bathymetries were taken into
ward lateral flattening and longitudinal elongation so
consideration. Litoscalpellum had an Indo-West Pa-
that the outer becomes effectively anterior and the
cific-austral distribution between 100 and 2000 meters,
inner, posterior. Cirral counts are customarily given
while Neoscalpellum had a West Pacific-East Pacific- with the outer (anterior) above and the inner (poste-
North Atlantic distribution between 2800 and 5000 rior) below in the numerical notation for each side.
meters. Representatives of both were taken by the This being understood, the designations are unneces-
Eltanin in the Antarctic at comparable depths. On the sary and are not given in the cirral formulae in this
other hand, the species forming Annandaleum are work. The caudal appendages, when present are al-
strictly Indo-Pacific in distribution, between 200 and ways uniramous, and when represented by one or more
1250 meters, and no antarctic representatives have articles, the counts are included in the cirral formula
been found. Thus it can be seen that this method is after the notation for cirrus VI.

SYSTEMATIC ACCOUNT
K C:I T O I t I + : ( : E h r l ' A N T A R(:'l'J (: Familv SYNACOC,II)$E
Gruvel. 1005
ll~agno,ss: Six pairs of thoracic appendages; ab-
1. Fectling of1 1~lank1o11 I)y thoracic l i m l ~ s (cirri) . . . . . . . . 2 domen of 4 or 5 di5~inc.t5ornitcs and furta; carapace
I . l'arasi~ic; feeding t;y r n ( ; u ~ lparts.
~ or al~sorption t l l r ~ u g h distinctly 1)ivalvetl. Ectoparasitic on antipatharians,
hotly wall or procrsses, never by cirri . . . . . . . . . . . . . . . 3
c~ir~oids,and uphiu~oids.
2. Small to large attaclirtl barnac:lrs wit11 calcarvous platrs;
appendages forming cirral net evcnly distributed along Genus Ascothorax 1)jakonov. 101 L
thorax: Thoracica ................................. 4
2. hlinutc burrowing l)ar~rac.lcswithout calcarrous platrs; Dsag~~osis:First thoracic somite fuqed uith head;
appendages forming c:irral n r t concentrated a t dii;tal thoracic limbs somewhat reduced, Lrst and sixth pairs
end of thorax: Acrothoracica (only antarctic rcprc- hiramous or uniramous; abdomen of 2 or 5 somites.
sentative, Cryptophialus tontlinsoni sp. nov.) . . . .p. 25 F(toparasitic on ophiuroicis.
3. Adult with vrsrigial appcndagcs, parasilic on cchino-
derrns ant1 coelenterates: Ascothoracica (only known Ascothorax bulbosus Heegaarrl, 1051
antarctic rcprrcscntalive, Ascothorax bulbosus I-Jcc- Text-fig. 4
gaard, 1951, parasitic on ophiuroitl) ............. .p. 23
3. Adult without apl~mdages, parasitic 011 crustaceans: Asco~horaxbulbosi~sHeegaard, 1951. p. 1G6, figs.
Rhizocepl~ala (only known antarctic representative, 6 , 7.
Briarosaccus callosus Boschma, 1930, parasitic on IJ)iagr~osis:Four abdomirlal segments in both male
lithoditl crabs) ................................ p. 183
and female; thoracic appendages not reduced.
4. External form bilaterally symmetrical ................ 5
4. External form asvmmctrical: hox-like wall of 4 calcified fifaterial: Lliscovery, Sta. 27, type-locality off South
plates closet1 lry a singlc pair of movablc valvcs: Georgia (approx. 55OS, 37"W), 100 meters, 1 speci-
Verrucomorpha (only known a n ~ a r c t i c rcpresentativc men. William Scoresby, Sta. 42, off South Georgia
V ~ r r u c agibbosn Hoek, 18.83) .................. p. 735 (approx. 55's. 37"W), 175 meters, 1 specimen.
5. Body divisible into pedunc:le and ca~)itulum,the latter
always arinorcd with calrarcous plates: Lepado- Original Description: "'The parasite lies in the bursa
morpha ..........................................6 of the host, Amphiura belgicae Koehler and Amphiurn
5. Shell divisible: into wall of 6 o r 4, calcareous plates microplax Mortensen ; on1y one specimen was found
and operculum of 2 pairs of calcarcol~svalves (except in each host, and in both specimens a complementary
X m o b n l a i ~ u s : 6 plates cmlredtlctl in skin of ccta- male [dwarf male] was placed dorsally to the female
ccans, operc~llar platcs ahscnt) : Ralanomorpha ..... 7
underneath the cuticle of the female.
6. Capitul~irn 5 or 2 calcareous plates, peduncle
nakcd, pelagic: Lepotlidae ...................... .p. 29 "The body of both the male and the female is more
6. Capitulunl supporting morc than 5 plates, peduncle with or ltss regularly globular; it is totally covered with
nnmcrous calcareous plates, brnihic:: Scalpellidae. ..p. 37 a large leathery shell, corresponding to the shell of
7. Living on cetaceans: Ralanitlae in part (Coronula and the cypris stage. The two halves of the shell are com-
Xenobalanus) ................................ p. 178
pletely coalesced at the dorsal side and here the male is
7. Rcnthic: Kalarromorplia (see key lo families) ..... .p. 137
found lying on its side and forming an angle of 90
Order ASCOTHOR AClCA Lacaze-Duthiers, 18:N degrees with the body of the female. The free margins
of the shell are bent inwards and lie close to each
Diagnosis: Cirripedia with prehensile first antennae; other. In the female the shell is filled up with the
body enclosed in bivalved carapace; sexes separate eggs of the parasite.
or combined; mouth parts forming oral cone modified "The thorax is fixed to the shell by a bundle of
for biting and piercing; thoracic appendages basically strong muscles. The body is strongly curved and
natatory, first pair not conspicuously modified to form when the shell is re~novedit may remind one of an
accessory mouth parts; fifth abdominal segment hear- amphipod. The frontal part is rather heavy and with-
ing furca. Ectoparasitic arid endoparasitic on coelen- out any distinct segmer~tatior~, and the abdomen is
terates and echinoderms. formed By four segments. Stephcnsen (1935) in A.

Fig. 4. Ascothoraz bulbosus Heegaard: A-I, female; J-L, male. A, carapace removed; B, first antenna; C, carapace;
D-I, natatory thoracic appendages I-VI respectively; J, carapace; K, carapace removed; L, first antenna (redrawn from
Heegaard, 1951, text-figures 6 and 7 ) .
ophiocter~isfound 5 abdominal se,pents in the female, exopod, and in Stephensen's figure the exopod is
and 4 in the male. I can only find 4 abdominal seg- broken so that only the basal part of it is shown. The
ments in both sexes, for the segment which appears four following pairs of legs are very flat and broad,
to be the first one without appendages in reality bears lying one after the other as leaves in a book. The
the small sixth pair of curved thoracic limbs. sympod is divided into two joints, the coxa and the
"On examining my specimen I first thought that basis, and the exopod as well as the endopod is un-
it was A. ophioctenis Djakonov, for no difference jointed, but tipped with long marginal setae. The
could be seen externally. But on dissecting out the sixth pair of legs also has a two-jointed sympod. The
appendages I found that they differed from those two branches, the exopod and the endopod, are
figured by Stephensen. In his figures there is a returned to one side and placed in a dorsal direction
markable reduction in the limbs. There are no setae along the side of the body segment.
on one branch of legs number 2, 3, and 6, and the "The antenna was described as six-jointed by
other branch is two-jointed in all the limbs except Djakonov, but Stephensen could only see five joints.
in the first one. Neither this reduction nor this ar- In the female specimen of A. bulbosus the antenna
ticulation are found in my specimens. The first pair can be said to have six joints, for a weak suture
of legs has an unjointed sympod and a long and showed a coalescence of joints number 3 and 4. This
slender exopod which extends towards the mouth line of coalescence could only be found in the female.
cone and may have the function of a maxilliped. The I was not able to recognise it in the male, since the
endopod is only short and rudimentary. 1)jakonov other joints of the antenna were more coalesced than
(19141, according to his description, only found the in the female" (Heegaard, 1951, pp. 186-189).

SYSTEMATIC ACCOUNT 25
Order ACKOTHORACICA Gruvel, 1905 1. Four pairs of terminal cirri; single median dorsal fila-
mentary appendage, Cryptophialus (austral group:
Suborder PYGOPHORA Berndt, 1907
= Australophinlus Tomlinson, 1969) ............... 2
Diagnosis: Females with biramous terminal cirri 2. Spines guarding aperture three or four times a s long as
and complete alimentary canal, accompanied by wide ............. . C r y p t o p h i a h s tomlinsoni sp. nov.
minute nonfeeding males. 2. Spines guarding aperture less than twice a s long as
wide ............................................. 3
Family CRYPTOPHIALIDAE Gerstaecker, 1886 3. Operculum without projecting spinous lower angle;
margirl with pronounced median n'otrtl; surficial spines
Cryptophialidae : Batham and Tomlinson, 1965, in vicinity of lateral bar simple ..................
p. 142, synonymy. ................... .Cryptophialus turbonis Barnard
Diagnosis: Pygophora with 3 or 4 terminal cirri, 3. Operculum with projecting spinous lower angle; margin
without pronounced notch; surficial spines in vicinity
no caudal appendages, e!ongate labrum, 1 or 2 dorsal
of lateral bar complex, bifid or trifid ..............
filamentary appendages, and alimentary canal with ................ .Cryptophialus melampygos Berndt
gastric mill.
Cryptophialus tomlinsoni sp. nov.
Genus Cryptophialus Darwin, 1854
Cryptophialus: Batham and Tomlinson, 1965,
p. 142, synonymy. L)iugnosis: Four pairs of terminal cirri, mouth cirri
Diagnosis: As for the family. as mere vestigial knobs; thorax supporting single
Type-species: Cryptophialus minulus Darwin, 1854, median dorsal filamentary appendage; spines guard-
p. 566. ing aperture 3 to 4 times as long as wide and margin-
ally serrate.
KEY TO THE AUSTRALGROUPOF
Distribution: Recent, Antarctic, in dead stylasterine
Cryptophialus
corals, bryozoans, and shells of Bathylasrna corolli-
1. Three pairs of terminal cirri; 2 dorsal filamentary ap-
forme (Hoek, 1883).
pendages ........................Cryptophialus, s.s.
(known from both sides of the equator, not included Material: Eltanin, Sta. 933, off Antarctic Peninsula
herein). (61°25'S, 56"30'W), 300 meters, numerous speci-
A Crypfophiolus melompygos Ber ndt Crypfophialus furhnis Bar nard

Crypfophiolus tomlinsoni sp, nov. 8 tBrochyzopfes elllptica Codez
Chart 3. Distribution of Acrothoracica. The austral group of Cryptophialus and the Cretaceous Brachyzapfes elliptica
Codez.

Fig. 5. Austral group of Cryptophialus: A-0, Cryptophialus tomlinsoni sp. nov., from Antarc-
tica ; A-K, females ; L, burrow; M, cyprid ; N-0, male. P-R, Cryptophialus melampygos Berndt,
from New Zealand; S,T, Cryptophialus turbonis Barnard, from South Africa. A, apertural end
from right side; B, apertural spine of A enlarged; C, aprrtural end from right side, comb collar
extended; D, intermvdiate segments of last cirrus; E, mouth field including elongate labrum and
stubs of first or mouth cirri, viewed from below; F, mandible; G, first maxilla; H, second maxilla;
I, median dorsal filamentary appendage; J, opercular hars of C, seen from above; K, entire speri-
men, from right side; L, section of burrow; M, cyprid larva, from below; 1U, first antenna of male;
0, male, viewed from left side; P , lateral bar and marginal thickening; Q, apertural end, from
right side; R, apertural spines of Q, enlarged; S, apertural end, from right side; T, apertural
spines of S, enlarged. A-L, N and 0, Eltunin, Sta. 993 (South Shetland Islands) ; hf, NZARP
Endeavour, Sta. A-465 (Ross Sea) ; P-R, New Zealand and S,T, South Africa, specinlens
courtesy of J. T. Tomlinson.
mens in dead stylasterine coral; Sta. 10S9 (60°47'S, Depository: USNM Cat. No. 125218 (holotype; Sta.
53O3030'W), 641 meters, in br~ozoan. NZARP En- 933) ; 125219 (paratype; Sta. 933) ; 125220 (para-
deauour, Sta. A-465, off Victoria Land, Ross Sea types; Sta. 1089) ; New Zealand Oceanographic In-
(72'555, 175"30fE), 399 meters, in Bathylasrna stitute, Wellington (Endeavour, Sta. A-465).
shells. Description: Female-A simple sac with broad at-

SYSTEh'lATIC ACCOUNT
tachment disc and slender neck extending to aper-

ture, measuring approximately 2.5 mm long and 1.5
mm wide (fig. 5K). Chitinous opercular bars with
strongly produced inferior angles, each terminating in
a pair of strong spines, with a row of as many as 20
strong, serrated spines standing approximately 4 times
as high as wide along outer marginal surface, with an
inner row of comparable spines and scattered, fine,
long setae (fig. 5A, C, J ) . Lateral bar simple, without
expanded distal spinous portion (cf. fig. 5A, P ) . Fig. 6. Cryptophinlus tomlinsoni sp. nov.: Outlines
Thorax with single median dorsal filamentary ap- of burrow openings. Eltanin, Sta. 993 (South Shet-
pendage (fig. 51). Mouth parts and cirri typical of land Islands).
the genus; labrum long, terminating in a tuft of fine
6 specimens) ; surface covered with minute chitinous
setae; mandibles with 3 major teeth along cutting
beads and a few scattered minute setae; possessing
edge-1 superior, 1 median and 1 inferior, with
a pair of compound eyes, red in color in preserved
minute teeth or spines between; mandibular palps
material (fig. 5M).
long, spatulate, supporting a few terminal setae; first
maxillae simple, blade-like, supporting terminal cut- Aperture of burrow ( n z 1 6 )
(see fig. 6 )
ting edge of setae; second maxillae broad, with soft -
setae over inner surface and along outer margin be- Range X
low apex; first or mouth cirrus reduced to mere Length 0.536-0.368 0.485 mm
knob, supporting few setae (fig. 5E-H). Four pairs Width 0.245-0.140 0.196 mm
of terminal cirri; 2 pairs of long setae per article Length :Width 2.5 :1
along lesser curvature and 1 long seta at every second Etymology: Named for Dr. J. T. Tomlinson, for
or third articulation along greater curvature; all kindly providing comparative material he collected
setae with scattered, widely spaced setules (fig. 5 0 ) . in South Africa and New Zealand, and for his in-
First terminal cirrus with rami unequal in length; formative studies and discussions on acrothoracicans.
following pairs essentially equal; counts for 2 speci- Remarks: The taking of Cryptophialus by the El-
mens : tanin and Endeavour in the Antarctic to depths of
600 meters is of considerable interest because here-
tofore acrothoracicans have been known only from
Specimen 1 shallow water. To our general knowledge, the larvae
(Ross Sea) of truly deep-sea cirripeds have not been described
as possessing eyes, while in the cyprid of C. tomlin-
soni, the eyes are well developed and as obvious
as in littoral forms. This suggests that the collec-
Specimen 2 -16 -
24 25
-
25
- tions on hand may be from near the lower bathy-
(off Antarctic Peninsula) 20 24 24 25 metric limit for the species and/or that the species
has only relatively recently entered deep water.
Cryptaphialus tomlinsoni is comparable in struc-
ture to C. melampygos from New Zealand and C.
Male-Approximately 0.1 mm long, 0.07 mm wide, turbonis from South Africa (in having 4 pairs of
appearing as a simple sac with much convoluted in- terminal cirri and a single dorsal filamentary ap-
ternal material suggesting intromittent organ; at- pendage) and therefore belongs to the austral group
tached to female by pair of first antennae (fig. 5N, 0 ) . of the genus. It is more similar to the former than
Males found on either side of female, in area between the latter; in particular in the opercular apparatus,
upper portion of attachment disc and lateral bar, in the inferior angle of which is strongly produced and
clusters of as many as seven. spined, and in the surficial spines being complex
Naupliar stages passed through in egg; larva rather than simple. Cryptophialus tomlinsoni is sim-
emerging as cyprid stage measuring approximately pler in the development of the lateral bar system than
0.5 mm long and 0.2 mm wide (ratio of 2.31:l for C. melampygos, which has the most complex system

seen in the family. Because the lateral bar system is half the length of the complete cast, is elongatt. and
in itself a specialization in the acrothoracicans, find- occasionally slightly arcuate. The tapered anterior
ing its highest development in Cryptophialus, it fol- may be twisted either to the left or right, while the
lows that while C. tomlinsoni is more specialized in posterior is proportionately broader and rounded.
r
this regard than C. turbonis, it is at an intermediate
7
I h e anterior projects dorsally as an excentric apex

grade of construction that could have readily given that constitutes the highest part of the cast.
rise to C. melampygos. Considering the biogeography "Below the aperture the casts expand downward
o l the genus, with the more generalized austral group and anteriorly into larger sacciform bodies. The
( 4 terminal cirri, 1 filamentary appendage) in the antero-dorsal margin slopes down evenly from the
Antarctic, New Zealand, and South Africa, and the apex to reach a rounded antero-lateral margin. The
more specialized group ( 3 terminal cirri, 2 filat maximum width of the cast is at the antero-lateral
tary appendages) tending north and across the equa- margin. The postero-lateral margin slopes almost
tor, it seems reasonable to look for a southern hemi- vertically downward from beneath the posterior end
sphere origin and radiation for the family. Since of the aperture. In longitudinal section the casts are
C. turbonis from South Africa is the most generalized virtually symmetrical half ellipses. The fairly pro-
member of the genus, arid the greatest number of nounced transverse and longitudinal ribbing probably
species is from that region, it is suggested that South reproduces the structure of the surrounding calcite;
Africa is the center of cryptophialid diversification the ornament is dissimilar from the striation described
arid radiation, with descendants circling the globe by Tomlinson (1955, p. 99) which he has attributed
around Antarctica by the West Wind Drift. to the action of the boring teeth.
Saint-Seine, 195 L
Family ZAPFELLIUAF: Measurements of Casts
Diagnosis: Burrow depth at least one-half length, Length of Width of Ovprall Overall Maximum
Aperture Aperture Length Width I-lcight
without peduncular slit or marginal flange. The ap-
(mm) ( mrn i (mmi (mm) (mmi
propriateness of applying Linnean nomenclature has - - -
3.6 0.8 7.2 1.8 4.0

been questioned by Rodda and Fisher (1962) and
3.6 1.0 7.2 2.4 4.2
Ross (19651 , and there is in fact good reason to 3.4 1.0 6.8 2.0 4.5
voice concern when it is recognized that only Crypto- 2.2 0.72 4.2 1.1 2.4
phialus arid Trypetesa, among extant Acrothoracica, "Although the infilled cavities could have been
can be recognized presently by the form of their bur- made by a number of known boring organisms (nota-
rows alone. bly clionid sponges), they seem to be indistinguish-
Genus Brachyzapfes Codez, 1957 able from the cavities left by acrothoracic cirripeds
(Codez and Saint-Seine, 1957, pls. xxxvii-xxxix) .
Diagnosis: Form of apcrture elliptical. Burrows in
The largest of the cirripede casts from Alexander
belemriites and pelecypods. Island is considerably larger than ariy previously de-
Brachyzapfes elliptica gigantea Taylor, 1965 scribed from the Mesozoic. Simonizapfes elongata
Plate IV Codez, the largest acrothoracic cirripede described by
Codez and Saint-Seine (1957) has a maximum length
Brachyzaphes elliptica var. gigantea Taylor, 1965,
of 4.5 mm and width of 1.1 mm. If the biometrical
p. 39, figs. 3a-c.
analysis of Codez and Saint-Seine is followed. this
Diagnosis: Burrow aperture approximately y2 size difference could be thought of as having a spe-
length of complete burrow ; aperture slightly arcuate;
cific or even generic importance. However, the au-
burrow length greater than 4.2 mm.
thor is riot satisfied that figures representing the
Material: East coast of Alexander Island, Antarctic maximum dimensions of the cavities are more impor-
Peninsula (approx. 73 S, 6S0W 1 , Upper Aptian, tant than general form and outline, since such meas-
Cretaceous; numerous burrows. urements probably reflect the maturity of the indi-
Original Description: "Casts of cavities exposed on vidual cirripede.
a belemnite rostra1 surface reveal the outline of the "The Antarctic acrothoracic cirripedes most clearly
aperture and, if some of the surrounding calcite has resemble Zapfella Saint-Seine and Brachyzapfes Codez,
been etched, part of the broader under surface of two genera in the Zapfellidae. Brachyzapfes has been
the burrow. The aperture, which is approximately observed in belemnites and lamellibranchs, while

SYSTEMATIC ACCOUNT
Lepos austrolis Darwin v Lepos fasciculoris Ell is & Solander

A Lepos anafi'fero Linn6 v Conchodermo virgofum (Spengler)
Chart 4. Distribution of Lepadidae.
Zapfella has been found boring into molluscs, brachi- Order THORACICA Darwin, 1854
opods, corals and solid rock. The cavities made by the Suborder LEPADOMORPHA Pilsbry, 1916
Alexander Island cirripedes are not so deep as those Diagnosis: Body differentiated into capitulum and
made by Zapfella and the nearest specific match is peduncle, generally armored with calcareous plates;
with BrachyzapJes elliptica Codez, a Lower Creta- sexes separate or combined, hermaphrodites of some
ceous species. The Alexander Island Acrothoracica, species accompanied by complemental males.
however, are larger than those described by Codez
and Saint-Seine (1957) and therefore this difference Family LEPADIDAE Darwin, 1851
has been accorded varietal status. Lepadidae: Nilsson-Cantell, 1921, p. 233, syn-
"The infilled cavities are both superficial and deep- onymy.
ly buried. They may be scattered over the rostra Diagnosis: Capitulum armed with 5 approximate
of belemnites or locally congregated but each perfora- plates, 5 or 2 reduced, widely separated plates, or
tion is discrete and, as there is no distortion of the no plates; peduncle naked. Caudal appendages
surrounding laminae, the cavities were probably minute uniarticulate thoracic extensions, not sup-
bored following the death of the host. The exclusive- porting setae; first maxillae with step-like cutting
ness of belemnite rostra as hosts is probably due to edges. Hermaphroditic, generally pelagic, on floating
the thickness of the shell which provided the young objects and nekton.
larva with a firm anchorage. In several specimens
KEYTO ANTARCTIC LEPADIDAE
the cavities were confined to one side of the rostrum,
presumably the one farthest away from the mud. 1. Capitulum completely covered by 5 calcified plates
( L e p a s ) .........................................2
Perforation of the suggests that 1. Capitulum with 5 or 2 plates, much reduced.. ......
the shell was rolled. (Conchoderma) ................................. 4
"Although the bored belemnites described in this 2. Carina bent abruptly at submedian umbo, basal portion
forming a large disc; juveniles on small floating ob-
paper were collected from only three field stations,
jects, adults producing gas-filled float, cirri acantho-
belemnite rostra throughout the succession were found ~ o as d in Conchoderma ...........................
to be similarly affected" (Taylor, 1965, pp. 3 9 4 1 ) . ....Lepas ( D o s i m a ) fascicularis Ellis and Solander

2. Carina with basal umbo, inferior portion produced into prosoma as long as those at the base of first cirri
a relatively small fork or knob, on floatiug objects, but thin and inconspicuous; caudal appendages as
cirri ctenopod [Lepas (Lepas)] .................. 3 low blunt projections ........L. ( L . ) australis Darwin
3. Right scutum usually with an internal umbonal tooth,
4. Capitulum with 5 reduced plates, usually with longi-
left without; 1 pair of filamentary appendages on each
tudinal purple stripes; on variety of pelagic and float-
side, 2 on prosoma as long and as conspicuous as those
ing objects.. ... .Conchoderma virgatum (Spenglerj
at base of first cirrus; caudal appendages as slender
rounded projections ............................... 4. Capitulum with 1 pair of plates (scuta), with longi-
............................. L. ( L . ) anatifera Linnb tudinal purple stripes and large pair of fleshy apical
3. Each scutum usually with an internal umbonal tooth; 1 projections; on whales ............................
pair of filamentary appendages on each side, 2 on .................... .Conchoderma auritum (LinnC)
Fig. 7. Lepas (Lepcrs) rtnatijera Linnk: A, labrum and right palp, left palp deleted; R, right mandible;
C, inferior angle of right mandible enlarged; D, right first maxilla; E, inferior angle of right first maxilla
enlarged; F, right second maxilla; G, right first cirrus; H, right second cirrus; I, intermediate articles of
outer ramus of cirrus 111; J, pedicel of sixth cirrus and caudal appendage; K, penis. Eltanin, Sta. 122.

Genus Lepas Linnk, 1753 de Graaf, 1952), but both L. anatifera and L . australis
have 2 similarly placed pairs. However, a difference
Lepas : Darwin, 1851b, p. 67, synonymy.
in relative size has been noticed in the present study
Diagnosis: Capitulum with 5 approximate or nearly that may prove u~eful,and this is taken up in the
approximate plates. discussion of L. australis. There is also a difference
to be found in the form and relative size of the
Subgenus Lepas s.s. caudal appendages in these two species (cf. figs. 75,
Lepas: Pilsbry, 1907b, p. 79; Annandale, 1909b, 9F).
p. 72, synonymy. Lepas anatifera has been treated here, and in the
Diagnosis: Capitulum flattened laterally ; basal mar- literature in general, as a well-defined species. This
gins of scuta curving inward; plates approximate, could be misleading, because populations differ mark-
well calcified, sometimes thin but not papery; basal edly in appearance. Unfortunately, descriptions are
portion of carina developed into a knob or fork, not commonly based on material taken from bottoms of
a broad flat disc; cirri ctenopod. wide-ranging ships, and where the fouling occurred
is usually unknown. Workers attempting to analyze
Lepas (Lepas) anatifera Linni, 1758 such data are confronted with a proverbial bag of
Plate V C, Text-fig. 7 worms. In some cases the differences in form reflect
Lepas anatifera: Jennings, 1915, p. 285, 288, text- the genetic composition of the populations involved
figs. 1, la-c; 1918, p. 57; Barnard, 1924, p. 50 (key and can aid in defining populations inhabiting par-
to species) ; Nilsson-Cantell, 192933, p. 484, text-figs. ticular water masses, but these cannot presently be
la-i; 1930b, p. 247; 1957, p. 5 (principal citations: distinguished from differences due to environmental
Southern Hemisphere). factors. The solution would require worldwide samp-
ling on one hand, and experimental work with the
Diagnosis: Plates thin, smooth or delicately striated; effects of the environment upon development on the
right scutum usually furnished with internal umbonal other. While such large-scale projects are not likely
tooth; 2 conspicuous filamentary appendages on each to be practical for some time, careful analysis of locaI
side, 1 at base of first cirrus short, the other on the populations can be extremely valuable in forming a
prosoma long; caudal appendages consisting of slen- framework of information pinpointing critical areas
der rounded projections. in need of study. For example, L. anatifera from
Material: Eltanin, Sta. 122, off Tierra del Fuego the California Current differs considerably from what
(surface; 2 specimens, largest 7 mm capitular length). would be known as the same species in the central
Pacific, and it seems likely that we are dealing with
Depository: USNM Cat. No. 125221 (Sta. 122). two distinct populations. On the other hand, L.
Remarks: Lepas anatifera, a cosmopolitan species, australis from the coast of Chile and Islas Juan Fer-
is not known at latitudes higher than 60"s. It has nLndez is distinguishable from the antarctic form,
previously been recorded from Cape Town, Kermadec and in this case it appears that the variation is due
and Chatham Islands, and from Cape Horn. The to environmental factors rather than to a restriction
present material, taken off Cape Horn, constitutes in gens flow from one (segment of the) population to
the most southerly record, in a area of mixing with the other (Nilsson-Cantell, 192933).
the closely related antarctic species, Lepas australis.
Lepas (Lepas) australis Darwin, 1851
Lepas anatifela and L. australis are very similar
Plate V A, B; Text-figs. 8, 9
in appearance. The latter generally has an internal
umbonal tooth on both scuta rather than only on the Lepas australis Darwin, 1851b, p. 89, pl. I, fig. 5;
right scutum, and the capitular plates are generally Hoek, 1883, p. 41; 1907, p. 1 ; Weltner, 1895, p. 290;
thicker and less brittle. 1896, p. 63; 1897, p. 245; 1898b, p. 13, in part
As Nilsson-Cantell has remarked, the trophi and (non L. australis from Honolulu) ; 1899, p. 442; 1900,
cirri of different species of Lepas are remarkably p. 305; Gruvel, 1905, p. 109; 1907b, p. 161; Stebbing,
similar and characteristics of systematic value have 1910, p. 564; Jennings, 1915, p. 289; Nilsson-Cantell,
yet to be found. The number and disposition of the 1921, p. 237; 1926, p. 7 ; 192933, p. 487, fig. 2a-j
filamentary appendages, on the other hand, have been ( L . australis var. weltneri) ; 1930a, p. 210; 1930b,
useful in separating species of the genus (Broch, 1959; p. 247, text-figs. 10a-j; Weltner, 1922, p. 76, fig. 2;

Fig. 8. Lrpas ( L r p a s ) australis Darwin: A, labrum, left side and palps deleted; B, right palp; (1,
right mandible; D, right first maxilla; E, right second maxilla. Eltanin, Sta. 1185.
deGraaf, 1952, p. 5 (key to species) ; Nilsson-Can- Lepas analifera, except the shell is thinner, quite
tell, 1957, p. 15. brittle, and the scutal umbones usually each bear an
Diagnosis: Plates thin, brittle, marked only by faint internal tooth.
growth lines and striae; generally an umbonal tooth It is generally conceded, for species of Lepas
on both scuta; 2 filamentary appendages on each side, ( L e p a s ) , that the mouth parts and cirri are too similar
1 at base of first cirrus conspicuous, the other on to be taxonomically useful at this time. On the other
prosoma remarkably thin and inconspicuous; caudal hand, great weight has been given to the number
appendages consist of broad terminal projections. of filamentary appendages. However both L. australis
and L. anatifera have 2 pairs. In the numerous speci-
Material: Eltanin, Sta. 375 (12 specimens, all stages) ; mens of L. australis dissected, the filaments on the
Sta. 718 (SOSC 104, 2 specimens) ; Sta. 1161 ( 3 on prosoma were difficult to see; although long, they
pumice) ; Sta. 1162 ( 5 on pumice) ; Sta. 1167 ( 3 on were very fine. In the two specimens of L. anatifera,
p m i c e ) ; Sta. 1170 (10 on pumice) ; Sta. 1185 (30, however, these filaments were much thicker and ob-
some cyprids, on kelp) ; Sta. 1188 ( 5 on pumice) ; vious.
Sta. 1196 ( 7 plus several cyprids, on pumice) ; Sta. Differences noticed in the caudal appendages are
1200 (10 on pumice) ; Sta. 1201 (15) ; Sta. 1203 (34% also helpful in distinguishing between these two
plus 3 cyprids, on pumice) ; Sta. 1206 ( 5 ) ; Sta. 1220 species. In Lepas they are represented by simple
( 5 on pumice) ; Sta. 122% (7 on pumice) ; Sta. 1235 extensions of the terminal portion of the thorax, in
(1 on pumice) ; Sta. 1270 (4) ; Sta. 1428 ( 2 on L. australis as a pair of broad, blunt knobs, and in
pumice). L. anatifera as a pair of slender, rounded projections
Depository: USNM Cat. No. 125222 (Sta. 375, off (cf. figs. 9F and 75).
~ a i k l a n dislands) ; 125223 (Sta. 718) ; 125224 (Sta.
Subgenus Dosima Gray, 1825
1161); 125225 (Sta. 1 1 6 2 ) ; 125226 (Sta. 1167);
125227 (Sta. 1170) ; 125228 (Sta. 1185) ; 125229 Dosima: Pilsbry, 1907b, p. 81; Annandale, 190!!h,
(Sta. 1196) ; 125230 (Sta. 1200) ; 125231 (Sta. p. 72.
1201) ; 125232 (Sta. 1203) ; 125233 (Sta. 1206) ; Diagnosis: Capitulum inflated rather than laterally
125234 (Sta. 1220) ; 125235 (Sta. 1224) ; 125236 compressed; plates nearly approximate, calcareous but
(Sta. 1235) ; 125237 (Sta. 1270) ; 125238 (Sta. 14'28). thin and papery; basal margin of scutum flaring out-
Remarks: This widely distributed antarctic species ward; basal portion of carina developed into a large
is well known. It is quite similar in appearance to disc; cirri acanthopod, as in Conchoderrna.

Lepas (Dosimn) fascicularis Ellis and cosmopolitan in distribution, and is not kriowri to
Solander, 1786 occur in antarctic waters. While it has been reported
Plate V D; Text-fig. 10 from New Zealand and South Africa, the present
record from south of Cape Horn is apparently the
Lepas fascicularis: Jennings, 1915, p. 286; Pilsbry, first from South America.
1907b, p. 81, pl. IX, fig. 6 ; Barnard, 1924, p. 50 Lepas fascicularis is the only pelagic barnacle
(principal citations for Southern Hemisphere) . known to produce a float of its own. The cyprid
Diagnosis: As for the subgenus. larvae attach to floating objects (feathers, bits of
pumice, colonial hydroids such as Velella, etc.) in
Material: Eltanin, Sta. 117, off Tirrra del Fuego
the usual way, but the selection is clearly for small
(engine room strainers; 9 specimens, largest 15 mm
rather than for large objects. Growth proceeds rapidly
in capitular length).
after metamorphosis, but before the weight of the
Depository: USNM Cat. No. 125239 (Sta. 117). barnacle exceeds the buoyancy of the object, a highly
Remarks: Lepas jascicularis, like L. anatifera, is vacuolatetl secretion is deposited around the attach-
Fig. 9. Lepas (Lepas) australis Darwin: A, first cirrus; B, second cirrus; C, pediccl and basal segments of
third cirrus; D, penis; E, intermediate articles of third cirrus; F, caudal appendage. Eltanin, Sta. 1185.

Fig. 10. Lepas (L)osirt~u)fascicularis Ellis and Solander: A, labrum, left side and palps deleted; R, right palp; C,
right mandible; I), right first maxilla; E, right second maxilla; F, first cirrus; G, pedicel and basal segments of
second cirrus; H, third cirrus; I, pcnii. Eltanin Sta. 117.
ment area and soon the barnacle's position at the sea necessity of locating and attaching to a small object
surface is self sustaining. Because virtually any ob- first, is an evolutionary advance yet to be achieved.
ject floating at the sea surface commonly becomes
Genus Conchoderma Olfers, 1814
fouled bv various s ~ e c i e sof Lepas, it can be inferred
that pop;lation density is limited in good part by the Comhoderma: Nilsson-Cantell, 1921, p. 2m,
availability of sites. Lepas fascicularis apparently Synonymy-
selects objects too small to be suitable for other species, Diagnosis: Capitulum usually bearing a few broad
and in this way has avoided competition with them. purple stripes and 5 or 2 reduced plates; cirri acan-
The ability to produce a float initially, without the thopod.

Conchoderma virgatum (Spengler, 1790) fork or knob; terga long and narrow rather than
Plate V E; Text-fig. 11 triangular; scuta Y-shaped; 2 or 3 purple longitudinal
Conchoderma virgatum: Nilsson-Cantell, 1921, p. stripes on sides of capitulum and peduncle.
242; Jennings, 1915, p. 287; 1918, p. 59; Barnard, Material: Eltanin, Sta. 117, off Tierra del Fuego
1924, p. 61; Nilsson-Cantell, 1939, p. 236; 1957, p. 6 ; (engine room strainer; 4 specimens, largest 17 mm
Clarke, 1966, p. 3 (principal citations for Southern capitular length).
Hemisphere). Depository: USNM Cat. No. 125240 (Sta. 117).
Diagnosis: Five reduced plates; carina without basal Remarks: This cosmopolitan species is known from
Fig. 11. Conchodermu virgatum (Spengler) : A, labrum, lateral margins and palps deleted; E, right palp;
C, right mandible; D, right first maxilla; E, right second maxilla; F, first cirrus; G, second cirrus; H, third
cirrus; I, whorl of spines around greater curvature cf sixth cirrus, enlarged; J, penis. Eltanin, Sta. 117.

New Zealand, South America and South Africa, but grow, the terga always increase a little, and sometimes
has not been reported from the Antarctic. It attaches to such a degree as to be even thirty or forty times as
to a wider variety of floating objects and nektorl long as carina. When most developed they are not
than L ~ p a sand apparently stems from the stock that above one third as long as the scuta, to which they
gave rise to C. auritum occurring on cetaceans, Alepas lie at nearly right angles; they consist of imperfectly
occurring on scyphomedusae, and perhaps Anelmma calcified plates, square at both ends, slightly broader
which is wholly parasitic on certain dog fish. Varieties and thinner at the end towards the carina, where they
have been described, but the present material is entire- are a little curled inu'ards, than at the opposite end;
ly typical. they are not quite flat in any one plane; internally
they are slightly concave; finally, they nearly resemble
Conchoderma auritum (Linn6, 1767) in miniature the terga of C. virgata. In full grown
Plate I11 specimens, the terga almost invariably drop out and
Conchoderma auritum Darwin, 1851b, p. 14.1, pl. 3, are lost; but ever1 in this case, a long brownish cleft
fig. 4<; Clarke, 1966, p. 4. in the membrane of the capitulum marks their former
position. The orifice of the capitulum is usually notch-
Diagnosis: "Capitulum with two tubular ear-like ed between the terga, or between the clefts left by
appendages, seated behind the rudimentary and often them; on each side of the notch there is a slight pro-
absent terga; scuta bilobed; carina absent, or quite minence. In some few cases, however, there is no
rudimentary; peduncle long, distinctly separated from trace of this notch. Behind the terga or the clefts.
the capitulum. the great ear-like appendages are situated.
"Filaments attached to the pedicels of the second "Carina, rudimentary and often absent; it is point-
cirrus; two upper spines of the maxillae pectinated" ed-elliptical, and is rarely above the 1/40th of an inch
(Darwin, 1S51b, p. 141). long. After arriving at this full size, calcareous matter
Supplementary Description: '"The capitulum is slight- is added to the under surface over a less and less
ly compressed, almost globular, composed of thick arca, so that it becomes internally pointed, and finally,
membrane, with two large, ear-like, flexible, tubular, in place of calcareous matter, continuous sheets of
folded appendages, at the upper end, opening into chitine are spread out beneath it; hence, during the
the sack. These appendages are seated behind the disintegration of the outer surface, the carina comes
rudimentary terga when such are present, or behind to project more and more, and at last drops out;
the spots which they would have held if not aborted. subzequently, even the little hole in which it was
In a young condition they are tubular, but not folded; imbedded, disintegrates and disappears.
and often either one or both are at first imperforate. "Peduncle, cylindrical, distinctly separated from the
They are formed externally of the outer membrane czpitulum, and generally twice or thrice as long as it:
of the capitulum (rendered thin where folded), and the thickness of the outer membrane generally great,
internally of a prolongation of the inner tunic of the but variable: surface of attachment variable, either
sack; between the two, there is, as around the whole pointed, or widely expanded, or formed into divergent
sack, a double layer of corium. These appendages projections.
are sometimes very nearly as long as the whole capi- "Filamer~taryAppendages, seven on each side, high-
tulum: a section near their bases is sub-triangular. ly developed, long and tapering; there are two beneath
"The Scuta, as well as the other valves, are im- the basal articulation of the first cirrus, and one on
perfectly calcified: shape, variable. They usually the posterior margin of the pedicel of each cirrus,
consist of two lobes or plates, placed at above a right excepting the sixth pair; the filaments on the pedicels
angle to each other, and rarely almost in a straight are nearly twice as long as the cirri themselves.
line; the lower lobe is more pointed and narrower "Mouth, mandibles, with the five teeth nearly
than the upper; the two correspond to the lower and equidistant, and towards their bases finely pectinated
middle lobes in the scuta of C. virgata, the upper one on both sides; inferior angle rudimentary, often repre-
being here absent. sented by a single minute spine: in one specimen,
"The Terga are developed in an extremely variable there were only four teeth on one side. Maxillae, with
degree; they are often entirely cast off and absent. five steps, not very distinct from each other, with the
In very young specimens, they are of the same length first step much curved. The larger of the two upper
with the carina, but after the carina has ceased to unequal spines is pectinated, like the teeth of the

mandibles; there is a third long finer spine beneath 5. Rostra1 latus wider than high ....................... 6
6. Tergurn and basal margin of scutum forked, capitular
the upper large pair. plates reduced . . . . . . . . . .Gyrnnoscalpellum gen. nnv.
"Cirri rather short, broad, with the anterior faces 6. Tergum and basal margin o l scutum not forked.. .... 7
of the segments protuberant, especially those of the 7. Capitular plates of adult approximate. ...Arcoscalpellum
first cirrus and of the anterior ramus of the second 7. Capitular plates of adult reduced ....................
pair: spines on the anterior cirri doubly serrated. ........................... Litoscalpellum gen. nov.
Posterior cirri, with the intermediate spines between Genus Smilium Leach, 1825
the pairs, long; dorsal tufts, minute. On the lower
segment of the pedicels of the four posterior cirri, Smiliuin: Withers, 1053, p. 168, synonymy.
there are two separate tufts of bristles. Remarks: The genus Srnilium was instituted by Leach
"Colours extremely variable; sometimes five longi- (1825, p. 209) for an undisclosed species. The month
tudinal bands of dark purple can be distinctly seen following the publication of Leach's paper, Gray
(as in C. virgata) on the peduncle, these bands be- (1825, p. 100) validated this taxon through assign-
coming more or less confluent on the capitulum: at ment of Smilium peronii, an Australasian species.
other times, the capitulum is more or less spotted, or During its history, Smiiium has held full generic rank,
often nearly uniformly purple: the sack, cirri and as at present, but more commonly subgeneric rank
trophi are, also, purple" (Darwin, 1851, pp. 142-144). under Scalpellum. Since the inception of Smilium
Family SCALPELLIDAE
Pilsbry, 1916 many workers have initially, or through subsequent
study by others, assigned species to this genus:
Scalpellidae: Withers, 1953, p. 100, synonymy.
S. acutum (Hoek) , 1G83, p. 80.
Diagnosis: Capitulum armed with more than 5 cal- S. scorpio (Aurivillius), 1894, p. 46 (syn.: Scalpel-
careous or partly calcareous plates; covered lum sexcornutum Pilsbry, 1897, p. 723, fide
with rows of calcareous scales. Hiro, 1933, p. 16; Calantica pedunculostriata
Remarks: The family Scalpellidae is a heterogeneous Broch, 1931, p. 3, fide Hiro, 1933, p. 16;
agglomeration of numerous groups. Many of these Scalpellum verticillutum Miers, MS, fide Gru-
groups represent obviously different and diverging vel, 1905, p. 34).
lineages but the revisionary work necessary to form S. squamuliferum Weltner, 1894, p. 80.
concise definitions has not been carried out. It is not S. pollicipedoides (Hoek) , 1904, p. 92.
deemed practical at this time to broadly subdivide the S. bengalense (-Annandale), 1906, p. 395 (=Euscal-
Scalpellidae, largely because it is beyond the scope pellum bengalense, fide Pilsbry, 1908, p. 108).
of this study. However, subdivision of this family S. aries (Hoek) , 1907b, p. 63.
into such groups as the pollicipoids, arcoscalpelloids, S. uncus (Hoek), 1907b, p. 67.
smilioids, etc., should be a major goal of future S. kampeni (Annandale) , 1909a, p. 267 (=Scalpel-
studies. In the present study certain groups have been lum pilsbryi Gruvel, 1911b, p. 290 = Calantica
removed from the largest genus among the Scalpel- pilsbryi, fide Hiro, 1932, p. 468).
lidae, Arcoscalpellum, and what were formerly Meso- S. parvulum Withers, 1914, p. 496.
scalpellum and Neoscalpellum have been extensively S. nudipes (Annandale) , 1916, p. 287.
revised. S. hypocrites Rarnard, 1924, p. 14.
S. sinense (Annandale) , 1910c, p. 2 11.
The Cretaceous species, S. parvulum, is questionably
1. Carina angularly flexed ............................. 2 assigned to this genus, and S. hypocrites, which pos-
1. Carina evenly curvet1 ............................... 4 sesses only 9 or 11 rather than 13 capitular plates, is
2. Scutuln and tergum lacking cleft in basal margin.. .... retained incertae sedis.
........................................ Scalpellurn
2. Scutum and tergum with cleft in basal margin.. ...... 3
In view of the fact that Smilium consisted of a
3. Inframedian latus essentially rectangular, higher than heterogeneous assemblage of generally unrelated spe-
wide ............................. .Brochia gen. nov. cies, Pilsbry (1907b, 1908), I-Iiro (1932), and With-
3. Inframedian latus V-shaped, wider than high ........ ers (1935, 1953) split off many of the recent species
........................ Australscalpellurn gen. nov. known to them (excluding S. hypocrites), and assign-
4. Capitulum with subcarina ................... .Smiliarn
4.. Capitulum without subcarina ....................... 5
ed them to CaZuntica (sensu lato). The remaining
5. Rostral latus higher than wide, capitular plates re- species, of which there are six ( S . peronii, S. acutum,
duced (see key, p. 9 3 ) . . . . . . . . . . . . . . . .Neoscalpellurn S. uncus, S. sinense, S. scorpio, S. a r i a ) , and question-

Fig. 12. S171ilirrnl crc.crtu~ti ( H o c k ) : A, side vicw of lierrr~apliroditr; LI-D, sidc virw of cornplcmcntal
malcs; E, labrurrr :ultl right palp; F, left palp; G, mandible; kl, enlarged view of third toorh and in-
ferior angle of rr~antlihlr; I, maxilla I ; J, maxilla I I ; I<, cirrus I ; L, intermediate artirlc. of cirrus
V I ; hl', pc%tlic:el of c,irrus VI showing relationshil)s of cirrus, cautlal apl~rntlagc ant1 penis; N, cn-
largcd vicw of c ~ ~ u d apperrdagc.
al Eltanin, Sta. 1429.
ably S. hypocrites, have been retained in Smilzum, ~ t u d yto undertake a rcvision of Sn~iliurnand Calan-
which still consists of an assrmhlage of only distantly lira since there is hut one representative of the genus
related species. The removal arid subsequent assign- Srniliurr~contained in the Eltanin collections.
ment of species of Sn~rlzurn to Calantica has only
served to weaken the tiefirrition of Culantica, and to Smilium rtcutum (Hoek, 1G83)
further complicate the drlimiting morphological fea- Plate V F; lext-fig. 12
tures of these two groups of primitive lepatlomorphans. Scalpellz~macuturn Hoek, 1883, p. 80, pl. 3, fig. 19,
This problem has plagued many taxonomists (see pl. 8, hg 12; lZ84, p. 20; Weltner, 1897, p. 246;
Hiro, 1933), but it is beyorid the scope of the present Gruvel, 1905, p. 56, Gg. 60; Hoek, 1907b, p. 64, pl.

7, fig. I ; Gruvel, 1012, p. 2 ; lloek, 1913, p. xiv. margins of about equal length; umho apical. Rostral
Scalpell~rrn lonsiroslr~~nz.Gruvel, 100011, 1). 100; latus 4-sidcd, and also essentially triangular; plate
19023, p. 70, pl. 2, figs. 1, 5 ; 1003, p. 57, fig. 61; smaller than carinal latus; rostra1 margin convex,
Weltner. 1022, p. 70. carinal margin slightly concave; rostra1 and c,arinal
S~railiurra acuturn: Pilsbry, 1008, p. 107; Icriiger. margirls allout equal in length; urn110 apical. Rostrum
1911. p. 15; Gruvel, 1020, p. 12, pl. 2, l ~ g 7. ; Uroch. large; nearly 3 times as largr as subcarina; plate
1022, p. 234; 1031, p. 11, Gg. 5 ; Hiro, 1033, p. 17, externally carinate; rhomboid uheri viewed head-on;
pl. 1, fig. 3 ; Stubbings, 1036, p. 57; Hiro, 1037. p. umbo apical.
300; Nilsson-Cantell, 1938, p. 6 ; Kriiger, 1010, p. l'eduncle about height of capitulum (pl. V
ll00; Utinomi, 1958, p. 283. F) ; surface completely covered by close-spaced, im-
Scall~ellurrz. (Snziliurn ) acutuna: Annandale, 101011, bricating scales; scales ~ i e a ~ lec~uidimensiorial:
y free
p. 154; 1916, p. 120, pl. 7, fig. 4; Calman, 101:;. p. edges somcwl~atevenly rounded.
101; Nilsson-Cantcll, 1921, p. 170, text-l~g.23.
Measurements of the single hermaphrodite in the
Scalpellurn has~aturn Weltner, 1022, p. 68, pl. 2,
Ellanin collections are as follows: overall height 13.1
fig. 3.
mm, c a ~ i t u l a rheight 9.4 mm, capitular width 5.3 mm.
Diagnosis: Carina simply bowed; tectun~ (roof)
Labrum bullate; periphery bearing short, stiff
evenly rounded; umbo apical. Upper latus largest
bristles; surface free of bristles; crest armed with
of latera; occludent margin same height as carinal
about 40 unequally spaced, triangular or M-shaped
margin of scutum ; plate occuy y ing completely space
teeth; area on both sides of thickened membrane of
between scutum and carina. Peduncle of hermaph-
mouth cavity lined with parallel rows of small spines
rodite covered with closely spaced, imbricating scales.
in clusters of 3-5. Palp very narrow, conical; superior
Anterior ramus of cirrus 1 about ?/$ longer than pos-
proximal margin bearing numerous, short setules;
terior ramus. Third tooth of mandible with subsidiary
distal superior margin with setae; lateral face and
teeth along superior slope. Capitulum of males covered
inferior margin free of setae. Mandible with 4 teeth
with 7 calcareous plates; peduncle unarmored.
including inferior angle; first tooth well separated
Materzal: El~anzn,Sta. 1429, Campbell Plateau, be- from second; second, third, and inferior teeth es-
tween Auckland and Antipodes Islands (45O15'S. sentially equidistantly spaced; superior margin of
172O14'E) 320-361 meters, 1 specimen with 3 at- thircl tooth and inferior angle serrate; about 12 teeth
tached malcs. on margin of inferior angle fig. 12G, H ) . Maxilla I
Llcpository: USNM Cat. No. 125211. highly spinose; cutting edge steplike; spines divided
Supplemental y Description: Hermaphrodite - Capi- into 3 clusters; apical cluster uith 2 large stout spines
tulum armed with 1 3 fully calcified, approximate anti 5 short, thin spines; moderate notch separates
plates; elongate-oval in outline; cuticle thin and not apical cluster from medial cluster; medial cluster con-
hirsute; plates ornamented with unequally spaced tains 9-11 long, thin spines. 1VIaxilla I1 triangular in
growth lirlcs only ; occludent margin essentially outline; marginal setae divided into distinct clus-
straight; carinal margin strongly convex (fig. 1 2 A ) . ters; setae of superior proximal cluster longer than
Tergum somewhat triangular; basal and carinal mar- those in other 2 clusters.
gins slightly convex; distal extremities of plate acumi- Rami of cirrus I grossly unequal (fig. 12K) ; pos-
nate; umbo apical. Scutum subparallelogrammic; terior ramus about 3 :; length of anterior ramus;
tergal margin concave; carinal, basal, and occludent intermediate articles not significantly protuberant;
margins convex; height of plate more than twice inner faces of both rami heavily clothed with random-
width; umbo apical. Carina bowed, roof simply and ly arrayed, long setae. Cirrus I not greatly modified;
evenly rounded; umbo apical. Upper latus of 5 cirrus 11 not modilied, approximatirig cirri 111 and
unequal sides; scutal margin concave, others convex; 1V. Each articulation along greater curvature of in-
scutal margin longest; tergal and basicarinal margins termediate articles of cirri 11-VI with 1 long and 1 or
of nearly equal length; umbo apical. Subcarina small; 2 shorter setae; at distal erld of each intermediate
triangular; umbo apical; apex of plate overlies base article parallel to articulation there is a row of ctenoid
of carina. Carinal latus large, 4-sided, although scales; interarticular areas along greater curvature
essentially triangular in outline; lateral and carinal free of setae; setae absent from inner lateral faces of
margins nearly equal in lcngth; basicarinal and lateral intermediate articles. Chaetotaxy ctenopotl; 4-5 pairs

on cirri 11 and 111; 5 pairs on cirri IV-VI, 1 or 2 northeastern Atlantic S. acutum ranges in depth from
short setae situated near bases of major setae. Cirral 174.0 meters to 24480 meters (Hoek, 1883; Gruvel,
counts are given below : 1902a, Calman, 1918; Gruvel, 1920; Weltner, 1922).
In the Japan archipelago it ranges from 6 1 meters to
287 meters; whereas, further south in the Moluccas,
and the immediately adjoining seas, the depth range
is 255 to 1264 meters (Nilsson-Cantell, 1921; Hoek,
1907b; Broch, 1922: Broch, 1931; Hiro, 1933;
Utinomi, 1958). Of the few widely scattered occur-
rences in the Indian Ocean, S. acutum ranges from
Caudal appendage moderately large, consisting of 768 meters, off southwestern Sumatra (Weltner,
a single article (fig. 12M, N ) ; approximately y2 1922 1 , to 896 meters, off the Andaman Islands in the
height of first segment of pedicel of cirrus VI; num- Bay of Bengal (Annandale, 1910a), to 2194 meters,
erous setae arrayed along both margins and apex; in the Gulf of Aden (Calman, 1918). In the south
apical setae about height. Pacific the bathymetric range of this species is 320-
Penis moderately long; surface weakly annulated 361 meters at Eltanin, Sta. 1429, and farther north.
and sparsely hirsute; numerous terminal bristles pres- in the region of the Kermadec Islands, it is 950-1152
ent. meters ( Hoek, 1883) .
Complemental males found attached externally to Based upon the foregoing, S. acutum can be con-
membrane between occludent margins of scuta im- sidered a cosmopolitan deep-water species, but it is
mediately below orifice; capitulum covered with 7 not known south of the Antarctic Convergence.
plates (carina, paired scuta, terga, and rostral latera;
Genus Euscalpellum Hoek, 1907b
peduncle devoid of scales; rostral latera often largest
of the capitular plates) ; ranging in height from 0.55 Diagnosis: Capitular plates 15; rostrum large and
mm to 0.95 mm (fig. 12B-D) . prominent ; subcarina well developed ; inframedian
Remarks: Morphologically, S. acutum is more close-
latus diamond-shaped.
ly related to several species of the Cretaceous to Re- Type-species: Scalpellum rostratum Darwin, 1851b,
p. 259, by subsequent designation.
cent genus Euscalpellum than to any other species of
Smilium. It can be assumed that they share a com- Euscalpellum antarcticum Withers, 1951
mon ancestry. The major difference between Euscal- Plate VI A-E
pelzum and S. acutum is that in the latter the in-
Eescalpellum antarcticum Withers, Is], p. 157,
framedian Iatus has dropped out, and is compensated 12, figs. 2a4b.
for by the enlargement of the diamond-shaped upper
Diagnosis: "An Euscalpellum with the peduncle com-
latus and migration of the rostral latus to a more
paratively wide, showing some curvature; plates de-
central position.
veloped for its whole length, usually formed of an
There is considerable confusion regarding the num-
oblong block of calcite extending inwards to the sub-
ber of plates covering the capitulum of the males in
median canal, close-set, and generally with a small
both Smilium and Calantica. Most workers report the
outer face, variable, but often elongated, somewhat
presence of six plates, i.e., carina, rostrum, and paired
rounded transversely, and tapering towards the apex.
scuta and terga. This is seemingly correct when the
Capitular valves unknown" (Withers, 1951, p. 157 ) .
male is mounted and viewed from the lateral side
only. However, what has been interpreted as the fii'aterial: On the slopes between Dagger Peak and
rostrum, when viewed from head-on, is seen to be Comb Ridge, The Naze (63'555, 57030fW), type-
paired, and consequently must be interpreted as being localit~, Senonian, Cretaceous. The saddle be-
the rostra1 latera. tween the two eminences, Humps IsIand (63'595,
Smilium acutum was first described by Hoek (1883) 57025'W) Upper Cretaceous.
from two widely separated regions, one near the Depository: British Museum (Nat. Hist.) In. 43813
Azores in the Atlantic, and the other near the Kerma- ( h o l o t y ~ eThe
, Naze) ; 4 3 8 1 u 3 8 1 5 (paratypes, The
dec Islands in the southern Pacific. The first locality Nazei ; 4390643907 (paratypes, Humps Island).
cited, Challenger, Sta. 78 (37"24'N, 25"13'W, 1828 Original Description: "The peduncles vary in width,
meters), is here selected as the type-locality. In the but are comparatively wide. There is considerable

variation in the shape of the outer faces of the plates, Original Description: "Carina oblong, narrowing
for they vary from as long as wide to three times as apically and widening basally. Inner surface thickened
long as wide. The plates are somewhat projecting, apically and crossed by slightly oblique growth lines.
transversely rounded, taper towards the apex, and the Outer surface not seen.
umbo often stands out prominently, as can be well "Scutum sub-rhomboidal, moderately convex trans-
seen in the holotype. The holotype represents part of versely with a rounded apicobasal ridge and a similar
a peduncle broken longitudinally down the middle, though less pronounced ridge extending from the
and shows the solid oblong plates almost horizontally apex to the middle of the basal margin. The basi-
inclined on the left side, and obliquely inclined up- lateral angle is acute, while the lateral margin appears
wards on the right side; they extend inwards nearly to be relatively straight. The apex is incomplete and
to the median canal which is fairly wide. This is the occludent margin and rostra1 angle are obscured.
the structure seen in three of the peduncles, and is Growth lines are fairly prominent. No internal moulds
probably normal. showing the adductor muscle scars are preserved. The
Measurements: "Holotype, length 44.5 mm, breadth terga are lozenge or sub-rhomboidal in outline, feebly
23.7 mm. convex transversely and acutely terminated apically
"The longest part of a peduncle is exceedingly curi- and basally. The apices are curved towards the as-
ously developed; it is not known which part of the sumed position of the scutum. A rounded, sub-medial
peduncle it represents, but is possibly the upper part. and slightly curved apico-basal ridge, which narrows
Outwardly the plates are very much elongated, some apically, divides each plate. A second broader and
are very large, long, and wide, and others long and more elevated ridge follows the occludent margin be-
narrow; in some cases the sutures between the plates fore joining with the scutal margin. The scutal mar-
are well seen, and in others they do not extend for gin, which is approximately the same length as the
the whole length of the plate, and certainly give the occludent margin, is straight. The upper carinal
appearance that some of the plates are incompletely margin, which is longer than the lower carinal margin,
fused together. Looking at the top of this peduncle joins the lower carinal margin in a rounded obtuse
it was thought at first that the plates were of the angle. Both halves of each tergum are ornamented
shape seen in the holotype, that is they were oblong with upwardly inclined growth lines and longitudinal
blocks laid one upon the other. Instead they are the ribs. Where growth lines and ribs intersect, small
inwardly directed portions of these outer, much tubercles are occasionally developed. No ribbing can
elongated and variably shaped plates; they differ be seen in specimen KG.11.29 but several of the
markedly in shape, length, and in the degree to which growth lines are thickened. The inner apical margins
they extend inwards. This is a somewhat different of several terga are ornamented with growth lines.
development from that of the holotype" (Withers, According to Withers (1928, p. l o ) , these growth
1951, pp. 157-158). lines are caused by recession of the corium. As a
result, the apices of the plates project freely beyond
Genus Cretiscalpellum Withers, 1922 the capitulum.
Diagnosis: Capitular plates 17, including 4 pairs of "In specimen KG.11.29 the prominence of the
large, overlapping and little-modified lower latera; tergum's apico-basal ridge, the absence of longitudinal
subcarina much larger than rostrum; all umbones ribbing and the variable thickness of the growth lines
apical. suggest that this may represent a separate species. An
Type-species: Pollicipes unguis J. de C. Sowerby, adjacent tergum and scutum may form part of the
1836, p. 335, by subsequent designation. right side of a capitulum.
Cretiscalpellum aptiensis antarcticum "The Alexander Island cirripedes are distinguished
Taylor, 1965 from Zeugmatolepas georgiensis on the basis of the
relative proportions of the terga (those of 2. georgien-
Plate VI F-H
sis being much broader than the plates described),
Cretiscalpellum aptiensis var. antarcticum Taylor, the relative lengths of all four tergal margins, the
1965, p. 37, figs. 1, 2 a, b. prominence of the tergal apico-basal ridge and the
Material: East coast of Alexander Island, Antarctic presence of an apico-basal and occludent ridge on the
Peninsula (approx. 73"S, 6S0W), Upper Aptian, scutum.
Cretaceous, 4 specimens. "The pedunculate cirripedes of Alexander Island

ANTARCTIC CIKRIPEDIA
(:hart 5 . ~ ~ n Arcos~alprlluv~.
(:irrumdntnrctic d i s t r i b u t i o n of the c o ~ n ~ o p o l i tEmus
differ from Scalpellurn iArcoscalpellum) comptum in approx. 55"S, 38OW I . U p p ~ rAptian; two incomplete
lacking prominent longitudinal ridges on the carinal capitula and numerous individual plates, particularly
side of each tergum. Moreover, the occludent ridge scuta. Senckenberg. Mus. Nos. 328 and 404. Only
terminates at the scutal angle, whereas in Scalpellurn species of genus of Aptian age.
( A . ) comptum, the ridge extends to the middle of the Diagnosis: Tcrgum short, wide; upper carinal mar-
scutal margin. Since Sralpellurn ( A . ) simplex and gin shorter than lower margin; occludent margin
Scalpellum ( A . ) accun~c~laturr~
are each represented by much shorter than scutal margin. Scutum with hardly
a solitary carina, they are unsuitable for comparison. any trace of an apicohasal ridge, tergal margin short.
"In Cre~iscalpellurnaptierlsis the tergal half of the Distinguished from Z . ( ? ) hausmanni (Koch and
scutum is wide and comparatively flat, whereas in the Uutiker) hy the much wider and shorter tergum and
Alexander Island cirripedes the tergal half of the virtual absence of apicol~asal ridge. Similar to Z.
scutum is narrow and steeply inclined. Therefore, concinr~a t Morris) as regards the tergum, but dis-
this difference has becrl accorded varietal status. No tinguishetl hy the proportionally longer carinal mar-
reconstruction of the complete capitulum of each speci- pin, shorter occludent margin ant1 less pronounced
men has been attempted" (Taylor, 1965, pp. 38-39). carinal angle.
Genus Zeugmntolepns Withers, 1913
Genus Arcoscalpellum Hoek, 1907
Iliagnosis: Capitulum like that of Scalpellurn, but
Arcoscalpellzcm: Withe~s,1053, p. 195, synonymy.
with at least 34 plates. including 3 or more ~ h o r l s
of lower latera; sculal urn110 apical in Jurassic forms, 1)iagnosis: Capitulum of female or hermaphrodite
subcentral in Cretaceous forms. a r m ~ d with 11 fully or partially calcified plates;
Type-species: Zeugrr~atolepas nhockleri Withers, carina with apical or slightly sul-)apical umbo, the
1013, p. 93G, by original designation. plate evenly curved, nevrr angularly flexed; tergum
never forked; scutum with apical, never subapical.
Zeugmatolepas georgiensis Withers, 1913 urn110 ; caudal appendage usual1y uniarticulate ; dwarf
Plate VII or complemental male sack-like, not divided into
Zeugmatolepas georgierzsis Withers, 1947, p. 18, pl. capitulum and prduncle, with or without 4 rudimen-
1 , figs. 1-2. tary calcareous plates.
Material: Annenkov Island, N E Coast (South Georgia, Type-sprcies: Scalpellum velutinum Hoek, 1883

SkSTEMATIC ACCOUNT 43
( = ScaliDeLlurr~rr~ic/le[ottiar~ulr~
Segueny,a, 1876 = 13. It~frarrrrtlian ldtus more thdn 3 t1rnc.5 as high as %id<.
. . . . . . . . . . . . . . . . . . . . . . . . . .A. brevecarinaturn (Hock
13. Inframctlian latus less than 2 limes as high as witle. .14
Aerr~arks: There are at present more than 200 fossil 14. Carina with subapical urnho, infrarriedian latus crlrri-
and recent species assigned to this genus. Attempts latcral ................. . A . recurvirnstrrim (Hock)
to organize this congery into natural groups have 14,. Carina witti apical urnbo, inframrtlian latus 11igt1cr
been made (cf. Pilshry, 1907b; Bloch. 1024; Withers. than wide ....................................... 15
1 0 5 3 ) , but by and large thrse do not embrace thc. 15. Inframcdiatt latrrs hipltcr than adjoining margins of
latrra ....................... . A . africarlum (Iloek)
plethora of available species and have otherwise riot
15. lr~frarnc-tlianIatus same 11rigl11a s adjoining n~arginsof
proved weful. latera ........................................... 16
In the key presented below. only those species pre- 16. Scuturn lrss than 55 I~rightof capitulum ............
viously repolted or collected by the Eltanin from ............................ . A . aciculnrunt sp. nov.
south of the Antarc.tic Convergeric~eare included. 16. Srutuin more than 54 hripht of capitulum ......... . I 7
17. Roof ol' caritra rouritled ...... . A . trialrgulnre (Hock)
17. Roof of carina flat .......... . A . nrrtarcticum (Hock)
I . IT~nhoof cariiial latus apical ....................... 2 18. Height of scutum morc tlian twice the width.. ......
1. limbo of carinal latus nledial to basal ............. 5 ............................. . A . bouvieri (Gruvcl)
2. Inframedian latus wider than high .................. 18. lleight of scutum less than twice the width ........
...................... A. liberrim (Nilsson-Cantcll) ...............................A. weltrteri (Gruvcl)
2. Iiiframcdian l a ~ u shigher than wide ................ 3
3. Inframediarr latus projccting from capitular surface Arcoscalpellum acicularum sp. nov.
...................... A. compactum (Borradaile) Plate VlIl C; Text-figs. 13, 14
3. Jrifrarnedian latus flush wilt1 cal~itular surface ...... 4
Diagnosis: Rostrum small, acicular ; upper latus
4. Caudal appendage mnltiarticulate . . . A . darwinii (Hoek)
4. Caudal appendage uniarticulate . . . . . . . . . . . . . . . . . .
about T3 as wide as high; height of carinal latus
............... . A . mngrraecnrinae (Nilsson-Cantell) more than q3 height of capitulum; mandible with 2
5. Carinal latus horn-shaped.. . .A. multicostatum sp. nov. or 3 teeth excluding inferior angle; setation of inter-
5. Carinal latus otherwise ............................ 6 mediate articles of cirrus VI ctenopod, with 5 pairs
6. Inframedian latus 8-10 times higher than wide.. .... of setae; dwarf males sack-like, spinose, and lacking
..............................A. latusculum sp. nov. calcareous plates.
6. Inframedian latus lcss than 4 times higher than wide.. 7
7. Inframedian and carinal latera projecting ......... Materiul: Eltanin, Sta. 112, type-locality, Drake Pas-
.....................A. angulare (Nilsson-Cantcll) sage, south of Falkland Island (56"02'S, 61°56'W),
7. Inframvdian and carinal latera not projecting.. . . . . . . 8 4006 meters, 1 9 specimens.
8. Upper latus about 54 and inframedian latus ?/lo height
Depository: USNM Cat. No. 125242 (holotype ) ;
of capitulum ....................................
........................... . A . ventricosum (T3oek) 125243 (paratypes).
8. TJpper latus less than ?h and ilifrarnedian latus morc Description: Female-Capitulum globose; elongate-
than Xo height of capitulum ...................... 9 oval; consisting of 1 4 fully calcified, approximate
9. Inframedian latus widcr t l ~ a nhigh, and highrr than
plates covered with hirsute membrane; occludent and
margins of adjoining latera ......................
....................... .A. bouveti (Nilsson-Cantcll) carinal margins convex; plates marked by u~iequally
9. Inframedian latus not wider than high, and not higher spaced, prominent growth ridges, and very low, nar-
than adjoining margins of latera ............... .10 row ridges emanating from region of umbo. Tergum
10. Carinal latera interdigitating; iriframcdian latus same triangular, acuminate at extremities; basal and oc-
height a s margins of adjoining latcra .............. cludent margins straight to slightly convex; about T3
..............................A. formosum (Hock)
length of straight carinal margin: umbo apical.
10. Carinal latera interdigitaiing; itiframetlian latus about
licight of margins crl' athoining latera . . . . . . . . . . Scutum subquadrate; occludent and lateral margins
............................... . A . vitreuln (Hoek) convex; basal and tergal margins y2 length of occlu-
10. Carinal latera not interdigitating . . . . . . . . . . . . . . . . . ..ll dent margin; immediately below juncture of tergal
11. Lower portion of capitulunl swollen and protruding and lateral margins plate is hollowed out; umbo
laterally ....................... . A . gaussi (C;ruvel) apical; apex of plate extending over basal margin of
11. Lower portion of capilulum flat, not protruding.. .. . I 2
tergum; internally there is one large and one small
12. Jnframcdian latus hourglass-shal~cd, nrnho snhapical
............................... . A . bernrlti (Gruvel) depression for males; major depression situated about
12. Illframedian latus kcyholc-shapetl, umbo suhapical.. . l 8 the length of the occludent margin from the apex;
12. Inframcdian lains triangular, umho apical.. ....... .13 small depression situated on apicolateral border of

Fig. 13. Arcoscdpelhn~acicularum sp. nov.: A, B, side view of females; C, external view of carinal latus;
D, external view of upper latus; E, internal view of sc:utun~;F, view of ahutment of carinal latera and basal
portion of carintl (note hole bored in right carinal latus) ; G, external view of inframedian latus; H I, ex-
ternal view of rostru1n; .I, apex of carina; K, extcrnal view of a peduncular scale; Id, labrum and palps;
M-0, selcctetl portions of crest of labrum showing form of teeth; P, intermediate article of left cirrus
VI; Q-S, caudal appendage, Eltanin, Sta. 112 (holotype, figs. A, F, G, L-P, K, S ; paratypes, figs. R-E,
H-K, Q ) .
major depression. Carina bowed; simple; roof evenly of slightly concave scutal margin; umbo apical; 2
rounded; crest with single broad ridge; lateral mar- major ribs emanate from umbo; one extends curvi-
gins of roof lacking ribs; umbo apical and set off linearly to basal margin parallel to scutal margin;
from main body of plate by slight stricture. Upper the other is situated a short distance from and parallel
latus 4-sided, but essentially triangular (fig. 13D) ; to tergal margin. Carinal latus largest of latera; 5-
tergal and basal margins straight; about T$ length sided; subpentagonal; umbo on upper carinal margin

SYSTEMATIC ACCOUNT $5
about of length of plate from basal margin; um- margins nearly of equal length; basal margin about
bonal region somewhat elevated above general level :% length of occludent; umbo at upper rostral angle.
of plate; plate extending around and below basal Rostrum small, narrow, elongate-oval, width of plate
margin of carina; no interdigitations where plates about !/I height; partially hidden by rostral margins
meet (fig. 13F). Inframedian latus triangular, nar- of rostral latera; umbo apical. Peduncle about $$
row; basal margin less than l/iL length of lateral mar- height of capitulum; 7-12 rows of imbricating pedun-
gins; plate slightly higher than rostral latus and cular scales; 5-7 scales per row; scales extremely
higher than adjoining margin of carinal latus; umbo broad and low; superior margins evenly rounded
apical. Rostra1 latus quadrilateral; scutal and lateral or flat; exposed portions with growth lines only.
lACEGIKMNOP 0 25 rnrn
BDFHJL 0.5
Fig. 14. Arcoscnlpellum ncicularun~sp. nov.: A, C, E, G, I, K, mandibles; B, D, F, H, J, L, enlarged

view of corresponding inferior angles of mandibles; M-0, maxilla I ; P, maxilla 11. Eltanin, Sta. 112
(holotype, figs. A-D, M, P ; paratypes, figs. E-L, N, 0 ) .

46 ANTARCTIC CIRNIl'EDIA
Me~suremcnts (in mm J of 10 sl~ecimensare as fol- for the number of segments in the anterior ( a ) and
lows: posterior ( p ) rami, n = 6 throughout:
Overall Capitular Capitular

Height lleight Width
R- 6.7-18.1 5.1-13.5 2.7-6.6
X 11.5 8.8 4.3
Labrum hullate; fine, short bristles scattered along

periphery; superior surface lree of bristles; crest
armed mith 50-50 terth: crests of teeth hifid, trifid or
multi-denticulate (figs. 13M-0). Palps dongate,
hroad, rounded distally; p ~ o x i m a l superior margin Caudal appendage of a single article; short; ranging
bears short stiff bristles; distal, and basal margins florn to '15 height of hasal segrnent of protopo-
and outer surface of palp bear long setae. Mandible dite of cirrus V1; shape vaiying from short and glo-
nith 2 or 2 terth (lrgs. 14A-L) : second tooth well hose to tall and slender; distal portion with 4-6
separated from first; third tooth, when present, proxi- short, stiff setae; length of terminal setae about
mal to inferior angle; inferior angle strongly denticu- length of appendage.
late, 5-20 teeth either subspatulate or triangular, Tlwarf males sack-like, elongate oval (0.7 x 1.15
commonly somr teeth are hilid. Maxilla I with slight m m ) , laclting calcareous plates, but covered with
to strong notch above midpoint of cutting edge (I~gs. coriccntric bands of short spines. First antennae only
11M-0) ; 2 lorig stout and 2-3 shorter, stout spines appendages visible.
above notch; 0-1 1 short to moderately long spines Remark,: This species is related to A. dubium
helow notch. Maxilla TI triangular in outline; 3 (Hork, 18::3), dredged by the Challenger from 2560
distinct clusters of marginal sctae present. Maxillary meters off the eastern side of Cape York Peninsula,
lohe moderately long. hroad, erect, cylindrical, and north Queensland, Australia (12°08'S, 145010rE), at
apically truncate. the northwest edge of the Coral Sea. Although Hoek
Cirrus I sepa~atedfrom posterior cirri; rami ahout reported the presence of only 1:3 capitular plates,
'"
.; length of rami of cirrus 11; anterior ramus slight- which may prove to he the most diagnostic character-
ly shorter than poste~iorramus; intermediate article- istic separating these t ~ species,
o no critical compari-
of anterior rarnus protuberant; posterior ramus about son car1 be made until A. dubiz~rnis ~cdescribed.How-
' 2 width of anterior. Cirrus I1 not modified; ap- rvel, A. acicularrrna ma) he separated from A. d~ebiuin
proximating cirri 111-V1. Cirri 11-VI essentislly equal l)y tlie following characters: the roof or the carina is
in length, with rqnal or rrearly equal rami; each artic - evenly rounded, with but a single rib extending the
ulation alorig grrater curvature of intermediate arti- lvngth of the crrst, rather than being llat and lacking
cles supporting 2-4 long setae and 1-3 extremrlv a (.lest: the height of the upper latus is less than twice
short. still 111istlc5 : intel artit ular areas along greater the ~ i d t hrathcr than more than twice the width;
curvature tlrvoid of srtac: setation of intermeciiatcl tlre caririal latus is more than 5;; the height of the
ctenopod; :l-l pairs o f setae with I or 2 short ?etCica capitulum, rather than less than 1Li; the rostra1 latus
at base of carh major pair; inner lateral facrs of is slightly wider than high rather than more than
proximal at tic lvs n ith I anrlomly arrayed setac dr- t n i t e as wide as high. In addition, in A. duhiurn
creasing in r1um1tc.r frorn cirrus IT to cirrus V. Cirral thvtc. appear to he fencr ~)oduncular scales in each
counts for (!I(. liolot\ pe .Ire as follows: r012. and the rocts are i~legularl) arranged. As for
the males, which Hoek examined, it need only be
poinlrd out that in A. o!ril~zuna the\ lack ?mall spincs
and apparently reach a sotnewh~t1argc.r size (length
1.;; m n l ) .
Additional factors that probably seive to separate
the two species are geographic distribution ant1 hathy-
metry. Altllough both specics are known only from
Data on the cirri of tlie h o l o t ~ p rarrd two paratypes the type-1oc.alities. A. ncscr11airrrr~occurs nearly 1500
are summarized below. Range ( R ) , mean values iX 1 , meters tierper than A. dubiurn.

The specific name, derived from the Latin, acicula, Depository: USKM Cat. No. 125244.
points out the unique shape of the rostrum. Supplementary Description: Female-The capitular
Arcoscalpellum africanum (Hoek, 1883) plates of this species were described by Hoek and
Plate VIII D, Text-fig. 15 Nilsson-Cantell (1930b, p. 242), and nothing further
need be added at this time. Measurements of the
Scalpellum ajricanurn Hoek, 1883, p. 87, pl. 6. single specimen are as follows: total height 8.7 mm,
fig. 14; Gruvel, 1905, p. 62: Nilsson-Cantell, 1930b, capitular height 7.5 mm, capitular width 3.8 mm.
p. 242, figs. 8a-j; 1931, p. 7. Labrum bullate; short, broad, gently rounded
Diagnosis: Rostrum large, ovotriangular; peduncle distally: superior surface and margins sparsely cover-
about scutum about I$$, and carina about %; ed with short, stiff bristles; crest armed with 45-50
height of capitulum; mandible with 3 or 4 teeth ex- broad, triangular teeth (fig. 15A). Palp long, slender,
cluding inferior angle; maxilla I with deep medial terminally acute; superior proximal margin clothed
notch; anterior ramus of cirrus 1 shorter than pos- with short, stiff bristles; inferior margin free of both
terior ramus; sack-like dwarf male lacking calcareous bristles and setae; distal extremity and lateral face
plates. sparsely clothed with long and short, stout setae.
Material: Eltanin, Sta. 289, off Adelaide Island, Mandible with 4 teeth including inferior angle; first
Antarctic Peninsula (65 5 3 5 , 70" 56'W) , 2782-2912 and second teeth widely spaced; third tooth equidis-
meters, 1 specimen. tantly spaced between second and inferior angle; in-
Fig. 15. Arcoscalpellum africanum (Hoek) : A, labrum and right palp; B, left palp; C, mandible;
D, enlarged view of second and third teeth and inferior angle of mandible; E, maxilla I ; F, maxilla
11; G, cirrus I ; H, intermediate article of cirrus VI; I, caudal appendage. Eltanin, Sta. 289.

ferior angle serrate; 7-12 moderately long, triangular Apparently this species enjoys a wider bathymetric
teeth present (fig. 15C, D ) . Maxilla I with deep range than most deep-water scalpellids.
medial notch; 2 long and 2 short stout spines above
Arcoscalpellum angulare (Nilsson-Cantell, 1930)
notch; 7 stout spines below notch; longest spine in
Plate IX G, Text-,fig. 1 6
lower cluster as long as long spines in the upper
cluster (fig. 15E). Maxilla I1 ovotriangular; marginal Scalpellurn angulare Nilsson-Cantell, 1930b, p. 239,
setae segregated into 3 clusters; setae moderately long fig. 7 ; Nilsson-Cantell, 1939, p. 228.
and covered with setules. Maxillary lobe low, broad; Diagnosis: Plates with few widely spaced growth
apex broadly rounded; surface free of setae or ridges, densely clothed with long bristles; rostrum
bristles. large, broadly triangular; inframedian latus triangu-
Rami of cirrus I subequal (fig. 15G) ; intermediate lar, umbo subapical, elevated above general surface of
articles of anterior ramus protuberant; intermediate capitulum, and when viewed from ventral or dorsal
articles of posterior ramus about y,width of those edge appears obtusely flexed; cutting edge of maxilla
of anterior ramus; both rami clothed with long setae. I with 10-12 spines; crest of labrum lacking teeth,
Rami of cirrus 1 about y2 length of rami of cirrus 11. anterior surface covered with ctenoid scales.
Cirrus I1 normal. Cirri IV-V1 essentially equal in
Material: Eltanin, Sta. 410, between Elephant Island
length; rami equal; each articulation along greater
and Aspland Island, South Shetland Islands (61°18'S,
curvature of intermediate articles with 1 long and 1
56'09'W), 220-240 meters, 17 specimens; Sta. 439,
short seta and 1-2 very short bristles; interarticular
north of Brabant Island, Palmer Archipelago (63
areas along greater curvature and lateral faces free
51'S, 62"3SrW), 12G-165 meters, 16 specimens.
of setae; setation ctenopod (fig. 15H) ; 3 major and
1 minor pair, ~ i t h1-2 short, slender seta at base of Depository: USNM Cat. No. 125245 (Sta. 410) ;
each major pair. 125246 (Sta. 439).
Caudal appendage uniarticulate; about $5 or less Supplerner~tary description: Female - Labrum bul-
the height of the first segment of the pedicel of cirrus late; long, narrow, acutely rounded distally; crest of
VJ; appendage laterally compressed; anterior and labrum free of teeth; superior surface and margins
posterior borders free of setae and bristles; apex covered with ctenoid scales except for a narrow zone
bears 4 5 short setae, none of which equals length immediately anterior to the crest; region below crest
of appendage (fig. 111). covered with several rows of comb-like teeth (fig.
1 6 H ) . Outer surface of palps free of ctenoid scales;
superior margins lacking setae or bristles; inferior
proximal margin armed with only short, stiff bristles.
Mandible (fig. 16L, Q) with 3 long, slender, more or
less equidistantly spaced teeth; inferior angle coarsely
pectinate, 3-7 teeth. Maxilla I with medial, or medial
and sub-basal notch in cutting edge (fig. 16N, P, R ) ;
One male cyprid was found in pouch in each 2 long, stout or 1 long, stout and 3 shorter, stout spines
scutum; each measured 0.65 X 1.30 mm. above medial notch; 5-6 moderately long, slender
spines below notch followed by 2-3 very short, thin
Remarks: This species was first collected by the
spines which may or may not he separated by a notch.
Challenger off Nightingale Island, Tristan da Curiha
Maxilla I1 ovate; marginal setae not clearly segregated
Group (37'25'S, 12"30fW) at a depth of 183 meters, into clusters; inner lateral face with a few clusters
and subsequently was taken by the Discovery from of ctenoid scales. Maxillary lobe very low and broad.
the same region at a depth of 80-140 meters (Hoek, Anterior ramus of cirrus I slightly shorter than
1983, p. 88; Nilsson-Cantell, 1930b, p. 242). Its posterior ramus; intermediate articles of anterior
occurrence at 2782-2912 meters off Adelaide Island ramus slightly produced; intermediate articles of
greatly extends not only its geographic range but its posterior ramus about y, width of articles of anterior
bathymetric range as well. The agreement in cirral ramus. Cirrus 11 slightly shorter than cirrus 111;
counts, and capitular and trophic morphology, leave cirri 111-VI essentially equal in length, rami subequal.
little doubt that the present specimen is conspecific Articulations along greater curvature of intermediate
with those described by Hoek and Nilsson-Cantell. articles of cirri 111-VI with 1-2 long, slender, setulose

SYSTEMATIC ACCOUNT
Fig. 16. Arcoscalpellun~nngulare (Nilsson-Cantell) : A-F, side view of females; G , K, side and front views
of inframedian latus; H, labrum; I, M, side and top views of cyprid, respectively; J, external view of
rostrum and adjoining plates; L, Q, mandibles; N, P, R, maxillae I; 0,maxilla 11. Eltanin, Sta. 439 and
Sta. 410 (figs. F, J only).

setae and 1-2 extremely short, stiff bristles; inter- Arcoscalpellum antarcticum (Hoek, 1883)
articular areas and inner lateral faces free of setae; Text-fig. 17
setation ctenopod, 2 major and 1 minor setulose pair,
Scalpellum antarcticum Hoek, 1883, p. 95, pl. 4,,
1-3 very short, slender setae at base.
figs. 16-17.
Specimens in the present collections range in overall
height from 0.9 to 7.7 mm. Measurements of the Diagnosis: "Capitulum covered by membrane, f w -
two species from Station 410 are as follows: total nished with numerous though very small spines.
height 7.7 mm, capitular height 5.2 mm, capitular Valves thirteen. Carina simply bowed, with the umbo
width 3.3 mm (specimen 1 ) ; total height 7.3 mm, at the apex and with a flat roof. Upper latus quad-
capitular height 5.0 mm, capitular width 3.5 mm rangular. Umbo of the carinal latus almost at the
(specimen 2 ) . Cirral counts follow: base of the valve. Peduncle cylindrical, narrower than
the base of the capitulum" (Hoek, 1883, p. 95).
Material: Challenger, Sta. 153, type-locality, north of
Specimen 1 6 9 11 11 12 12 1 Ingrid Christensen Coast (65042'S7 70°49'E), 3062
- - - - - -
7 10 12 12 12 12
meters, one specimen.
Original Description: "This is the most southern
species of the genus taken by the Challenger. It is a
medium-sized species, and it is represented by a single
Specimen 2 5 9 10 10 10 11 1 specimen only.
- - - - - -
6 10 11 11 11 12 "The capitulum is rather robust, convex. It con-
tains thirteen valves; there is a trace of a fourteenth
(the rostrum), but it is too rudimentary to be noticed.
The valves are covered by a brown chitinous mem-
Cyprids in mantle cavity, about 50 to 60 in each brane, which bears numerous very small spines. Be-
of 3 specimens, measured 0.25 X 0.75 mm; each had tween the valves there are very narrow chitinous
paired, long, slender, appendage-like structures poster- interspaces.
ior to antennae 1, the distal ends of which bear ex- "The scutum is quadrangular, with the umbo at
tremely short and few bristles (fig. 161). the apex. The whole valve is convex, especially in the
Dwarf males sack-like; ranging from 0.30 X 0.55
mm to 0.35 x 0.67 mm; lacking cirri and calcareous
plates; antennae present; surface lacking concentric
bands of short spines.
Remarks: The single most diagnostic feature of the
present species is the strongly flexed inframedian
latus, the umbo of which is greatly elevated from the
surface of the capitulum (fig. 16G). However, the
characteristic development of this plate does not ap-
pear until after the capitulum reaches a height of
about 1 mm.
The specimens described by Nilsson-Cantell bere
dredged near Clarence Island, the northeasternmost
of the South Shetland Islands. The present specimens
were recovered from the southernmost islands in this
chain. The Eltanin specimens occurred at slightly
shallower depths (128-240 meters) than those taken
by the Discovery (342 meters). No information is
available insofar as substrate preferences of the col-
lections here discussed. Nilsson-Cantell's material oc-
Fig. 17. Arcoscalpellum antarcticum (Hoek) : A,
curred on a tunicate and a pycnogonid (Colossendeis view from left side; B, carinal view (redrawn from
sp.). Hoek, 1883, pl. 4, figs. 17-18),

SYSTEMATIC ACCOUNT
uridermost part. 'The occluclent margin is arched. the

latrral margin also; the basal margin is slightly
arched, almost straight; the tergal margin, finally, is
hollowed out.
"The tergum is triangular, of a greater area than
the scutum, with the apex recurved. The occludent
margin is slightly arched.
"The carina is rather stout. Its umbo is at the apex,
arid the whole valve is simply bowed. It has a flattened
roof, which does not increase very much in width
from the upper to the lower extremity. The sides of
the cariria are at more than right angles with the
roof; they increase much in width from the lower
to the upper extremity.
"The upper l a ~ u sis quadrangular, with the lower
angle truncated, hence pcntagonal. The umbo is at
the apex, which is slightly prominent. The scutal
margin is hollowed out, the tergal margin slightly
arched. Fig. 18. Arcoscalpellum herndli (Gruvel) : A, view
"The rostral latr~s is of considerable size, quad- from right side; R, rostral view (redrawn from
rangular. Scutal and basal margins nearly parallel; Gruvel, 1909, pl. 23, figs. 6-7).
the whole valve is distinctly convex and divided into
two triangular parts by a not very prominent ridge Diagnosis: "Capitulum laterally compressed, of 1 4
running from the umbo to the basal lateral angle. completely calcified, approximate plates covered with
"The infra-median lalus is small, triangular. The a very thin smooth cuticle. Carina regularly curved,
umbo is at the slightly protuberant apex. of scarcely half the height of the entire capitulum.
"The carinal l a ~ u shas an irregular shape, resembl- Dorsal margin convex, without lateral edges, but with
ing that of a shoe. The umbo is at the carinal basal a very weak central dorsal ridge that terminates be-
angle and represents the heel of the shoe. The carinal low with a sharp angle. Tcrga triangular, elongate,
margin (the sole of the shoe) is slightly hollowed out. with sharp, slightly forward bent apex.
The upper margin is straight, the lateral margin is "Scuta quadrangular, elongate; the umbones situ-
very short, the basal margin straight, only slightly ated at the apices which are sharply and scarcely
upturned towards the umbo. reflexed. The upper latera are elongate, triangular
"Length of the capitulum, 20 mm. and bear their umbones very near to the apices. The
"The peduncle is short (5.5 m m ) , cylindrical, umbones of the carinolateral plates lie on the apices
much narrower than the base of the capitulum. and are somewhat recurved. The anterior margin of
'6'
1 he scales are covered by membrane, very narrow,
7
the rostro-lateral is straight and bends with the unibo

transverse, calcareous stripes only being visible. They slightly forward a little.
are not numerous, and do not form very distinct rows; "Infra-median latus elongate, rectangular with slight
they are at considerable distances from one another. central constriction.
"This species may be easily distinguished from the "Rosirum elongate, pentagonal, completely free.
nearly related Scalpellum velutinum, Scalpellum ped- "Peduncle narrow, moderately long, almost cylin-
unculatum, etc., by the form of the carinal latus. drical, with eight pairs of small irregular scales, which
Though this valve also may perhaps present variations par~iallycover one another.
within the limits of a species, in the present case it "Locality: Winter station of the GAUSS, north of
can be safely made use of" (I-loek, 1353, pp. 95-96). Kaiser-Wilhelm 11 Land, at 380 m depth. Only one
example found.
Arcoscalpellum berndti (Gruvel, 1907)
"Relationship: By the form and the shortness of the
Text-fig. 18
carina, this species approaches Scalpellum brevecal-i-
Scalpellurn berndti Gruvel, 1907b, p. 160; Gruvel, r~aturnHoek, Sc. sessilc IIoek and Sc. pentacrinarr~rn
1909, p. 207, pl. 23, figs. 6-7. Pilsbry, however, iL can he united with none of these

because of the form of their carinolateral plates and margin concave. Rostrum triangular, elongated, umbo
particularly the infra-median latera, the umbones in- apical. Rostra1 latus quaclrangular, a little wider than
deed being on the apex, which have however a more high. Inframedian latus triangular, umbo apical, the
or lcss distinct triangular form" (Gruvel, 1909, pp. margins of the same length. Carinal latus pentagonal
207-2081. with the umbo at the middle of the carinal margin.
Peduncle with transversely elongated scales, cuticle
Arcoscalpellum. bouveti (Nilsson-Cantell, 1939) hairy. Mandil~le with three teeth and a pec~inated
Text-fig. 10 inner angle. Maxilla 11 \+ith a concave front edge
S c a l p e l l ~ ~ mbouveti Nilsson-Cantell, 1930, p. 233, with bristles and a posterior lohe with bristles. A
fig. 4. distinct maxillary lobe is formed. Caudal appendage
Diagnosis: "Female. Capitulum with fourteen valves single-jointed without bristles. of about the same
separated by distinct chitinous interspaces. Growth length as thc proximal segment of the protopodite.
lines weakly marked. Capitulum covcred by a thin "Complemental male sack-like, without cirri arid
cuticle. Scutum quadrangular, occludent margin slight- ~ d v e s " (Nilsson-Cantell, 1939, p. 233) .
ly convex. Tergum triangular, occludent margin con- Materictl: Discovery, Sta. 456, type-locality, east of
vex. Carina regularly bent, unlbo apical, dorsal roof Eouvet Island (approx. 5b0S, 3OE1, 40-45 meters,
convex. Upper latus pentagonal, umbo apical, scutal 1 specimen.
Fig. 19. Arco~ca/~~rllrcnr boui~rti (Nilsson-Cantcll) : A, view from right side; R,

rostruln and rostrolatcrals; C, I-section of carina; L), palp; E, mantlil)lc; 12, second
maxilla; 6, pedicel of cirrus V I and caudal apprndagc (retlrawn frorn Nilsson-
Canlell, 1939, fig. 4,).

Original Description: "Female. Capitulum covered mature. As earlier stated (Nilsson-Cantell, 1921)
by a thin cuticle. Valves fourteen in number, separated several Scalpellurn species from Antarctic and sub-
by distinct chitinous interspaces. The growth lines in Antarctic seas as for instance S . gibherum, convexurn,
this species ~ ~ e a k lmarked.
y compacturn and ventricosum, like this new species,
"Scutum quadrangular with somewhat convex oc- have the first development until metanauplius or cypris
cludent margin. stage in the mantle cavity.
"Tergum triangular with convex occludent margin "One Scalpellurn specimen I cannot refer to any
and recurved apex. previously described species from corresponding parts
"Carina regularly bent with the umbo at the apex. of the ocean. I think it convenient to erect a new
Dorsal roof convex. Sides well developed with wide species hoping it will be possible later on to refound
areas in the upper part. it from richer material. It shows some resemblance
"Upper latus pentagonal with the scutal margin to younger specimens of S . convexum, represented in
concave and the tergal margin convex. The carinal this collection, but is different especially in the in-
margin and the margin against the inframedian latus framedian latus and carina" (Nilsson-Cantell, 1939,
short. The umbo is apical. pp. 233-235).
"Rostrum triangular, elongated with the umbo at
Arcoscalpellum bouvieri (Gruvel, 1906)
the apex.
Text-fig. 20
"Rostra1 latus quadrangular with short basal mar-
gin. Width of plate exceeds height. Scalpellurn bouvieri Gruvel, 1906, p. 272 ; 1907a,
"Inframedian latus triangular, with margins of the p. 3, figs. 7-9; Nilsson-Cantell, 1926, p. 2, fig. 1 ;
same length. The umbo is apical. Zevina, 1964, p. 252; 1968, p. 92, fig. 5.
"Carinal latus pentagonal with the umbo at the Diagnosis: "Capitulum with 14 plates, well calcified,
middle of the carinal margin. Lateral upper and basal with small chitinous interspaces. Growth lines weakly
margins nearly of the same length. indicated. Scutum rectangular, with, as a rule straight
"Peduncle of the same length as the capitulum, occludent and lateral margins. Umbo at the apex.
with transversely elongated scales separated by chitin- Carina regularly bent, with the umbo in the vicinity
ous interspaces. The cuticle a little hairy. of the apex. Lateral ridge distinct, but weakly indi-
"Measurements (in mm) of holotype: length of cated. Upper latus pentagonal, umbo a little below
capitulum 6, breadth of capitulum 4, length of ped- the apex. Rostrum elongated, umbo at the apex.
uncle 6, breadth of peduncle 2. Rostro-latus quadrangular, the height equal to the
"Mouth parts. Palpus conical with bristles at the width. Infrarnedian latus nearly hourglass shaped,
point. Mandible with three teeth and a pectinated with the umbo at the middle or a little below the apex.
inner angle. Maxilla I could not be studied in the Carino-lateral high, with the umbo near the base of
holotype. Maxilla I1 with bristles along the whole the carina. Peduncle with small, sparsely situated
concave front edge. A posterior lobe with bristles is plates. Mandible with three teeth and a pectinate
differentiated like a maxillary lobe. lower angle. Maxilla I with a distinct notch in the
middle of the front margin. Maxilla I1 with bristles
in three groups. Caudal appendage small, one seg-
Holotype 6 12 12 12 12 14 1
- - - - - - mented, conical without bristles" (Nilsson-Cantell,
8 12 12 13 13 14
1926, p. 4 ) .
"Cirrus I shorter than the other cirri, rami un-
Arcoscalpellum brevecarinatum (Hoek, 1883)
equal in length with two segments.
Text-fig. 21
"Cirrus IV with six pairs of spines on the front
edge of the segments. The caudal appendage is single- Scalpellurn brevecarinatum Hoek, 1883, p. 82, pl. 3,
jointed and pointed without bristles. The appendage fig. 22; Zevina, 1964, p. 253.
is about the same length as the proximal segment of Diagnosis: "Valves fourteen, covered by a very thin
cirrus VI. membrane only. Carina very short, simply bowed,
"The complemental male [dwarf male] is of the with the umbo at the apex. Upper latus quadrangular.
reduced sack-like type. Valves of the lower whorl, and especially the carinal
"In the capitulum a number of cypris stages were latera, large. Rostrum wedge-shaped" (Hoek, 1883,
found, indicating that the specimen dissected was p. 8 2 ) .

Fig. 20. Arcoscalpellum bouvieri (Gruvel) : A, view from riglit side of

entire individual, carina alone, and rostral side; B, inframedian latus from
another individual; C, palp; D, mandible; E, first maxilla; F, second
maxilla; G, ~ e d i c e lof cirrus VI and caudal appendage (redrawn f r o m
Nilsson-Cantell, 1926, fig. 1, A-13).
Material: Challenger, Sta. 146, Atlantic-Indian Ridge, margin is arched, the lateral margin also; the tergal
between Prince Edward and Crozet Islands (46"46'S, margin behind the projecting triangular part is
45"31fE), 2514 meters, 2 specimens; Sta. 147, type- straight.
locality, Atlantic-Indian Ridge, between Prince Ed- "The tergum is triangular; its occludent margin
ward arid Crozet Islands (46"16'S, 4g027'E), 2925 arched; the apex is recurved, the carinal margin
meters, 3 specimens. slightly arched, the scutal margin almost straight. It
Original Description: "Of this curious little deep sea surpasses the scutum in area.
species t ~ large
o and three very small arid young "The carina is short, simply bowed. The umbo is
specimens were collected. They agree M ith Scalpellum at the apex. The roof is not quite flat, and slightly
balanoides in the shortness of the carina. furrowed longitudinally. In the upperinost portion
"The capitulum is flat, and consists of fourteen small parts which must be considered as sides are
valves covered by a very thin chitinous membrane; in visible.
consequence of this the white colour of the valves has "The rostrum is elongate, narrow. The umbo is at
become slightly yellowish. The high development of the upper extremity, which is slightly narrower than
the valves of the lower whorl is especially characteris- the base.
tic of this species. "The rostral latus is trapeziform. The scutal margin
"The scutum is trapeziform; about twice as long as is much longer than the basal margin. The latter is
broad. The apex is slightly recurved, and projects over straight, the former slightly hollowed out.
the tergum as a small triangular part. The occludent "The infra-median latus has about the same shape as

the rostrum. It is slightly protuberant over the surface most anterior or;e is shorter and very thick, the pos-
of the other latera. Its umbo is at the upper extremity. terior one is much more elongate; the latter has seven,
C<
The cnrinal latus is very large and of an elongate the former five segments. A distinct interval separates
pentagonal shape. The lateral margin is long and the first cirrus from the five posterior pairs, which
straight, the basal margin is rather short; the carinal are much more elongate.
margin is divided into a superior portion which is "The caudal appendages are uniarticulate" (Hoek,
hollowed out, and into a rather long and slightly con- 1883, pp. 82-83).
vex inferior portion; the upper margin is straight. The
umbo projects slightly over the carina. The two cari- Arcoscalpellum buccinum sp. nov.
nal margins touch one another over a long distance Plate VIlI I ; Text-figs. 22, 23
beneath the carina. Diagnosis: Capitular plates ornamented with strong
"Length of the capitulum, 7 mm. radial ridges; rostrum small, partially hidden by ros-
"Peduncle short, conical. Scales large, calcareous; tral latera, essentially triangular in outline when viewed
the edges very prominent. They are not numerous, and externally, but in side view shaped like a horn with the
are placed in horizontal rows; vertically no distinct umbo directed inward toward the mantle cavity; ad-
rows can be made out. joining margins of carinal latera interdigitating be-
"The labrum is bullate; its free edge is furnished low carina; inframedian latus triangular, and shorter
with a row of small but distinct and pointed teeth; than occluding margins of latera; mandible with 4
the palpi are not very elongate, rather robust, furnished teeth including inferior angle, margin of third tooth
with hairs not only at the extremity but also at the serrate; maxilla I with deep notch in medial portion of
sides. The mandibles have three teeth and a very short cutting edge; maxilla I1 triangular, but not trilobate;
inferior angle, which is strongly pectinated. The maxil- anterior ramus of cirrus I shorter than posterior
lae have the edge nearly straight; only a trace of a ramus; caudal appendage uniarticulate, clothed with
notch behind the very broad upper spines. The outer 14-22 setae; dwarf males sack-like, lacking calcareous
maxillae have the bristles not divided into separate plates.
tufts. The so-called olfactory orifices [maxillary gland Material: Eltanin, Sta. 1429, type-locality, on Camp-
openings] are not placed on highly protuberant excres- bell Plateau, between Auckland and Bounty Islands
cences. (49"15'S, 172"14'E), 320-361 meters, 35 specimens;
"The first pair of cirri has unequal branches. The Sta. 1430, on Campbell Plateau, between Auckland and
Bounty Islands (4g019'S, 171°36'E), 165-192 meters,
1specimen.
Depository: USNM Cat. No. 125247 (holotype; Sta.
1429) ; 125248 (paratypes; Sta. 1429) ; 125249 (para-
types; Sta. 14$30).
Description: Female-Capitulum with calcified, ap-
proximate plates; long and oval; widest in the middle;
both margins convex, the carinal border more so than
the occludent; cuticle thin, sparsely hirsute; plates
sculptured with unequally spaced growth lines, and
low, approximate, radial ridges (pl. VIII, I ) . Tergum
narrow; projecting apically; extremities acuminate;
carinal margin more than twice length of occludent
margin (fig. 22B). Scutum quadrate; higher than
wide, occludent margin convex and commonly more
than twice the width of the basal margin; carinal mar-
gin straight. Carina bowed; extremely broad basally,
but tapering towards the apex; roof flat, bounded by
high, moderately broad ridges; crests of ridges ribbed
Fig. 21. Arcoscalpellum brevecnrinatum ( H o e k ) :
Individual viewed from right side (redrawn from (fig. 23B). Upper latus with 5 sides of unequal length;
Hoek, 1883, pl. 3, fig. 22). umbo apical; scutal margin convex; tergal and basi-

Labrum bullate; no soft setae present; crest armed

with about 50 triangular teeth. Palp sparsely setose;
setae distributed over lateral face, superior border, and
distal extremity, but not on inferior margin. Man-
dible with 4 teeth, including the inferior angle (fig.
2 3 6 5 ) ; first and second teeth well separated; third
tooth and inferior angle more closely spaced; inferior
angle serrate; teeth on upper margin but not lower.
Maxilla I highly spinose; deep medial notch present
(fig. 23K-0) ; number of spines above notch variable;
but commonly 4-5 large, stout, and 1 or 2 smaller,
narrow spines; 9-13 spines below notch. Maxilla I1
essentially triangular in outline; 3 well-segregated
clusters of marginal setae present; setae of the superior
proximal cluster longer than the setae of the other 2
clusters. Maxillary lobe extremely small; tubular;
Fig. 22. A r c o ~ c a l ~ e l l u mbuccinum sp. nov.: A, left rounded apically.
side of malformed female, the upper latus lacking and Cirrus I separated from cirrus 11, and about 3/4 its
the scutum now occupying the position normally filled length. Anterior ramus of cirrus I slightly shorter than
by this plate; B, right side of normal female. Eltanin,
posterior ramus (fig. 2 3 4 ) ; intermediate articles of
Sta. 1429 (holotype, fig. B) .
posterior ramus about width of those of anterior
carinal margins straight and of nearly equal length. ramus. Each articulation along greater curvature of
Carinal latus essentially triangular, but with 5 unequal intermediate articles of cirri 11-VI with 1 long and 1
sides; apex projecting slightly beyond the carinal bor- or 2 shorter setae; interarticular areas along greater
der; umbo at extremity of projection; basal and cari- curvature devoid of setae; setation ctenopod, 3 pairs
nal margins longest; upper and lateral margins shorter, on cirri 111-V, 3-4 pairs on cirrus VI.
subequal and convex; carinal margin concave; plates Cirral counts for the holotype are as follows:
interdigitating below basal margin of carina (fig.
23B). Inframedian latus nearly triangular; margins
of nearly equal length; plate less than height of
lateral margin of rostral latus. Rostral latus essen-
tially quadrangular; occludent and lateral margins
convex; scutal margin concave; plate slightly wider
than high; highest at lateral edge. Rostrum partially
Counts of the cirral segments are summarized below
hidden beneath rostral latera (fig. 23C) ; nearly as
for the holotype and two paratypes. The range ( R )
high as occludent margin of latera; plate nearly as
high as broad; shaped like a horn when viewed from and mean values (x) for the number of segments in
side; umbo apical; directed touards body cavity (fig. the anterior ( a ) and posterior (p) rami are as follows:
23D, E).
Peduncle about qj or less height of capitulum;
armored with closely packed, imbricating, broad
scales; about 4-6 scales per row.
Measurements of the holotype are as follows: total
height 18.9 mm, capitular height 15.3 mm, capitular
width S.9 mm. Measurements of the paratypes from
Station 1429 are summarized below:
Overall Capitular Capitular
Height Height Width
n 31 34 34
6.2-15.8 3.3-9.2 Caudal appendage extremely short, conical, 1 seg-
-R
~ 7.6-20.1
X 14.39 11.65 6.42 ment; anterior margin armed with long setae; distal

SYSTEMATIC ACCOUNT
1:ig. 23. Arcosculpellurrz buctinurn sp. nov.: A, external view of inframcdian l ~ t u s ;B, view of abutment
of carinal latera and basal portion of carina; C, rostrum and adjoining margins of rostra1 latera; D, E,
internal and extcrnal views of rostrum, respectively; F, labrum and right palp; G-J, mandibles; 13, J, third
tooth and inferior anglcs of respective mandibles; K-0, maxillae I ; P, maxilla 11; Q, cirrus I ; R, inter-
mediate articlc of cirrus V I ; S, caudal appendage. Eltnnin, Sta. 1429 (holotype, figs. A-B, F-K, N, P,
R, S only).
extremity with 3 setae of equal length; setae less than there is no upper latus on the left side of the capitu-
lh height of appendage. lum (fig. 22A). However, in this malformed speci-
Dwarf male sack-like, elongate, somewhat constricted men the space normally occupied by the upper latus
medially; consequently, pear-shaped, covered with con- is filled by the scutum. The tergum, also somewhat
centric bands of fine spines; 4 chitinous plates sur- malformed, does not fill any portion of the gap left
round mantle opening; antennae altached above medial open by the scutum. The absence of the upper latus
constriction. is uncommon, although several specimens from differ-
Remarks: In one of the paratypes from Station 1429 ent genera have been examined by the writers.

Arcoscalpellum buccinum is related to the following

species :
A. australicum (Hoek), 1883, p. 118.
A. elegans (Hoek), 1907b, p. 107.
A. elongatum (Hoek) , 1883, p. 93.
A. praeceps (Hoek), 1907b, p. 114.
A. rigidum (Aurivillius) , 1898, p. 194.
A. vitreum (Hoek), 1883, p. 115.
In each of these species the inframedian latus is
shorter than the adjoining margin of the carinal latus,
which is higher than wide, the rostra1 latus is as high
as or higher than wide, and the plates are ornamented
with both growth and radial ridges. The unusual horn-
shaped rostrum of A. buccinum separates it from A.
praeceps in which it is narrow and elongate, A. vitreum
in which it is ovate or acicular, and A. elegans in Fig. 24. Arcoscalpellum compactum (Borradaile) :
which it is just a narrow slip. The tergum of A. aus- A, individual viewed from right side; B, inframedian
tralicum is erect, and y2 as wide as high, and the latus from left side (redrawn from Borradaile, 1916,
fig. 3 ) .
inframedian latus is about y5 the height of the adjoin-
ing margins of the latera, but in A. buccinum this plate
is about the height of the adjoining margins of the rower and more widely separated" (Borradaile, 1916,
latera, and the tergum is inclined and about 1/ as wide p. 130).
as high. In A. elongatum and A . rigidum the carinal Arcoscalpellum darwinii (Hoek, 1883)
latera meet below the carina, but apparently do not Plate IX E, Text-fig. 25
interdigitate as they do in A. buccinum, and the terga
of these two species are erect rather than inclined. Scalpellum darwinii Hoek, 1883, p. 110, text-fig. 3,
The specific epithet is derived from the Latin, pl. 5, figs. 1, 2, pl. 10, figs. 3-5; Weltner, 1897, p. 247;
buccin-, a trumpet, and alludes to the horn-shaped 189Sb, p. 7 ; Gruvel, 1905, p. 78, fig. 87; Pilsbry,
rostrum. 1907b, p. 25.
Scalpellum (Arcoscalpellum) darwini [sic] : Hoek,
Arcoscalpellum compactum (Borradaile, 1916) 1907b, p. 85.
Text-fig. 24 Diagnosis: Capitular plates fully calcified, approxi-
Scalpellum (Arcoscalpellum) compactum Borradaile, mate; carina simply bowed, roof evenly rounded, lat-
1916, p. 130, fig. 3. eral margins ornamented with 5 low, longitudinal ribs
extending from apex to base; rostrum of 4 sides but
Material: Terra Nova, Sta. 356, type-locality, off
essentially triangular, broad apically, tapering toward
Granite Harbor, western entrance to McMurdo Sound
base, umbo slightly subapical; umbo of carinal latus
(approx. 7 7 ' 5 164OE), 92 meters, 1 specimen.
apical; teeth of mandible uniformly spaced; maxilla I
Original Description: "An Arcoscalpellum of about with poorly defined step-like cutting edge armed with
the same size as the supposed young stage of S. dis- 32-38 spines arranged in 3 groups; caudal appendage
coveryi (length of capitulum 5.5 m m ) , but differing with 5 segments. Dwarf males sack-like, lacking cal-
from it as follows: the lateral border of the scutum is careous plates.
not notched; the produced angle of the upper lateral
Distribution: Eltanin, Sta. 803, southern border of
plate is much sharper; he carino-lateral is not notched
Southeast Pacific Basin (65'50'S, 8Z040'W), 4 3 1 G
where it meets the shoulder of the carina; the umbo of
4328 meters, 1 specimen; Sta. 945, off Peter I Island,
the rostrolateral projects beyond the outline of the
Bellingshausen Sea (67'5 5 3 , 90 "43'W) , 4005 meters,
capitulum, but transversely, not with an upward trend,
1specimen.
as in S. nymphonis; the inframedian plate is triangu-
lar with the apex distal (except on one side of one Depository: USNM Cat. No. 125250 (Sta. 803) ;
specimen, where it is very narrow, ~ i t ha spear-head 125251 (Sta. 945).
at the distal end) ; the plates of the peduncle are nar- Supplementary Description: Female-The capitulum

SYSTEMATIC ACCOCJNT 59
has heen fully described by Hoek (1883) ~ i t hthe ex- rounded distally ; proximal superior arid inferior mar-
ception of the rostrum. Rostrum 4-sided, but essen- gins free of setae and bristles; distal extremity and
tially triangular, very narrow, broad apically arid margins densely armed with setulose setae. Mandible
tapering basally, plate about same height as lateral \\ith 4 teeth including inferior angle, teeth uniformly
margins of rostra1 latera and partially hidden by them; spaced, long, slender; inferior angle serrate, teeth
umbo slightly subapical and elevated. 18-22 in number (fig. 25B, C ) . Maxilla I with step-
The specimen from Station 803 is too poorly pre- like cutting edge and no distinct notch (fig. 25D) ;
served to provide meaningful measurements; the speci- spines arranged in 3 poorly defined clusters; apical
men from Station 945 measures as follows: total height cluster with 2 long, stout and 5-7 short, slender spines;
85 mm, capitular height 51 mm, capitular width 34.5 n~iddlecluster with 13-17 short, s l e n d ~ rspines; basal
mm. Both specimens were growing on basaltic cobbles. cluster with 2 long, slender and 13-15 short, sleritler
Labruni bullate; long, broad, obtusely rounded dis- spines; superior and inferior margins and lateral face
tally; surface and margins free of bristles; crest not covered with numerous clusters of bristles. Maxilla I1
armed with teeth. Palp long, broad, triangular, acutely short, broad, and trilobate; marginal setae in 3 dis-
Fig. 25. Arcosculpellum darzvinii (IIoek) : A, palp; B, enlarged view of third tooth and inferior angle of
mandible; C, same mandible; I), maxilla I ; E, maaiila 11; F, caudal appendage; G, intermediate article
of left inner ramus of cirrus VI; I-I, male cyprid; I, antenna I of same cyprid; J, antenna I of dwarf
male; K, same dwarf malc. Eltanin, Sta. 945.

tinct clusters. Maxillary lobe extremely low and broad, covered with concentric bands of long, acicular spines,
apex evenly rounded (fig. 25E), apical opening slit- length of antennae about length of sack.
like. Remarks: This species is known only from the de-
Anterior ramus of cirrus I about %< length of pos- scription of Hoek (1883). The single Challenger speci-
terior ramus, intermediate articles strongly protuber- men was dredged from the Peru-Chile Trench off Val-
ant; intermediate articles of posterior ramus y2 to q< paraiso, Chile (33"31'S, 74"43'W), at a depth of 3950
width of articles of anterior ramus. Cirrus I about meters. The present specimens are from a considerably
y2 length of cirrus 11. Cirrus I1 sligh~ly modified; more southern region, and from slightly deeper water
slightly shorter than cirrus 111. Cirri IV-VI essen- (4000-9328 meters).
tially equal in length with slightly subequal rami; each Arcoscalpellz~rndarwir~iihelongs to the A. michelot-
articulation along greater curvature of intermediate tzarurn group of scalpellomorphans (Pilsbry, 19071,,
articles ~ i t h2-3 long and 1-2 short, slender setae; p. 2 6 ) , which iriclucles the following species:
cirrus I11 with 2-3 long and 3-5 short slender setae. A. michelo~tianum(Seguenza), 1876, p. 3G1.
Interarticular areas of intermediate articles along A. giganteurn (Gruvel), 1902a, p. 153.
greater curvature of cirri 111-VI free of setae; setation A. regium (Thomson), 1873, p. 4.
ctenopod (fig. 25G), 3 major and 1 minor pair; at A. regium latidorsum (Pilsbry), 1907b, p. 29.
and between bases of major s ~ t a ethere are a few long, A. regium ovale (Hoek), 1883, p. 109.
slender setae. Lateral face (inner) of intermediate A. regina (Pilsbry), 1907b, p. 31.
articles of cirri IV-VI with few scattered, long, slender A. gigas (Hoek), 1883, p. 102.
selae. A. tritonis (Hoek), 18G3, p. 122.
Cirral counts for the two specimens collected are as A. alcockianurn ( Annandale) , 1906, p. 392.
follows: A. moluccanum (Hoek) , 1883, p. 10 1.
A. ar~nandalei(Calman), 1918, p. 109.
A. regulus (Calman), 191& p. 113.
Station 803 9 24 23f 34 38 40 5 A. judcli (Calman), 191G, p. 116.
- - -
-
Representatives of this group are c.haracterized by
13 25+ 32 32 35 38
their large overall size. which ctrmmonlj exceeds 75
mm. The capitulum is covered mith a thick, dense coat
of 1,ristles. The iriframedian latus is triangular. and
its height never exceeds that of the caririal latus. The
rostra1 latus is about twice as wide as high, arid corn-
morily the plate is not higher than the adjoining irifra-
median la~us. The carinal latera have an apical uml-10.
and they do not interdigitate here they meet below
the carina. 111 addition, taxa in this g ~ o u pcommonly
Caudal appendage of 1--5 segnlerlts (fig. 25F) ; about occur at depths in excess of 1500 meters.
44-jheight of pedicel of c.ir~usVJ; setae present along
lateral face and anterior margin of each articulation; Arcoscnlpellum formosum (Hoelc, 1907)
terminal segment supports apical cluster of 9-11 long Plate VIII G, Text-fig. 26
setulose setae about same length as distal 2 segments oI S(~all)cllurnformosum Hoek, 1907b, 1). 110, pl. 8,
appendage. figs. 11. I l a ; Nilsson-Cantell, 1921, p. 187, fig. 27;
Ovigerous frena attac,hed near hasilateral angle o f Hiro. 1933, 1). 30, text-fig. 6, pl. 1, figs. 10-lob;
scutum; each ovigerous lamella contained approwi- Nilssori-Cantell. 1938, pp. 7, 16,21.
matel y 150 large eggs (1.5 X 1.0 mm) . nor1 S(.alpc~ll~~m jorrnosun~ Pilsbry, 1007b, p. 58
Dorsal surface of carapace of dwarf-male cyprid (=Scalpellum bellum Pilsbry, 1908, p. 111, replace-
convex; ventral margin straight; posterior margin ment name = Arcoscalpellum vitreurn (Hoeki , 1883).
truncate and normal to ventral margin; anterior por- Scnlpellz~~n (Scalpellurn) forrnosum: Stubbings,
tion acute (fig. 25H) ; carapace of individual figured 1936, p. 26.
from Station 803 measures 1.0 X 2.5 mm. Dwarf Scalpellum (Arcoscalpellum) jormosum: Tarasov
male sack-like (fig. 25K I , elongate-oval (0.75 X 2.13 and Zevina, 1957, p. 140, text-fig. 114, 111. 1, fig. la-c.
mmi , without calcareous plates and cirri; surface Diagnosis: Capitular plates ornamented with weak

radial ridges; carinal margins of carinal latera inter- riorly; margin and superior surface lightly clothed
digitate; inframedian latus extends to and nearly with short, stiff bristles (fig. 26D) ; crest armed with
touches upper latus; rostrum large, rectangular, broad, about 24 teeth. Mandible with 4 teeth including in-
completely exposed, and always same height as ad- ferior angle; superior slope of second and third teeth
joining margins of rostra1 latera; anterior ramus of bear a few small teeth (fig. 26E) ; inferior angle
cirrus I longer than posterior ramus; caudal append- coarsely pectinate, 7-8 teeth. Maxilla I with well-
age uniarticulate, clothed with 12 to 14 setae; man- defined medial notch; 2 long, stout and 2-3 short,
dible with small teeth on upper slope of second and stout spines below notch (fig. 26F) ; commonly lower
third teeth; maxilla I with a medial and commonly a 2-3 spines segregated off by shallow notch. Maxilla I1
suprabasal notch in cutting edge, 5-6 spines below ovotriangular in outline, but not trilobate.
medial notch; maxilla I1 triangular, but not trilobate; Anterior ramus of cirrus I slightly longer than pos-
dwarf male sack-like with 4 calcareous plates. terior ramus; intermediate articles of both rami not
Material: Eltnnin, Sta. 268, west of Renaud Island, significantly protuberant and more or less equal in
Biscoe Island, Antarctic Peninsula (64"01'S, 67' width (fig. 26H) ; both right and left cirrus I1 in pres-
485'W), 2763-2818 meters, 1 specimen; Sta. 480, Scotia ent specimen malformed. Cirri 111-VI essentially
Sea, north of South Orkney Islands (58'065, 44' equal in length and with equal rami; each articulation
56'W), 2800 meters, 1 carina. along greater curvature of intermediate articles with 1
long and 1 shorter, slender seta and a cluster of 4-6
Depository: USNM Cat. No. 125252 (Sta. 268) ; extremely short, stiff bristles; interarticular areas along
125253 (Sta. 480). posterior margins with 1 short, stiff seta; distal end
Supplementary Description: Female-Labrum bul- of each intermediate article on the lateral face, and
late; moderately long, broad, obtusely rounded ante- parallel to the articulation, with row of ctenoid scales;
Fig. 26. Arroscalpellum ,formosum (Hoek) : A, side view of female; B, view of abutment of carinal
latera; C, external view of rostrum and adjoining plates; D, labrum and right palp; E, mandible;
F, maxilla I ; G, maxilla 11; H, cirrus I ; I, intermediate articles of cirrus VI; J, caudal appendage.
Eltanin, Sta. 268.

setation ctenopod, 4 pairs of setae, 3 major and 1

minor (fig. 261).
Caudal appendage of a single segment about
height of first article of pedicel of cirrus VI; sur-
face covered with 12-14 setae of varying lengths (fig.
26J).
Measurements of the single complete specimen are
as follows: total height 8.4, mm, capitular height 6.8
mm, capitular wlidth 3.8 mm. The cirral count of this
specimen follows (Sta. 20E) :
Dwarf male sack-like, elongate and oval (0.5 X 0.9

mm) , with 4 calcareous plates surrounding mantle
opening; surface covered with concentric bands of
short bristles and a few scattered, relatively long setae. Fig. 27. Arcoscalp~llum gauasi (Cruvel) : ,4, in-
First antennae situated distal to mantle opening. No dividual virwcd from right side; B, from rostra1 side
cirri present. (rcdrawn from Gruvel, 1909, pl. 23, figs. 8, 9 ) .
Remarks: Of the specimens dredged by the Dutch

irregularly quadrangular, forming anteriorly and at
man-of-war Siboga, Hoek (1907b, p. 110) described
their base, a strong projection. Apex poirited and on
A. / o ~ n ~ o s u rfrom
n two localities in tlie Banda Sea,
the occludent margin.
Moluccas (Sta. 227, 4"5O.ti'S, 127'59'E, 20G1 meters;
"Umbo of the carinolateral plates rounded and at
Sta. 241, 4O24.3'5, 129"4O.SrE, 1570 meters). Sub-
the same height as the apex.
sequent collections have been reported by Nilsson-
"Anterior margin of the rost~o-lateralsprojecting
Cantell ( 1921, p. 190) from Misaki, Japan, at a depth
tonards the apex.
of 600 meters, by I-iiro 11933, p. 31) froni between
ICuro-shima and Udi-shima, Kagoshima-ken, Japan "Jnfra-median lateral plates in the form of an
( 3Oo50'15"N, 12P44'31)"E I . at a depth of 55E meters,
isosceles triangle, a little irregular with the umbo at
hy Stubbings (1936, p. 20) from the region of Zanzi- the apex which forms a strong lateral projection.
bar at 1792 meters. arlti I)v Tarasov and Zevinia (1957. "Peduncle nearly cylindrical, short. nith 4 alternat-
p. 142) from the Sea oI Okhotsk at a depth of 1076 ing rows of scales, strong, large arid well calcified,
meters. The Eltanin collections were dredged off the perfectly imbricate.
Biscoe Islands, and from the Scotia Sea at depths of "Dimensions: length of capitulum 7 mm; width 3
2763-2818 meters. mm.
length of peduncle 2 mm; width 1.3
Arcoscalpellum gaussi (Gruvel, 1907) mm.
Text-fig. 27 "Habitat: Region of GAUSS wrinter station to the
Scalpellum gaussi Gruvel, 1907b, p. 159; 1909, p north of Kaiser Wilhelm I1 Land, from 380 meters.
204, pl. 23, fig. EL9. One specimen.
Diagnosis: "Capitulum compressed in its upper two- "Aflinities: Closely related to Sc. breuecarina~um
thirds, a little swollen at its base, with 14 strong plates, Hoek" (Gruvel, 1907b, p. 159).
entirely calcified, without apparent striae.
Arcoscalpellum hirsutum (Hoek, 1883)
"Carina regularly curved with umbo at apex, the
Text-fig. 25
dorsal margin very slightly concave, with the lateral
margins rounded but without ridges. Scalpellum hirsutum Hoek, 1883, p. 88, pl. 4, fig. 19;
"Terga triangular with the apex straight. Scuta Gruvel, 1905, p. 66, fig. 74; Pilsbry, 1907b, p. 25.

SYSTEMATIC A C C O U N T 63
Diagr~osis: Surface of capitulum clothed with long acute; second tooth me11 separated from first tooth;
bristles ; rostrum large, well exposed, elongate and third tooth well separated from inferior angle; inierior
oval; iriframedian latus taller than rostra1 latus; ter- angle serrate, of 12-14 long subspatulate teeth (fig.
gum extends downward nearly to whorl of lateral 28U). Maxilla I with concave cutting edge; high,
plates; maxilla I with concave cutting edge; anterior moderately shallow notch above center; 2 long, stout
ramus of cirrus I shorter than posterior ramus; caudal and 1-2 short stout spines above notch; 3-5 short,
appendage consisting of 3 fused segments. slender spines in notch; region immediately below
Material: ELanin, Sta. 63, Peru-Chile Trench, off notch bears 2 4 long, moderately stout spines; basal
Taltal, Chile (25'445, 70°58'W), 1863-1965 meters, cluster of lowest step contairis 5-6 short, slender to
2 suecimens. stout spines (fig. 22%). Maxilla I1 long, ovotriangular
Depository: USNM Cat. No. 125254. arid trilobate; marginal setae segregated in 3 distinct
clusters. Maxillary lobe low, broad, rounded apically.
Supplementary Description: The capitular plates of
Anterior ramus of cirrus I slightly shorter than pos-
this species were described by Hoek (fig. 28A), and
terior ramus (fig. 2SGI ; intermediate articles of ante-
the arthropodal structures are described below. The
rior ramus strorigly p-otuberant, those of posterior
specimen described lacks the peduncle; capitular height
7.9 mm, width 3.9 mm. Female-Labrum bullate; ramus about $6 width of anterior ramus. Cirrus I1
moderately broad and long, broadly rounded distally; modified; both rami heavily clothed with long setae.
surface and margins free of bristles; crest armed with Cirrus I about $:$ length of cirrus 11. Cirri 11-VI
30-35 simple conical teeth (fig. 286). Palps long, essentially equal in length, but with subequal rami.
slender, acutely rounded distally, superior margin free Cirri 111-VI with each articulation along greater
of bristles and setae; inferior border and distal end curvature of intermediate articles supporting 1 long
bear moderately long, stout setae. Mandible with 4 and 1 short seta and 1 extremely short, stiff bristle;
teeth including inferior angle; teeth slender, strongly interarticular areas free of setae; lateral faces with a
Fig. 28. Arcoscalpellz~m hirsutum (Hoek) : A, view of right sidc of femalc; B, crest of labrum; C,
palp; D, mandible; E, maxilla I ; F, maxilla 11; G, cirrus I ; IT, intermediate article of cirrus VI ;
I, caudal appendage. Eltanin, Sta. 63.

few short setae; setation ctenopod, 2 major and 1 Falkland Trough 153"01'S, 55051fW), 1953-1971
minor pair, at bases of major setae there are 1-2 short, meters, 1 specimen, holotype, with dwarf male.
slender setae (fig. 28H). Depository: U S N M Cat. No. 125255.
Description: Female-The unique specimerl measures
as follows: total height 8 mm, capitular height 4.5
mm, capitular width 2.9 mm.
Capitular plates fully calcified, approximate or im-
bricating; elongate oval; both margins of capitulum
convex; epicuticle thin, conspicuously hirsute; plates
sculptured with more or less equally spaced growth
Caudal appendage cor~sistsof 3 segments that ale ridges and broad. rounded, ridges emanating from
totally fused; long, very slender, about ?: height of urnbonal regions ( fig. 29A). Tergum triangular; api-
first article of pedicel of cirrus VI; apex with 5-1, cally acute and projecting; basal and occludent mar-
moderately long, stout setae; longest seta about :d; gins about :jh length of carinal margin; umbo apical.
length of appendage. Scutum quadrate; width length; occluder~tmargin
Dwarf males u ere not found. strongly convex; tergal margin strongly concave;
Rernrrrks: Aside from similar bathymetry, capitular umbo apical ; apex produced, free, strongly curved and
configuration and size, the relative proportions arid frorlting basioccludent angle of tergum. Carina bowed;
shapes of the capitular plates all point to the present extremely broad basally, tapering uniformly towards
specimens as being A. hsrsu~um. This is the second apex I fig. 290 1 ; grow ~h lines of roof strorlgly imbri-
reported occurrence of A. hirsutum, originally dredged cate; radial ridges high, broad, obtusely rounded; ele-
by the Challenger from 150:: meters in the Makassar vated so as to form knobs or beads proximal to edge
Strait between Rorneo and the Celebes (0°48'S, of each growth ridge; umbo apical. Upper latus essen-
120°58'E). The present specimeris were dredged from tially triangular; tergal and carinal margins slightly
comparable depths (1363-1905 meters) and extend the convex; scuta concave; tergal and scutal margins over-
distribution of this specirs to the other side of the lapping adjoining plates; umbo apical. Caririal latus
Pacific Ocean. Unfortunately, no comparison of the largest of lower whorl of plates; irregular in outline,
arthropodal structures can be made because of Hoek's but essentially triangular; bulk of plate free from ca-
reluctance to dissect the siugle specimen. pitular surface; apical portion of plate extending up-
An analysis of fac.ies similarity indicates that A. wards and outwards from basal margin; hasic.arina1
l ~ i r s u ~ u risn most closely related to the A. michelot- marpirls opposing below basdl margin of carina;
liarsurrs species group. The nlair~differences are the umbo apical. Inframedian latus broadly triangular;
relatively small size and lack of a dmse, velvety cover- slightly hroatlcr than high; lateral margins overlap-
ing on the capitulum and peduncle. Assigr~mer~t of this ping adjoining plates; apex overlyirlg l~asilateralangle
taxon to the A. micltelo~tianz~rngroup is telitdtivt oI scuturn; urnbo apical. Rostral latus quadrate;
uritil such time as atlditional specirnms become avail- higher than wide; umbo situated at upper angle on
able for study. occludent margin. Rostrum trii~r~gular(fig. 29C) ;
I~roatlerthan high; partially covered by rostra1 lstera;
Arcoscnlpellurn i m b r i c o t e c t ~ ~sp.
m nov. umho apical.
Plate 1X A, 'Text-fig. 29
Ptdunclr A o u t $5 height of capitulum; scales inl-
1)ingnosis: Growth ridgrs on roof of cariria strongly bricating. 2-3 times uider than high, superior 1)ortions
imbric.ate; coarse ridges err~anate from umbonal re- free of s u face. ~
gion of all capitular plates; margins of plates scal- Lahrum hullatr; broad, distally acute; superior sur-
loI~ed;basal margin of inf~arnetiiar~ latus more than face covered with ctenoid scales; crest not armed with
1", uidth of basal margin of capitulum; rostrum broad, teeth; anterior and parallel to crest are 2-3 rows of
triangular; maxilla I lacking notch in cutting edge; M-shaped or comh-like teeth surrounded by ctenoid
maxilla I1 globose, and oval in outline; surface of scales ( fig. 29D) . Palps elongate, distally acuminate;
caudal appendage covered with ctenoid scales; d ~ a r f proximal superior and inferior margins free of setae
male sack-like, and lacking calcareous plates. but covered with fine bristles; distal extremity sup-
Mtrte~ial: Eltanin, Sta. 372, type-locality, edge of porting a few setulose setae; inner superior-lateral sur-

SYSTEMATIC ACCOUNT
Fig. 29. Arcoscalpellum imbricotectum sp. nov.: A, view of left side of female; B, view of
carina and left carinal latus; C, external view of rostrum, left rostra1 latus and basal portion
of scuta; D, crest of labrum; E palp; F, mandible; G, maxilla 1; 13, maxilla 11; I, intermediate
article of cirrus VI; J, caudal appendage. Eltanin, Sta. 372 (holotype).
face covered with ctenoid scales. Mandible with 3 and 1 or 2 extremely short, stiff bristles; interarticular
long, sharp teeth and coarsely pectinate inferior angle; areas along greater curvature free of setae and bristles;
first tooth well removed from second tooth; inferior outer lateral face with 1-3 short, slender setae; setation
angle with 8-9 long, acicular teeth (fig. 29F). Maxilla ctenopod (fig. 29I), 2 major and 1 minor pair, at base
I globose, short; cutting edge not notched (fig. 29G), of each pair there is 1 short, thin, soft bristle; antero-
slanting basiproximally, 11-12 stout spines present; distal and distal margins of each article support a few
lateral surface covered with long setae. Maxilla I1 glo- cterloid scales. Cirral couilts for the holotype are as
bose, oval; marginal spines stout, and setulose; com- follows :
plete lateral face of appendage covered with ctenoid
scales; maxillary lobe low, broad, apex evenly round-
ed; surface free of setae and bristles (fig. 29H).
Anterior ramus of cirrus I about % length of poste-
rior ramus; intermediate articles of anterior ramus
slightly protuberant; intermediate articles of posterior
ramus significantly less protuberant and about q> Caudal appendage short, triangular; laterally com-
width of intermediate articles of anterior ramus; both pressed; consisting of a single segment about height
rami densely clothed with setulose setae; anterior and of first segment of pedicel of cirrus VI (fig. 295) ;
distal margin of each article supporting several ctenoid apex supporting very short setae; anterior and pos-
scales. Rami of cirrus I about y2 length of cirrus 11; terior margins and lateral faces completely covered by
rami of cirrus I1 subequal, not modified; cirri 111-VI ctenoid scales.
essentially equal in length but with subequal rami. Dwarf male sack-like, lacking cirri and calcareous
Each articulation along greater curvature of intermedi- plates; elongate and oval (0.25 x 0.70 mm) ; covered
ate articles of cirri 11-VI with one long setulose seta with concentric bands of extremely fine bristles.

Remarks: The present species is allied to Arcoscal- slightly subapical (fig. 30B). Kostral latus quadrate;
pellum liberum, to be described below. I t is, however, lateral margin higher than occludent; 1 strong ridge
easily distinguishable from this species by its wider extends from umbo to basicarinal angle. Rostrum
inframedian latus, which is not curved and free from small, narrow, less than ?Lz height of adjoining mar-
the capitulum, the hider and heavier imbricate carina gins of latera; umbo apical (fig. 30C).
and the somewhat oval outline of the capitulum. Measurements of the holotype are as follows: total
The specific name, imbricotectum, denotes the un- height 25.4 mm, capitular height 19.2 mm, capitu-
usual development of imbricate growth ridges on the lar width 11.5 mm.
roof of the carina. Labrum bullate; crest armed with about 60 triangu-
lar teeth arranged in 2 rows. Palps sparsely setose;
Arcoscalpellum latusculum sp. nov. setae limited to superior margin and distal extremity.
Plate VIII A; Text-figs. 30, 31 Mandible with 4 teeth including inferior angle (fig.
Uiagrzosis: Internal surface of scutum with several 30D, E) ; gap separating upper 2 teeth from third
adjoining parallel pits for males; inframedian latus tooth greater than that separating first from second,
extremely narrow, about 8-10 times higher than broad; and third from i n f ~ r i o rangle; inferior angle pectinate,
rostrum extremely small, less than 1/!Lheight of rostral 12 prominent teeth. Maxilla I highly spinose; cutting
margins of rostral latera; roof of carina flat with mar- edge deeply notched medially (fig. 30F). Maxilla I1
gins bounded by high prominent ribs; carinal latera triangular in outline; 3 well-segregated clusters of mar-
interdigitating below basal margin of carina; maxilla I ginal setae present; lateral face moderately setose;
with deep medial notch; distal setae on intermediate setae relatively long.
articles of anterior ramus of cirrus VI hypolasiopod; Cirrus I well separated from and about 3 length of
caudal appendage uniarticulate; dwarf males sack-like cirrus 11. Anterior ramus of cirrus I slightly shorter
and lacking calcareous plates. than posterior ramus (fig. 301) ; intermediate articles
of anterior ramus about l / broader than those of pos-
Material: Eltanin, Sta. 1150, type-locality, southwest-
terior ramus; both rami clothed with long setae ran-
ern edge of Southeast Pacific Basin (65O37'S, 121'
domly arranged on inner lateral face. At each articu-
OG'W) ,4758-4804 meters, 1specimen.
lation along greater curvature of intermediate articles
Depository: USNM Cat. No. 125256. of cirri 11-VI there are 1 long and 1 or 2 short setae,
Description: Female-Capitulum elongate and oval; and on the distal margin of each article there is a
widest at middle; both margins convex; plates orna- single row of ctenae, the number increasing from an-
mented with distinct growth ridges and low radial terior to posterior; no spines present along interarticu-
ridges on paired plates (pl. VIII A) ; all plates covered lar areas of greater curvature. On outer ramus of cir-
with thin, short, and moderately pilose cuticle. Ter- rus I11 there are 1-3 spines, but none on inner ramus.
gum somewhat crescentic ; protuberant apically; plate Setation ctenopod, on outer rami of cirri 111-VI 4
about height of capitulum; lateral extremity not pairs, on inner rami 5 pairs (fig. 305) of setae.
reaching to carina. Lateral and occludent margins of Cirral counts for the holotype are as follows:
scutum convex; plate longer than wide, basal margin
slightly concave; plate irregularly subquadrate, umbo
apical; several pits for males present along occludent
margin (fig. 30A). Carina evrnly curved; umbo api-
cal; roof flat, bordered laterally by prominent high
ridges. Upper latus with 5 unequal sides; scutal mar-
gin longest and slightly concave; margin bordering
carinal latus about 3 times length of margin bordering Caudal appendage short; about 1/:5 height of first
inframedian latus; umbo terminal. Carinal latus trape- segment of pedicel of cirrus VI; 2 terminal setae of
zoidal; somewhat greater in area than upper latus; equal length present; setae about as long as appendage
umbo at lower third of plate; not projecting beyond (fig. 30K).
the carina; plate with 2 strong radial ridges extending Dwarf males sack-like, without calcareous plates
from umbonal area to upper and lower lateral margins (fig. 31A) ; surface of sack ornamented with concen-
of plate. Inframedian latus same height as lateral mar- tric bands of fine spines; males aligned in single row
gin of carinal latus; width of plate % height; umbo along occludent edge of scutum, each in separate pit,

ranging in size from 0.85 X 1.15 mm to 0.90 X dredged from 512 meters south of Yensacola, Florida,
1.25 mm. On one specimen there mere uhat appear in the Gulf of Mexico. The major feature uniting these
to be vestigial cirri armed with long setae. three species is the extremely narrow inframedian latus.
Remarks: The present species is closely related to I n the present specimens it is rectangular, but it ap-
A. tenup (Hoek, 1883, p. 119) dredged by the Chal- pears to be virtually triangular in the above two spe-
lenger from 2914 meters on the Atlantic-Indian Ridge cies. Arcoscalpellurn tenue lacks the ribs bordering
between Prince Edward arid Crozet Islarids ( 4604GfS, the roof of the carina, but these are prcsent in A . semi-
4S031'E). and A . semisculplum iPilsbry, 1907a, 11. 6 2 ) sculp~urnand the new species. The caririal latera meet
Fig. 30. AT-coscalpellumlatusculun~sp. nov.: A, internal vipw of left scutum showing parallel depres-
sions for males above circular adductor muscle depression; B, view of right inframedian latus; C, rostrum
and adjoining margins of rostra1 latera; D, mandible; E, cnlargcd view of inferior angle of same man-
dible; F, maxilla I ; G, maxilla 11; H, pllp; I, cirrus I ; J, intermediate article of cirrus VI; K,
caridal appendage. Eltanin, Sta. 11501 (holotype).

Z L ~ n~ w . : A, dwarf male; B, enlarged view of concentric hands

Fig. 31. Arcoscall~ellum I U ~ I L S C I L ~ sp.
of spines covering mole; C, cnlarged biew of w h a ~appcars to be a specialized attacliment organ
found on right side of male shown in fig. A ; D, cnlarged view of rudimentary natatory appendages
of male; E, enlarged view of what appears to be reproductive system of male. Eltenin, Sta. 1150.
below the carina in "an irregular suture" in A. semi- mer Archipelago (63051fS, G203gfW), 128-165 me-
sculpturn, as they do in the present specimen, but ap- ters, 6 specimens.
parently do riot in A. Irnue. In neither of these pre- Depository: USNM Cat. No. 125257 (Sta. 410) ;
viously desc,ribed species is there a chitinous bridge 125258 (Sta. 426) ; 125259 (Sta. 439).
betneen the tergum and carina. arid this feature should
Szrpplementnry ljcscription: Female-Lateral surface
serve to readily separate them. The upper latus in
of palp sparsely covered with ctenoid scales. Mandible
A. L ~ I L Uis~ esserltially the same as that found in the
present specimens, but dilIers from A. semisculpturn with 3 teeth and serrate inferior angle (fig. 32H) ;
inferior angle variable (cf. fig. 321-L). Maxilla I with
in being pentagonal rather than Lriangular.
shallow subapical notch in cutting edge (fig. 32M) ;
The specific name, clerived from the Latin, denotes
inferior angle slightly recessed. Maxilla I1 ovotriangu-
the small size of the inframediari latus.
lar; somewhat bilobed. Maxillary lobe low, broad,
Arcoscalpellum liberum (Nilsson-Cantell, 1930) apex broadly rounded; surface free of setae and
Plate IX H, Text-Gg. 32 bristles.
Scnlpcllum liberurn Nilsson-Caritell. 1930h, p. 233, Anterior rarnus of cirrus I about $5 length of poste-
figs. 3, 4 ; 1931, p. 7. rior ramus, intermediate articles slightly protuberant;
intermediate articles of posterior ramus about 4 h width
Diagnosis: Capitulum somewhat triangular in out-
line; rostrum large, clearly exposed, protuberant and of anterior ramus. Distal end of each segment with
not overlapped by rostra1 latera; inframedian latus row of ctenoid scales. Kami of cirrus I1 unequal; cirri
large, triangular, convex apic.obasally with apex free 111-VI subequal. Citrus I1 not greatly rnodilied. Each
of capitulum; cutting edge of maxilla I short, subapi- articxlation along greater curvature of intermediate
cally notched; caudal appendage uniarticulate. laterally artic.les of cirri 11-VI with 1-2 long, slende~setae and
compressed, apex free of long setae, arid lateral fares 1-2 extremely short, stiff bristles; interarticular areas
lacking ctemoid scales. along greater curvature free of setae arid Lristles; dis-
Material: Eltanin, Sta. 410, between Elephant Island tal end of each intermediate article with row of cterloid
and Aspland Island, South Shetland Islands (61 I r S , scales parallel to the articulation. On the inner face of
50°09'W), 220-240 meters, 2 specimens; Sta. 426, be- the inner rarnus of cirri 111-V there are 1-2 short,
t ~ e e nElephant Island and Joinville Island. South Shet- setulose setae. Setation of posterior cirri ctenopod; 2
land lslands (62'275, 57"58'W), 809-1 116 meters. major and 1minor pair all uith setules, at base of each
ti specimens; Sta. 4.39, north of Brabant Island, I'al- pair of setae there are 1-3 short, slender setae.

Measurements of the two dissected specimens from Caudal appendage uniarticulate, triangular, laterally
Station 439 are as follows: compressed, acutely rounded distally (fig. 32P) ; an-
Total Capitular Capitular terior and posterior margins support short, thin, stiff
Specimen Height Height- W-~ d t-
h bristles; no terminal setae present; appendage about
.
- - -
1 8.2 4.7 3.6 :% height of first segment of pedicel of cirrus VI.

2 9.7 4.6 3.7 Dwarf male sack-like, without calcareous plates; one
Cirral counts for these two specimens are: male found in each pouch on sides of aperture; measur-
ing approximately 0.4X 0.8 mm; no cirri visible; sur-
face covered with concentric rows of minute bristles.
Remarks: Nilsson-Cantell (1930b, p. 233) reported
A. liberum from Neumayer Channel, Palmer Archi-
pelago (64'4S'S, 63"31'W) at a depth of 259 meters.
The Eltanin collections are from localities immediately
north and east of the type-locality and bracket (128-
1116 meters) the previously known depth.
The capitular morphology agrees in all essentials
with the description presented by Nilsson-Cantell.
However, he did not point out that the scutum is over-
Fig. 32. Arcoscalpellum liberum (Nilsson-Cantell) : A-F, side view of females; G, dwarf male in sack
attached to arthrodial membrane on occludent margin of scutum; H, mandible; ILL, enlarged view of
third tooth and inferior angle of mandibles; M, N, maxillae I ; 0, palp; P, caudal appendage. Eltanin,
Sta. 426 (figs. A-F only) and Sta. 439.

lapped by the upper latus, adjoining portions of the Material: Discovery, Sta. 187, Neumayer Channel,
inframedian latus and rostral latus, and in turn over- Palmer Archipelago (64048'30r'S, 63 31'30"W) , 259
laps the tergum. meters, 1 specimen; Sta. 190, type-locali~y,Bismarck
Dwarf males were found in both the specimens Strait, Palmer Archipelago (64'56'S, 65'35'W), 315
dissected. The long projection at the end opposite to meters, 2 specimens.
the first antennae, mentioned and illustrated by Nils- Original Description: "Female. Capitulum wholly
son-Cantell, was not observed. covered by a thick and very hairy cuticle, and this
The specimens were situated on spines of an echi- must first be removed before one can study the valves.
noid and the stalk of an antipitharian. There are fourteen plates separated by chitinous inter-
spaces, sometimes rather widely as in the larger indi-
Arcnscalpellum magnaecarime
viduals, especially round the upper latus. The valves
(Nilsson-Cantell, 1930)
are distinctly sculptured by growth ridges and longi-
Text-fig. 33
tudinal radiating lines.
Scalpellum magnae-carinae Nilsson-Cantell, 193Db, "Scutum quadrangular, with recurved umbo, occlu-
p. 236, figs. 5-6. dent margin slightly convex.
''r
Diagnosis: "Female. Capitulum hairy, with fourteen rergum triangular, with the apex acute, occludent
well calcified plates, separated by chitinous interspaces, margin like the others a little convex.
the valves sculptured. Scutum quadrangular, apex re- "Carina very typical, rather broad and regularly
curved and occludent margin convex. Tergum triangu- arched. Umbo apical. Dorsal roof very convex, with-
lar, with nearly straight occludent margins. Carina out lateral ribs, and merging into the well-developed
regularly bowed with apical umbo. Dorsal roof sides.
strongly convex. Upper latus pentagonal with apical "Upper latus pentagonal, with the umho at the apex.
umbo. Rostrum triangular with apical umbo. Rostral The scutal margin, which is the longest, hollowed out,
latus quadrangular, wide rather than high. Inframe- the carinal margin convex.
dian latus triangular with apical umbo. Carinal latus "Rostrum triangular, rather small with the umbo at
quadrangular with apic.al unibo. Peduricle with trans- the apex, not overlapped by the rostral latera.
versely elongated scales with chitinous interspaces. "Rostral latus quadrangular, wide rather than high,
Mandible w ith three teeth arid a pectinated inner angle. with the umbo at the upper rostral corner.
Maxilla I with an iridistirict notch. Maxilla I1 ~ i t h "Inframedian latus triangular, with the umbo at the
bristles in two groups. Caudal appendage single- apex.
jointed, with bristles, of the same length as the proxi- "Carinal latus quadrangular with the umbo also at
mal segment of the protopotlite of cirrus VI" (Nilsson- the apex. IJpper margin hollowed out; carinal margin
Cantell, 1930b, p. 237). convex.
"Peduncle shorter than the capitulum, with trans-
versely elongated scales separated by chitinous inter-
spaces. The cuticle with many hairs.
Lcngth of Breadth of Length of Breadth of
Capitulum Capitulum Peduncle Peduncle
Hulot~pe 13 8 6 4
Paratype 19 14 11 6
"Mouth parts : Palpus conical, pointed and with bris-
tles along one edge. Mandible with three teeth and
a pectinated inner angle. Maxilla I with straight front
edge: a very small, sometimes barely developed notch,
can be traced on the front edge. Maxilla I1 with bris-
Fig. 33. Arcoscalpellum rnagnaecari~~ae(Nilsson-Cantcll) : A,
view of carina, entire individual from left side, and rostrum
tles along the whole concave front edge. In the poste-
and rostrolaterals of holu~ype; B, palp; C, mandible; D, first rior Part a small lobe with bristles is differentiated-
maxilla; E, pedicel of cirrus VI and caudal appendage of Behind this is a large maxillary lobe with the opening
paratype; I?, sccond maxilla; G, pedicel of cirrus VI and for the maxillary grand.
caudal appendage of holotype (redrawn from Nilsson-Cantell, "cirrus I with rami of different lengths. cirrus 11
1930, figs. 5, A-C; 6, A-F). also with unequal rami, shorter than the following

cirri, in which the rami are equal. Cirrus V l with six carinal latus it is much related to S. regium, Wyv.
pairs of spines on the front edge of the segments. Thomson, 1873, S. moluccanum, Hoek, 1883, and S.
"The caudal appendage is single-jointed, broad and gigas, Hoek, 18li3, of the Challenger Expedition. These
flat and covered with marly bristles. The appendage is species I have studied in the British Museum, and have
of the same length as the proximal segment o f cirrus found them in other characters well separated. I n the
VI. In the paratype it is even shorter. internal parts this new species differs especially by the
"The com1)lemental male [dwarf male 1 could riot be single-jointed caudal appendage. In S. rcgium the ap-
studied here. pendage is four-jointed (according to Hoek) ; in S.
I 11 111 IV V VI Ca gigas I found in the type material four to six segments;
in S. rnoluccccnurn the appendage has five to six seg-
Holotype G 13 14 15 16 17 1
- - - - - ments (Hoek, 1883, and Nilsson-Cantell, 1927) " (Nils-
11 14 15 17 17 18 son-Cantell, 1930b, pp. 237-239).
Paratypc 6 13 xx xx xx xx 1
- - - - - -
11 17 15 xx xx xx Arcosculpellum michelottianum (Seguenza, 1876)

Plate IX B, Text-fig. 34
rhis new species, which I have nained S. magnae-
'6,
carinae, from the large and well developed carina, be- Scalpellum rnichelot~ianurn Seguenza, 1876, p. 381,
longs, like S . liberum, to the group of S . velutinurn. pl. 6 , figs. 15-25, p. 464, pl. 10, fig. 26; de Alessandri,
From this species it is bell differentiated by several 1895, p. 263, in part; 1897, p. 47; 1906, p. 251;
external arid internal characters. In the shape of the Withers, 1953, p. 225.
Fig. 34. Arcoscalpellum rnichelottianu~n (Seguenza) : A, lahrum and palps; E, mandihlc; C, maxilla
I; D. ~naxilla 11; E, left cirrus I ; F, intcrnlcdiatc article of cirrus VI; G, caudal appendage of 2
fused segments; H-J, view of right side ol fcmales. Eltanin, Sta. 18.

Scalpellurn velutinum Hoek, 1883, p. 96, pl. 4, figs. simply bowed; roof evenly rounded, with slight medial
10, 11, pl. 9, figs. 7-9; Weltner, 1897, p. 251; Gruvel, and bordering ridges; basal margin rounded, and not
1902a, p. 56, pl. 2, figs. 3c, 10a, lob, 14, pl. 3, figs. 1, reaching to basal margin of capitulum; umbo apical.
27-31, pl. 4, figs. 6, 11-22; 1905, p. 73, fig. 83; An- Upper latus narrow; 4-sided; essentially same height
- -
nandale, 1906, p. 83; Hoek, 1907b, pp. 59, 79, 87; as adjoining margin of scutum; umbo apical. Carinal
Pilsbry, 1907b, p. 26, pl. 3, figs. 2, 3 ; Annandale, latus tallest of plates in lower whorl; carinal border
1908, pl. 4, fig. 7 ; 1911, p. 588; Gruvel, 1912, p. 2 ; of plate not projecting; carinal margins of latera
Annandale, 1913, p. 229; Hoek, 1914, p. 4 ; Calman, abutting below basal margin of carina, but not inter-
1918, p. 108; Gruvel, 1920, p. 27, pl. 1, figs. 8-10, digitating; umbo apical. Inframedian latus triangu-
pl. 7, fig. 4; Weltner, 1922, p. 75; Barnard, 1925, p. 1; lar, taller than adjoining margins of latera; umbo
Nilsson-Cantell, 1927, p. 743, fig. 1 ; 1928, p. 4; Broch, apical. Rostra1 latus shortest of lower whorl; apical
1931, p. 18; Nilsson-Cantell, 1931, p. 7 ; Stubbings, and basal margins essentially parallel; basioccludent
1936, p. 28, fig. 12; Tarasov and Zevina, 1957, p. 24, angles approximate, except at supero-occludent um-
fig. 9H; Stubbiqs, 1967, p. 234. bonal angle. Rostrum very large, wholly exposed, tri-
Scalpellurn exirniurn Hoek, 1883, p. 100, pl. 4, figs. angular, acute basally; umbo apical.
6, 7, pl. 9, fig. 10; Weltner, 1897, p. 247. Peduncle about 1/5 to l/q height of capitulum; sur-
Scalpellurn sordidurn Aurivillius, 1898, p. 190. face covered with relatively close-spaced, large scales;
Scalpellum erecturn Aurivillius, 1892, p. 192; Gru- exposed portions of scales with irregular borders.
vel, 1905, p. 74. Measurements of the 3 specimens collected by the
Scalpellurn alaturn Gruvel, 1900b, p. 192; 1902a, Eltanin:
p. 57.
Total Capitular Capitular
Diagnosis: Rostrum large, elongate, triangular, acute Specimen Height Height Width
basally; inframedian latus higher than adjoining mar- --
1 10.5 8.8 3.8
gins of latera; adjoining margins of carinal latera not 2 6.0 4.6 2.5
interdigitating below carina, umbo apical; roof of 3 3.2 2.8 1.4
carina broad, gently rounded, and basal margin of
plate evenly rounded; anterior ramus of cirrus I Those of the Eastward are:
shorter than posterior ramus, and articles of both rami Total Capitular Capitular
moniliform; labrum broad and greatly max- Specimen Height Height Width
illa I with subapical notch; maxilla I1 triangular and 1 11.8 8.9 4.3
trilobate; caudal appendage with 2 fused segments. 2 11.4 8.9 4.5
Material: Eltanin, Sta. 18, Labrador Basin, off south- Labrum bullate; moderately protuberant and broadly
ern tip of Greenland (58' 15'N, 4Z03G'W), 3404-3422 rounded (fig. 34A) ; surface and borders free of stiff
meters, 3 specimens; Eastward, Sta. 7552, Hatteras bristles; crest armed with about 20-25 simple, conical
Plain, east of Charleston, South Carolina (33'39.5'N, teeth. Palps moderately long, triangular, broad, acutely
75041fW), 3010 meters, 2 specimens. rounded distally; superior proximal margin free of
Depository: USNM Cat. No. 125260. setae; inferior margin, lateral face and distal extremity
Supplementary Description: Female-Capitulum with armed with stout setae. Mandible with 4 teeth includ-
fully calcified approximate plates covered with moder- ing inferior angle (fig. 34B) ; teeth long and acute;
ately long bristles, not villous; elongate and oval in second tooth closer to third than to first tooth; third
outline, width approximately !$ height (fig. 34H-J) ; tooth equidistant between second tooth and inferior
occludent and carinal margins slightly convex; plates angle; inferior angle slender, protracted, comb-like,
ornamented only with irregularly spaced growth ridges. of 5-9 long, subspatulate teeth. Maxilla I with 3 dis-
Tergum somewhat ovate; carinal, occludent and basal tinct clusters of spines on cutting edge (fig. 34C) ; deep
margins convex; width slightly less than length; basi- notch present above center; 2 long, stout and 1 short,
carinal angle extending below midpoint of capitulum; stout spine above notch; occasionally 1 short, stout
umbo apical. Scutum with occludent and lateral mar- spine at side of notch. Maxilla I1 short, triangular;
gins essentially ~ a r a l l e l ;basal margin slightly convex margin deeply notched with setae occurring in 3 dis-
or straight; tergal margin concave; width y2 or less tinct clusters; maxillary lobe extremely broad and
height; y2 height of capitulum; umbo apical. Carina short, apex flat; surface free of setae and bristles.

S1 STEMATIC ACCOUNT 73
Cirrus I with subequal rami; intermediate articles arthropodal structures of these stages have never been
of anterior ramus strongly protuberant (fig. 34E ) ; described. In addition, there is seemingly much con-
a ~ ~ i c l eofs posterior ranlus about 24; ctidth of anterior fusion regarding the identity, hiogeopraphy and bathy-
ramus; both rami heavily clothed ~ i t hlorlg setae. metry of A. rr~icheloltiar~um even though it has beer1
Cirrus 11 normal, rami about twire length of rami of dredged and reported rlumerous times.
cirrus I. Cirri gradually increasing in length from Alrhough the Eltanin collections consist of only
cirrus TI posteriorly; rami subequal. Each articulation young stages of this species, certain details warrant
along greater curvature of intermediate articles of cirri cliscussion at this time. Most reports indicate that the
Ill-VI ~ i t l 1-2 i long setae; commonly orle short seta rostrum is small: whereas it is exceedingly large in all
at the articulation on the lateral face; interarticular three of the specimens reported here. I n all likelihood
areas along greater curvature free of setae. Setation negative allometry woultl account for the fact that this
ctenopod (fig. 3 4 F ) ; 2 major and 1 minor pair on structure is refcrred to as being extremely small. As
cirri 111-V; 3 major and 1 minor pair on cirrus \TI; for the shal)e of the basal margin of the carilia, M hich
at base of major setae there are 1-2 short, slender also seems to vary considerably, it need orily be pointed
setae. out that either there are several species involved or
Caudal appendage consists of 2 stout, fused seg- confused as A. velutinum, or different stages of growth
ments and 6-9 terminal setae of varying lengths (fig. are represented. As for the roof of the carina, which
34G) ; appendage I,($ to slightly less than !h height of in the present species is gently rounded with only in-
first segment of pedicel of cirrus VI. dications of bordering ribs, it would logically appear
that this plate is subject to certain changes in shape
1 I1 111 IV v VT C.1
with continued growth. What is needed at this time is
Ellarrin, Station 18 a complete series of specimens, in order to clarify these
Spccinien 1 7 13 xx xx xx 19 2 points of discrepancy.
9 14 xx xx xx xx Of the arthropodal structures, the number of seg-
7 13 13 17 xx xx 2 ments in the caudal appendage is also quite variable,
9 14 16 17 xu xx ranging Irom 2 to allout 6 or 8 in the Eltanin speci-
mens. Certain differences in the manner of counting
Specimen 2 5 9 12 13 14 15 2 appear in the literature, which is espzcially evident
- - -
7 10 12 14 14 15 when the segments that are fused are counted as only

5 9 12 13 14 17 2 one segment. If at all possible cou~itsshould be based
7 11 12 14 11 10 on the total number of segments, fused or not. Insofar
as the notch in maxilla I. Nilsson-Cantell 11927,
Enstzonrd, Station 7552
p. 745) states that there is none, although there is
Spccimen 1 6 19 19 21 23 25 2
some indication of a slight notch or induntation in the
10 17 19 21 xx 22
culting edge in the illustraLion he presents of this ap-
7 xx 19 22 23 24 2 pendage. In the present specimens this iridentatiori is
- - - - -
10 16 20 20 22 22 more and to all intents and purposes the
record should indicate that there is a distinct notch
Remarks: In describing A. velutinurn, Hoek (1E83,
in the first maxillae. Although Nilsson-Cantell (1927)
p. 96) neither described nor illustrated the arthropodal
reports the presence of 4 pairs of setae on the inter-
structures of this species. Subsequent workers, notably
mediate articles of the posterior cirri, there are only
Gruvel (1902a) and Nilsson-Cantell (1927), attempted
3 on the specimen here figured. This difference may
to rectify this situation, but their descriptions are brief
be attributable to age, but once again additional speci-
and the illustrations are inadequate for present-clay
mens are necessary to clarify this point.
requirements. Withers (1953), without offering fur-
ther discussion, equated A. velutinum with A. rnichelol-
Arcoscnlpellum multicostatum sp. nov.
tianum. I n checking this synonymy the writers are
Plate IX F; Text-figs. 35, 36
forced to concur with Withers.
The present specimens are only tentatively assigned Diagnosis: Capitular plates ornamented with strong,
to A. michelottianurn largely because so little is known narrow radial ridges emanating from region of um-
about the early stages of this species, and because the bones; carinal latera produced, extending curvilinearly

specimens; Sta. 1003, off Joinville Island, Antarctic

Peninsula (62"41rS, 54'43'W), 210-220 meters, 2
specimens.
types; Sta. 1003).
Description: Female-Capitulum elongate, ovotri-
angular; strongly acuminate apically ; occludent mar-
gin straight; carinal margin strongly convex; all ca-
pitular plates approximate, ornamented with poorly
discernible growth ridges and strong, numerous, nar-
row, acute, radial ridges emanating from the umbonal
region (pl. IX F) , rendering a scalloped appearance to
the margins of the plates. Tergum triangular, strongly
acuminate at extremities; carinal margin slightly con-
vex; approximately twice length of straight occludent
margin and y3longer than slightly concave basal mar-
gin; umbo apical. Scutum quadrilateral; plate twice
as high as wide; carinal and occludent margins slightly
convex; scutal and basal margins either straight or
concave; umbo apical, slightly produced and project-
ing over basioccludent corner of tergum. Carina
bowed, roof evenly rounded; tectum, paries and intra-
paries not clearly set off from one another; apex of
plate extending to a height slightly more than the mid-
point of the tergum; umbo apical. Upper latus with 5
unequal sides; scutal margin longest folloued by ter-
gal, carinal, basal and basiscutal; umbo subapical;
slightly elevated. Carinal latus largest of lower whorl
of plates; plate hook- or scimitar-shaped; carinal ex-
tremity extending beyond border of capitulum; lateral
extension of latera approximating along midline of
Fig. 35. Arcoscalprllurn multicostatum sp. nov.: A- carina, but not interdigitating; umbo at tip of projec-
D, view o l right side of females with normal com- tion. Inframedian latus essentially rectangular, slightly
plement of rapitular plates; E, view of right side of
higher than broad; umbo subcentral (fig. 35) ; greatly
female lacking upper latus. Eltanin, Sta. 671
(Iiolotypc, fig. A). elevated above level of capitulum; ridges emanating
from umbonal region in all directions except for basal
pie-shaped wedge. Rostra1 latus 4-sided, but essentially
and approximating, but not interdigitating below basal
margin of carina; umbo of inframedian latus elevated triangular in outline; scutal margin concave; basal
above level of capitulum, initially pointing directly and lateral margins convex; plate slightly wider than
outwards, but in later growth stages directly dorsally high; umbo situated at rostral angle. Rostrum moder-
toward carinal margin; mandible with 4 teeth includ- ately large, triangular; clearly exposed and not cov-
ing variably serrated inferior angle; maxilla I with ered by rostral latera; umbo apical.
deep subapical notch; maxilla I1 ovotriangular; caudal Peduncle less than height of capitulum; armored
appendage uniarticulate; dwarf males sack-like, lack- with few scales; scales not fully calcified, about 5 times
ing calcareous plates. broader than high, superior margins smooth and evenly
Material: Eltanin, Sta. 671, type-locality, south of rounded.
Soutll Georgia (5/E041'S, 3803SrW), 220-320 meters, 5 Measurements of the 7 specimens as follows:

Total Capitular Capitular crest armed with 30-35 M-shaped or comb-like teeth,
Specimen Height Height Width above which are 4-6 subparallel rows of short setae
- - -
Station 671 1 (Holotype) 13.9 11.1 5.9 in clusters of 3 4 (fig. 36A). Palp tubuloconic; su-
2 13.9 10.1 5.5 perior margin with short, stiff bristles; inferior mar-
3 11.7 9.3 4.4
4 7.8 6.5 3.2
gin free of setae and bristles; long setulose setae pres-
5 3.9 3.3 1.8 ent on outer face and distal extremity. Mandible with
Station 1003 1 5.7 3.8 2.2 3 teeth excluding inferior angle; second tooth more or
2 5.2 3.4 2.4 less equidistant between first and third tooth (fig.
Labrum bullate; superior surface and margins 36B) ; inferior angle serrate, with 2-11 triangular to
sparsely covered with extremely short, stiff bristles; spatulate and moderately long spines (fig. 36C-E) ;
Fig. 36. ArcoscaZpellum multicostatum sp. nov.: A, portion of crest of labrum; B, left mandible; C-
E, inferior angles of right mandibles; F, left maxilla I ; G, enlarged view of right maxilla I ; H, max-
illa 11; I, right palp; J, right cirrus I ; K, intermediate article of right outer ramus of cirrus VI; L,
caudal appendage. Eltanin, Sta. 671 (holotype, figs. A, B, E-L).

maxilla I with shallow or deep notch at or above mid- heavily armored with imbricating plates, and the cau-
point of cutting edge (fig. 36 F, G) . Maxilla I1 triangu- dal appendage consists of 5 segments and is longer
lar in outline; basal and superior proximal margins than the pedicel of cirrus VI. In many respects A.
and distal extremity covered with long setae; maxil- condensum is related to the boreal species A. cornutum
lary lobe moderately high. stout, rounded apically, free (Sars), which also has a heavily armored peduncle,
of setae and bristles (fig. 36H). sulcate roof on the carina, but a uniarticulate caudal
Anterior ramus of cirrus I about 4/5 length of poste- appendage. Furthermore, the inframedian latus in A.
rior ramus (fig. 365) ; intermediate articles protuber- condensum is essentially triangular whereas it is essen-
ant; posterior ramus about % -!i width of anterior. tially quadrate to pentagonal in A. cornutum as well as
Cirrus I1 not modified. Rami of cirrus I about $$ A. multicostaium. Of the many new South African spe-
length of rami of cirrus 11. Cirri 111-VI essentially cies names introduced by Barnard (1924), A. subala-
equal in length, with subequal rami; each articulation t u m is readily separable from the present species by
along greater curvature of intermediate articles with 2 its heavily armored peduncle, the basally projecting
long setae; interarticular areas free of setae; setation umbo of the somewhat triangular inframedian latus
of intermediate articles along lesser curvature of cirri and triangular rostra1 latus. Chaetotaxis of the inter-
11-VI ctenopod; 3 major and Z minor pair, 1 or 2 mediate articles of cirrus VI in Barnard's species is 5
short setae at bases of each pair of long setae (fig. whereas it is 4 in A . multicostatum and A. capense
36K). (Barnard). The latter species may be distinguished
Cirral counts for 3 specimens from the type-locality from the present one by its dwarf male having 4 cal-
follow : careous plates, the caudal appendage consisting of 3
fused segments, the inframedian latus having a basal
umbo projecting basally toward the occludent margin,
Specimen 1 (Holotype) 8 12 14 14 15 14 1 and the tergum having a strongly convex occludent
- - - - - -
10 13 14 15 15 15 and concave carinal margin. The dwarf male of A.
7 13 13 14 15 15 1 eumitos (Barnard) also has 4 calcareous plates. Al-
- - - - - -
10 13 13 14 xx 15
though the tergum of this species approximates that
found in A. multicostatum, the roof of the carina is
Specimen 2 10 xx xx 19 23 20 1
- - - - - - concave, and the umbo of the inframedian latus is
8 xx 17 xx xx 18 situated at the basirostral angle. In none of the pre-
viously described species does the carinal latus project
as far, nor does the projection curve apically.
Specimen 3 x 12 13 13 14 14 1
Arcoscalpellum parallelogramma (Hoek, 1883)
8 12 13 14 14 15 Plate IX C; Text-figs. 37, 38
Scalpellum parallelogramma Hoek, 1383, p. 83, pl.
3, figs. 15-16, pl. 9, figs. 1-5; 1884, p. 21; Weltner,
Caudal appendage uniarticulate; about lh to TJ 1897, p. 249; Gruvd, 1905, p. 61, fig. 67; Nilsson-
length of both segments of pedicel of cirrus VI (fig. Cantell, 1931, p. 8.
36L) ; slender, oval in section; posterior border clothed Diagnosis: Capitular plates ~ i t hfaint radial ridges;
with extremely short bristles and long slender setae; rostrum rectangular in outline, pyramidal, with umbo
terminal cluster of last segment consists of 6-10 long subapical; carinal latera interdigitating below carina;
thin setae about length of appendage. inframedian latus projecting free, umbo basal; pedun-
Dwarf males sack-like, lacking calcareous plates and cle with minute scales; maxilla I with deep subapical
cirri; sack elongate-oval (0.5 X 1.0 mm) , and covered notch; maxilla I1 triangular in outline with small patch
with close-spaced concentric bands of fine bristles. of short basal bristles; caudal appendage multiarticu-
Remarks: Arcoscalpellum multicostatum is related to late, with 3-6 articles; dwarf males sack-like, with 4
A. candensum Nilsson-Cantell (1921, p. 202) from calcareous plates.
Goto-rett6, Japan. However, in the Japanese species Material: Eltanin, Sta. 340, between Burdwood Bank
the ~ l a t e slack the strong and uniform ribbing, the and Falkland Island (53'08'S, 59'23'W), 567-578
roof of the carina has a deep sulcus, the peduncle is meters, 5 specimens; Sta. 557, northeast of Falkland

SYSTEMATIC ACCOUNT
Fig. 37. Arcoscalpellum parallelogramma (Hoek) : A, view of right side of female; B, view of abut-
ment of carinal latera and basal portion of carina; C, internal view of scutum showing depression
for dwarf males above oval depression for adductor muscle; D, E, external and internal views, re-
spectively, of carinal latus; F, L, external and internal views of inframedian latus, respectively; G,
I, internal and external views, respectively, of rostra1 latus; H, K, front and lateral views, respec-
tively, of rostrum; J, apex of c~arina. Eltanin, Sta. 558.
Island (51'565, 56'39'W), 8 5 5 4 6 6 meters, 3 speci- ated close to occludent margin and at midpoint of
mens; Sta. 558, northeast of Falkland Island (51'58'S, valve height; depression for dwarf males about 1/4
56'38'W), 646-845 meters, about 140 specimens. distance from apex of plate (fig. 37C). Carina bowed;
Depository: USNM Cat. No. 125264 (Sta. 340) ; roof flat; bordered by high, broad ridges; umbo apical
125265 (Sta. 557) ; 125266 (Sta. 558). (fig. 37J). Upper latus essentially triangular; tergal,
scutal and basal margins concave; carinal mar,'win con-
Supplernenlary Description: Female-Capitulum ro-
vex; scutal and tergal margins nearly equal in length;
bust; roughly triangular in outline (fig. 37A, B) ; cuti-
cle covering plates nonhirsute; plates ornamented with umbo apical. Carinal latus large; triangular projec-
growth ridges and very faint radial ridges; occludent tion extending beyond border of capitulum; umbo api-
and carinal margins convex. Tergum triangular; basal cal; lateral extensions (fig. 37B) interdigitating below
margin nearly as long as carinal margin; occludent basal margin of carina. Inframedian latus triangular
margin about length of carinal margin; low, weak in outline, basilateral alar-like projections occur ex-
ridge extending from apex to basicarinal angle. Scu- tending below external surface (fig. 37F, L) ; umbo
tum quadrate; twice as high as broad; umbo apical basal; apical margin of plate about ?L2 length of cari-
and slightly projecting over basal margin of tergum; nal margin; basal half of plate elevated above general
lateral margin indented just below tergal margin; ad- surface of capitulum. Rostra1 latus quadrangular (fig.
ductor muscle depression large, oval, and deep, situ- 37G, I ) ; broader than high; umbo situated just below

Fig. 38. A~~coscalpr!lurr~

paml!elogramma (Hoek) : 11, mandiblr: E-D, enlarged \-ietv of inferior angle of
mandibles; E-G, maxillae I ; H, maxilla 11; I , labrunl and palps; J, selected portion of crest of labrum;
K, intermediate article of cirrus V I ; L-N, caudal appendages shorn-ing range in number of srgments;
0, male. E!/at~irz,Sta. 538.

occludent-scutal angle. Rostrum small; pyramidal (fig. greater curvature devoid of setae. Setation ctenopod;
37H, K ) ; roughly quadrate in outline; externally visi- 5-6 pairs on cirri 111-VI; one shcrt seta commonly at
ble; exposed portion hourglass-shaped. bases of each major pair.
Peduncle cylindrical; fully covered with minute Caudal appendage of 2-6 articles (fig. 38L-N) ;
scales. first and second proximal articles commonly fused;
Measurements (in mm) of the specimens: height of appendage from shorter than first article of
pedicel to slightly longer than entire pedicel of cirrus
Total Capitular Capitular VI; terminal article with at least 4 long setae.
Height Height Width
Cirral counts are summarized below for four speci-
Station 340 mens from Station 558. Range ( R ) , and mean values
n 5 5 5 ( X ) for the number of segments in the anterior ( a )
R 8.3-24.4 6.3-17.1 3.4-9.4
X 15.50 11.08 6.16
and posterior (p) rami:
Station 557
n 3 3 3
R 10.6-17.5 8.9-15.0 4.8-7.8
x 13.03 11.13 5.9
Station 558
n 35 36 36
R 3.8-22.4 3.5-17.9 1.9-9.5
X 12.81 10.33 5.51
Labrum bullate; 30-40 short, fine bristles scattered

on each lateral margin; dense cluster of 3 0 4 0 fine,
short bristles present on extremity; crest armed with
Dwarf males sack-like, elongate and oval (0.60 X
about 35 teeth, central teeth triangular, lateral teeth
0.95 mm to 0.65 X 1.0 mm) ; surface of sack covered
either M-shaped or comb-like (fig. 381). Palps elon-
with concentric bands of short bristles; 4 calcareous
gate; narrowly triangular; setae absent from superior,
plates, oval to circular in outline; 2 plates small
but present along basal margin. Mandible with 4 teeth
(0.030 X 0.049 mm) and 2 large (0.060 X 0.075 mm) .
including inferior angle; second tooth well separated
Male cyprids ovate; measuring 0.65 X 1.15 mm; pos-
from first tooth; third tooth equidistant between sec-
teroventral angle somewhat produced; 4 circular plates
ond tooth and inferior angle; inferior angle serrate;
appearing at posterior end of body (fig. 3 8 0 ) .
teeth either spatulate or triangular, from 13-23 in
Remarks: This species was first dredged by the Chal-
number (fig. 38A-D) ; one small isolated cluster of
lenger from 1097 meters off Mar del Plata, Argentina
setae occurring at base of the first tooth. Maxilla I
with deep notch slightly above the midpoint of the cut- (37"17'S, 53"52'W), but apparently has not been
taken since. The Eltanin records from shallower water
ting edge; 2 long, moderately stout and 2 short, thin
extend the southern limit of its range.
spines occur above the notch; 14-16 long slender
spines below notch (fig. 38E-6). Maxilla I1 ovotri- Arcoscalpellum parallelogramma is superficially re-
angular in outline; with 3 distinct clusters of marginal lated to those species with produced, wing-like carinal
setae; patch of short bristles occurring on the basal latera. However, it is readily distinguished from them
surface; maxillary lobe tall, erect, cylindrical and by the total lack of large discrete peduncular scales, the
apically truncate (fig. 38H). umbo of the inframedian latus free from the surface
Cirrus I separated from and about 2/3 length of of the capitulum and projecting basally, and the inter-
cirrus 11. Anterior ramus of cirrus I about y5 length digitating of the adjoining margins of the carinal
of posterior ramus; intermediate articles of anterior latera below the carina.
ramus protuberant, nearly twice the width of those of Arcoscalpellum recurvirostrum (Hoek, 1883)
the posterior ramus; inner lateral faces of both rami Text-fig. 39
densely clothed with long setae. Cirrus I1 normal.
Each articulation along the greater curvature of the Scalpellum recurvirostrum Hoek, 1883, p. 77, pl. 3,
intermediate articles of cirri 111-VI with 1 or 2 long figs. 11, 12, pl. 8, figs. 9, 10.
and 1 or 2 shorter setae; interarticular areas along Diagnosis: "Capitulum covered by membrane. Valves

umbo is at the apex, which slightly projects outwards,

and which has a triangular, recurved, and pointed
form.
"The tergum is triangular. The occludent margin
is much arched, hence the apex is distinctly recurved.
The inferior part is a little produced.
"The carina is bowed and internally concave; the
portion above the umbo is very short and not more
than one-sixth of the total length. Since the roof is
slightly convex, the sides of the carina pass over into
the portion above the umbo.
"The upper latus is trapeziform, with the scutal mar-
gin slightly hollowed out and not quite twice as long
as the carinal margin. The apex is sharply pointed.
"The rostrum is small, triangular, wedge-shaped.
"The rostra1 latus is convex, four-sided, with the
scutal and basal margins almost parallel and slightly
hollowed out.
"The infra-median latus is extremely small, triangu-
lar, with the umbo at the apex.
"The carinal latus is flat, four-sided; basal margin
nearly straight. Superior portion of the carinal margin
slightly hollowed out, inferior portion short, lateral
margin straight. The umbo is seated at the base of the
carina and projects slightly outwards.
Fig. 39. A r c ~ s c n l ~ e l l u recuruirostrum
m (Hoek) : A, "Length of the capitulum 13 mm.
individual viewed from right side; B, carinal view; "The peduncle is cylindrical, and has a length of
C, first maxilla; D, pedicel of cirrus VI and caudal 7.5 mm. Very small and not numerous calcareous
appendage (redrawn from Hoek, 1883, pl. 3, figs.
11, 12, and pl. 8, figs. 9, 10 [in part] 1. scales are scattered over its surface. At the place of
attachment the foot of the peduncle grows wider.
fourteen, separated by broad membranous interspaces. "Mouth.-Mandibles with three teeth, and the infer-
Umbo of the carina at a little distance from the apex, ior angle pectinated, almost as in Scalpellum inter-
hence the valve is slightly angularly bent. Upper latus medium. Maxillae with a rather deep notch behind the
trapeziform. Inframedian latus triangular. Peduncle first four spines, and a second less deep notch in front
cylindrical, with the calcareous scales scattered and at of the inferior angle. The second maxillae have the
a considerable distance from one another" (Hoek, so-called olfactory orifices at the end of long processes.
1883, p. 77). "Cirri.-First pair with very unequal branches; the
Material: Challenger, Sta. 150, between Heard Island anterior and shorter branch has six rather broad seg-
and Icerguelen Islands (52"04'S,71 22'E) ,274 meters: ments, the posterior eight longer and more slender
several specimens. segments. The outer surface of the segments is ex-
tremely hairy. Of the other cirri nothing very charac-
Original Description: "The capitulum is robust. about
teristic has been observed.
twice as long as broad, and not very flat. It is covered
by a thin membrane. and the different valves are sepa- "The caudal appendages are elongate and distinctly
rated by broad chitinous interspaces. A distinct white four-jointed; each segment bears two small spines at
line divides these interspaces into two equal parts, and the extremity; the fourth is crowned by a tuft of about
marks the area of the different valves. Of these there five stouter and two smaller bristles.
are fourteen. The lines of growth are not distinct. "Penis short, rudimentary.
"The scutum is elongate quadrangular; it is some- "Complemental males not observed" (Hoek, 1883,
what broader near its base than in the upper half. The p. 77-78).

S\1 STIZR.1ATIC ACCOUNT 81
Arcosrcrlpellum sp., c.f. A. s i n n n t ~ r m trated on margins of distal half; superior arid inferior
(Pilsbry, 1907) margins of proximal half of appendage free of setae.
Plate I X D, Text-fig. 40 Maritlible with 4 teeth inclutling inferior angle; teeth
Sralpellum sinualum Pilsbry. 190711. 1). 50, fig. 10; sharp arid widely spaced; second tooth closer to third
Uarnard, 1924, p. 40. tooth than to first; iriferior angle overhung by third
tooth; irlferior angle serrate, 7-8 long, spatulate teeth
D i a g ~ ~ o s i s :Rostrum minute, triangular i r ~ outline,
(fig. 40Ki. Maxilla I slender; cutting edge hith deep
fully exposed; Inframedian latus with IT-shaped sub-
and broad medial notch; 2 long, stout arid 2 short,
cuticular apical extension; roof of carina flat, with
stout spines above n0tc.h; 1 long, stout, 1 short, stout
prominent bordering ribs; caririal latera not interdigi-
and 1-2 short, slender spines below notch; 1 short,
tatirig helow basal margin of carina; maxilla I with
stout spine at side of notch (fig. 4 0 D ) . Maxilla I1
medial notch; intermetliate articles of cirrus V1 armed
short, triangular; marginal setae segregated into 3
with 3 pairs of setae; caudal appendage with 4 seg-
distinct sparse clusters; setae moderately long, stout;
ments, and about same height as pedicel of cirrus V l .
maxillary lobe cqlindlical, tall, broad. apex everlly
Material: Eltnnin, Sta. 18, 1,abrador Basin, off south- rounded ; surface free of setae (fig. 40E ) .
western tip of Greenland (52' lSrN, 4ti036'W), 3404- Antcrior ramus of cirrus I about I/r, length of pos-
3422 meters, 1 specimen. terior ramus, intermediate articles protuberant; seg-
Depository: USNM Cat. No. 125267. ments of posterior ramus orily slightly protuberant.
Description: Female-Measurements of the single Rami of cirrus I about l,!J length of rami of cirrus IS.
specimen: total height 7.0 mm, capitular height 5.6 Cirrus I1 normal; rami about y$ le~igthof rami of
mm, capitular width 2.8 mm. cirrus 111. Cirri 111-VI essentially equal in length,
Labrum bullate ; broad and moderately short ; mar- rami subequal; each articulation along greater curva-
gins and superior surface free of fine bristles; crest ture of intermediate articles with 1 or 2 long slender
armed with 20 conical teeth in a single row (fig. 4 0 A ) . setae and 1 or 2 extremely short, stiff bristles; inter-
Palps short, triangular; sparsely setose; setae concen- articular areas along greater curvature devoid of setae;
Fig. 40. Arcoscalprllu,nz sp. cf. A. sinuatum (Pilsbry) : A, portion of crest of labrum and right palp;
P,, mandible; C, lower two teeth and infcrior angle of mandible; D, 111axilla I; E, n~axilla11; F,
left cirrus I ; G, intermediate article of Cirrus VI; N, caudal appendage; I, vicw of right side of
fcrnale; J, rostrum. Elmnin, Sta. 18.

lateral face with axially aligned short setae; number the present specimen agrees in details w i ~ hthe holotype
of setae per article decreasing from cirrus I1 to cirrus of A. sinuatum with one exception-the upper latus.
VI. Setation ctenopod (fig. 40G) ; 2 major and 1 minor The sinuate or excavated basicarinal margin of the
pair on cirri 111-VI; 2-3 major and 1 minor pair on upper latus of A. sinuatum appears to be one of its
cirrus 11, 1 or 2 short setae at base of each major pair; more diagnostic features, but the present specimens
3 setae in each terminal cluster on outer ramus, only have not developed sufficiently for this feature. Bar-
2 in terminal cluster of inner ramus. nard has questioned whether or not the development of
Caudal appendage of 4-5 articles ; slender, tall, this plate is normal, but this question cannot be an-
slightly longer than pedicel of cirrus VI (fig. 40H) ; swered until additional collections become available
2 long and 3 short setae on terminal article; long seta for study.
about length of complete appendage. In comparing the arthropodal structures of the El-
Cirral counts for this specimen are as follows: tanin specimens with those A. s i n u a ~ u mdescribed by
Barnard from South Africa, certain similarities be-
came apparent. The segments of cirrus I in the present
specimens are also somewhat moniliform; the caudal
appendage is the same height and has the same num-
ber of segments; the mandible has 4 teeth including
the inferior angle which is slightly serrate; maxilla I1
is also ovotriangular; maxilla I has 4 spines above a
Remarks: All indications point to the present speci- shallow notch or indentation. On the other hand, the
mens as being a juvenile stage of A. sinuaium, figured intermediate articles of the posterior cirri are armed
by Pilsbry (1907b, p. 50). However, the differences with only 3 pairs of setae on each article rather than 4
between the juvenile and adult forms figured by Pils- as in the South African form. However, it should be
bry, and the absence of any description of the arthro- noted that Uarnard did not state which of his speci-
podal structures, does not permit an exact assignment mens were dissected, so that the problem remains.
of the Eltanin specimen. The species was initially de-
scribed from specimens taken at a depth of 3164 meters Arcoscalpellum sp.
off the coast of Maryland (3S053'N, 69'23'30"W). Plate VIII H, Text-fig. 41
The only subsequent record for A. sinuatum is that
of Barnard (1924, p. 40) from off South Africa, at Diagnosis: Roof of carina flat and bordered by ribs;
depths ranging from 1462-1828 meters. This southern rostrum small, equitriangular, with acute basal point
record is at the depth of the Maryland record for and umbo at apex, completely hidden by carinal
A. sinuatum arid the EDanin specimens from off Green- latera except for umbo; inferior angle of mandible
land. sparsely toothed; maxilla I with medial notch; distal
The rostrum of the present specimen is broad at the setae on intermediate articles of outer ramus of cir-
top, tapering to a point basally-that is, triangular. rus VI hypolasiopod, normal on inner ramus; cau-
The plate is fully exposed, and the adjoining latera are dal appendage with 4-5 segments; dwarf males sack-
hollowed out near the apices of the rostra1 margin. like, and with 4 calcareous plates.
The height of the plate is about 1/(? that of the adjoin- Material: Eltanin, Sta. 18, Labrador Basin, off south-
ing latera, as figured by I'ilsb~y ( 1 907b; fig. 16b). western end of Greenland (58' lSrN, 4S036'W) , 3404-
The specimens described by Rarnard were reported as 3422 meters, 4 specimens.
extremely variable, with regard to the development of
Ilepository: USNM Cat. No. 125268.
the rostrum, and it is possible Barnard was actually
confronted with more than onc species. Description: Female-RIeasurements (in mm) of the
The inframedian latus of the holotype of A. sinua- 4 specimens collectetl are as f o l l o ~ s :
tum, as figured by I'ilsbry, may be misleading. The
Total Capilular Capitular
caret-shaped line drawn in the middle of the valve is Sperimrn Heigll t Height Wicith
. -
quite likely the median umbo. The U-shaped portion ----------
-
1 17.7 12.6 7.0

of the plate above this line would then be interpreted 2 - 14.8 7.8
as a subcuticular apical extension, and as such the 3 10.6 8.3 4.2
u ~ n b owould be apical. If this is indeed the case, then 4 5.4 4.4 2.1

SYSTEMATIC ACCOUNT
Fig. 41. Arcoscalpellum sp.: A, labrum and palps; E, mandible; C, maxilla I; D, maxilla 11; E,
left rirrus I ; F, internledidtc article of outer ramus of rirrus VI; G, interrncdiate article of inner
ramus of cirrus VI; H, caudal appendage; I-L, lateral view of females; M, rostrum. Eltanin, Sta. 18.
Labrum bullate; broad and moderately short; su- short, slender spines below notch; long, stout spine of
perior surface and periphery free of bristles; crest not lower group longer than spines in superior group.
armed with teeth (fig. 41A). Palps triangular, moder- Maxilla I1 short, triangular; marginal setae segregated
ately short; superior margin and distal extremity with into 3 distinct but sparse clusters; maxillary lobe tubu-
short setae; inferior margin with 1-2 setae distally. lar, erect, extremely tall, apex flat; surface free of setae
Mandible with 4 teeth including inferior angle (fig. and bristles (fig. 41D).
416) ; teeth more or less equidistantly spaced; inferior Anterior ramus of cirrus I about y4length of poste-
angle bifid, with 1 or 2 extremely small teeth along rior ramus (fig. 41E) ; intermediate articles of ante-
inferior edge of major spine; superior margin of ap- rior ramus protuberant. Posterior ramus about y5
pendage free of setae and bristles. Maxilla I with width and about l h length of anterior ramus, and vz
broad notch in center or just above center of cutting length of posterior ramus of cirrus 11. Cirrus I1 nor-
edge (fig. 41C) ; 2 long, stout and 2 short, stout spines mal but rami grossly unequal, outer being about ??
above notch; 1 long, stout, 3-4 shorter, stout and 3-4 length of inner ramus. Cirri 111-VI essentially equal

in length, rami unequal. Each articulation along apparently lacks the bordering ribs present in A. in-
greater curvature of intermediate articles of cirri 111- cisum, A. atlanticum, and the present specimens. In
VI with 1 stout seta on outer ramus and 1 stout and 1 A. antarcticum there is evidently a rudimentary ros-
very slender shorter seta on inner ramus; interarticu- trum, but Hoek gave it nothing more than passing
lar areas along greater curvature bear 1-3 stout setae mention. Gruvel (1902a, p. 74) reported the presence
increasing in length distally. Inner lateral face with of a rostrum in A. atlanticum, stating that it is, "Ouale,
1 or more rows of axially arranged setae; 3 rows on u n peu recouvert late'ralement par les bords internes
cirrus 11, 2 rows on cirrus 111, 1 row on cirri IV-VI; des plaques rostro-late'rales. Absolument ii de'couvert
number of setae per row decreases from cirrus IV to dans ses autres parties, et, par conse'quent, distincte-
cirrus VI. Setation ctenopod (fig. 41G) ; 4 pairs (2-3 ment visible ii la surface." Aurivillius (1898, p. 194)
major pairs and 2-3 minor pairs) ; distalmost cluster states that in A. incisum the "Rostrum superficie mi-
of setae of each article on outer ramus hypolasiopod, nuta triangulari (lateribus rostrolateralium verisimi-
with 3 4 setae of various lengths (fig. 41F). liter occulta) ." What the rostrum looks like, and its
Cirral counts : relation to the adjoining plates in A. antarcticum re-
mains to be determined. Although the plate is small
and triangular in the present specimens, it has an acute
Specimen 1
-
8
-
15 19
-
xx
- -
xx
-
xx 5 basal point rather than an acute apical point as illus-
11 18 21 22 23 23 trated by Gruvel (1920, pl. 6, fig. l l c ) for A . incisum.
In both of these species the umbo is apical, but in the
Greenland specimens the umbo is situated in the cen-
ter of the apical margin and only it is exposed. In the
Specimen 2 7 13 17 xx 19 xx 4 drawing of A. incisum presented by Gruvel, the ros-
- - - - - -
10 xx 19 xx xx 22 trum is clearly and totally exposed, although there is
no indication of this from the photographs presented
(Gruvel, 1920, pl. 2, fig. 3 ) .
One additional point of difference among the mem-
Caudal appendage with 4 5 segments (fig. 41H) ;
bers of the present complex of species should be men-
appendage approximately same height as first article
tioned, namely the manner in which the carinolateral
of pedicel of cirrus VI; 6-8 terminal setae, long-
plates abut below the basal margin of the carina. In
est approximately $5 length of appendage; setae with
the present specimens, the basal margins of the plates
setules.
are parallel, above which they form the typical V. In
Nauplii present in mantle cavity of specimen 2 ;
the illustrations of A . incisum Gruvel figures the rela-
obovate in outline, ranging from 0.65 X 0.85 mm to
tionship of the plates as a simple V, as does Hoek for
0.75 X 0.90 mm; frontolateral horns short and stout;
A. antarcticum.
appendages short, with long, soft, natatory setae.
The present form cannot be readily assigned to any
Dwarf male sack-like, oval (0.45 x 1.0 mm) ; sur-
of the aforementioned species, but perhaps this is only
face covered with concentric bands of extremely small
because they were inadequately described. By the same
spines; 4 calcareous plates present at one pole; testes
token, it cannot be determined whether the present
large and V-shaped.
form is sufficiently distinct to be proposed as a new
Remarks: The present form is closely related to
species. Therefore, we feel it better to leave the Eltanin
A. antarcticzcm (Hoek, 1883, p. 95), A . atlanticum specimen unnamed until the previously named species
(Gruvel, 1902a, p. 7 4 ) , and A . incisum (Aurivillius, are redescribed.
1898, p. 194) . Arcoscalpellum antarcticum was first
dredged off Ingrid Christensen Coast (65'425, Arcoscalpellum triangulare (Hoek, 1883)
7g049'E) from 3062 meters, A. atlanticum from 1187 Text-fig. 42
meters, south of Ilha de SBo Miguel, Azores (37'37'N,
25O20'45"W), and A. incisum from 1022 meters, Scalpellum triangulare Hoek, 1883, p. 130, pl. 3,
north of Ilha Graciosa, Azores (38047'40"N7 28'17' figs. 17, 18, pl. 11, fig. 4 ; Zevina, 1964, p. 253.
05''W). The Eltanin specimens are from much farther Diagnosis: "Valves thirteen; surface of the valves
north and markedly deeper water. covered by a rather thick chitinous membrane, fur-
The roof of the carina in A. antarcticum is flat, and nished with short woolly hairs. Carina simply bowed,

shaped. Striae much more distinct, and separated by

broader interspaces than on the other valves. The
lines of growth are oblique. The flattened roof of the
carina in most other species is totally wanting in the
present species.
"The upper latus is quadrilateral, or rather, trapezi-
form, the scutal margin being much longer than the
carinal margin; the latter is straight, the former is
distinctly hollowed out, with the angles at the upper
and under extremities very sharp.
"The rostral latus is nearly twice as long as broad,
widening considerably from the rostral extremity to
the opposite end.
"The infra-median latus is small. triangular, with
the umbo at the apex.
"The carinal latus is large, greater in area than the
rostral and infra-median latera together. Basal margin
straight, carinal margin very concave in the superior
half; the umbo of the carinal latus is placed almost in
Fig. 42. Arcoscalpellum triangulare (Hoek) : A,
individual viewed from right side; B, carinal view the middle of the carinal margin; inferior half of the
(redrawn from Hoek, 1883, pl. 3, figs. 17, 1 8 ) . carinal margin very convex. Upper margin straight,
lateral margin also straight, making with the upper
with the umbo at the apex; boat-shaped, without a margin an angle a little greater than 90".
flattened roof. Upper latus quadrangular, trapeziform. "Length of the capitulum about 9 mm, of the pedun-
Umbo of the carinal latus in the middle of the carinal cle 3.5 mm.
margin. Peduncle short. Males, one on each side" "The peduncle is short, with the scales in about
(Hoek, 1883, p. 130-131). seven longitudinal rows covered by the chitinous mem-
brane, each row containing transversely about eight
Material: Challenger, Sta. 320, type-locality, east of
scales.
Mar del Plata, Argentina (37" 1 7 5 , 53"5ZtW), 1097
"Mouth.-The labrum is angularly bent in the mid-
meters, 4 specimens.
dle, the angle being directed towards the adductor mus-
Original Description: "The capitulum is very thick cle of the scutum. The margin of the labrum is furnished
and robust, especially near the peduncle, growing flat- with extremely small teeth. The palpi are triangular
ter towards the apex. The margins of the valves and and small, with a few hairs only at the extremity. The
the structure of the surface of the valves cannot be mandibles have three nearly equal and equi-distant
easily made out, as they are covered by a rather thick teeth; the inferior angle is bluntly truncated, and bears
chitinous membrane, which is woolly, with short hairs small and numerous short spines or teeth. The maxil-
at the surface. Valves distinctly striated when the lae have a simple edge with numerous spines; a very
chitin is taken away. small notch is visible a little behind the middle. The
"The scutum is broad, its breadth being almost two- upper spine is large, much larger than the second one.
thirds of its length, convex, quadrilateral, with the The part above the notch forms a distinct projection
occludent margin slightly arched. Tergal margin hol- over the lower part. Second maxillae only slightly
lowed out, lateral margin arched, basal margin almost developed, covered almost over its whole surface with
straight. The umbo is at the apex, which projects con- numerous slender hairs.
siderably over the tergum. "Cirri.-First pair short, with unequal rami; the
"The tergum is triangular, flat, with the inferior longest ramus has seven, the shortest five segments.
part produced and the occludent margin slightly con- The other cirri in the specimen I have dissected have
vex. Its area is a little greater than that of the scutum. at least one of the rami truncated, and the truncated
"The carina is boat-shaped, narrower at the upper extremity swollen in a curious way; it looks almost as
than at the lower extremity. Sides only developed at if each ramus was furnished at its extremity with a
the superior half of the valve, and here semilunar number of globular vesicles.

'6
Caudal appendages rudimentary, of one segment dible with 5 teeth, as did Broch (1953, fig. 4d) in
only. A. vitreum. The bathymetric occurrence of Utinomi's
"The specimens are attached to a Sclerodermous specimens also argues against inclusion in A. vitreum.
Zoantharian (Flabellum? ) The shallowest occurrence of A. vitreum is 3192 me-
"A small complemental male [dwarf male] is pres- ters whereas the deepest record for Utinomi's speci-
ent at both sides, attached near the occludent margin mens is 230 meters.
of the scutum. It has a length of 0.88 mm, and shows The specimens Weltner (1922, p. 95) briefly de-
a well-developed testis with a receptaculuin seminis. At scribes are tentatively referred to this species solely
the anterior extremity it is furnished ~ i t hwell-de- on the basis of locality and bathymetry.
veloped antennae; at the other extremity a circular
space is to be distinguished, enclosed by a thickening Arcoscalpellum ventricosum (Hoek, 1907)
of the chitinous wall of the body. Text-fig. 43
"Among the eggs, which entirely fill the cavity of Scalpellam arcuatum Hoek, 1907b, p. 98, pl. 8, figs.
the capitulum, I observed a larva in the Cypris-stage. 3-3a (=Scalpellum ventricosum Hoek, 1907b, p. 275,
The ova ere fecundated, and happened to be in one replacement name), non Scalpellum arcuatum Darwin,
of the later cleavage-stages" (Hoek, 1883, pp. 130- 1851~1,p. 40; Nilsson-Cantell, 1921, p. 205, fig. 32.
132).
Diagnosis: "Valves fourteen. Carina simply and
Arcoscalpellum utinomii sp. nov. feebly bowed, with a flat roof bordered by ridges.
Upper latus large with a short carinal and long, ir-
Scalpellum formosum?: Weltner, 1922, p. 95.
regularly bowed scutal margin. Rostrum very small,
Scalpellum vitreum: Utinomi, 1958, p. 283, text-figs.
1 0 triangular.
- Inframedian latus narrow, with the umbo
at the apex. Carinal latus with the umbo not far from
Diagnosis: Capitular plates ornamented with strong the base" (Hoek, 1907b, p. 9 8 ) .
radial ridges; carinal margins of carinal latera smooth
and not interdigitating; rostrum small, diamond- Material: Siboga, Sta. 295, type-locality, Timor Sea
shaped, wholly exposed; anterior ramus of cirrus I (10°35.6'S, 124"11.7'E'I, 2050 meters, 1specimen.
shorter than posterior ramus; caudal appendage sup- Original Description: "The capitulum is flat with the
porting 2 setae; mandible without small teeth on upper carinal side slightly and the occludent side strongly
slope of second and third teeth; maxilla I with medial
notch in cutting edge; maxilla IS ovate with a basal
notch; dwarf male sack-like without calcareous plates.
Type-locality : Amadaiba, Sagami-wan, Japan;
July 16, 1934; 128 meters.
Depository: l3iological Laboratory of the Imperial
Household, Crust. No. 138.
Remarks: Utiriomi (1958, p. 286) described several
specimens of an arcoscalpellid which he believed ". . .
may be referable to Scalpellum oitreum Hoek . . . ."
However, there are several outstanding differences be-
tween Utinomi's specimens and A. vitreum. Foremost
is the development of the carinal latera, which do not
interdigitate helow the basal margin of the carina. In
both A. vi~reumand A. formosum, these plates inter-
digitate. In addition, the rostrum in these two species
is higher than broad and either elongate-oval or rec-
tangular, whereas in Utinomi's specimens the rostrum
figured is diamond-shaped, in good part as he states, Fig. 43. Arcoscalpellum ventricosum (Hoek) : A,
individual viewed from left side; B, view of rostrum
(6
. . . roughly quadrangular or lanceolate." The man- and surrounding plates (rostrolaterals and rostra1
dible in both of Hoek's species has 4 teeth including angles of scuta) (redrawn from Hoek, 19017, pl. 8,
the inferior angle whereas Utinomi pictures the man- fig. 3-3a).

arched. It is covered by a thin membrane, which bears but especially because the valves of the lower whorl
short hairs along the carina, forming tufts at the base are much more developed in Sc. gracile than is the
of that valve and in a less degree at the apex. The case in the present species. The carinal margin of the
carina is separated from the other valves by a distinct carinal latus is moreover slightly more hollowed out
chitinous interspace. The valves of the lower whorl in Sc. arcuatum than in Sc. gracile" (Hoek, 1907b,
are less than medium-sized. p. 99).
"The scutum is quadrilateral and more than twice
Arcoscalpellum vitreum (Hoek, 1883)
as long as broad. It has an arched occludent margin;
Plate VIII E, F; Text-figs. 44-47
its tergal margin is short; its lateral margin slightly
hollowed out in the upper part and strongly convex in Scalpellurn vitreum Hoek, 1883, p. 115, pl. 5, fig.
the lower. The apex is pointed, but not produced. 14; Weltner, 1897, p. 251; Gruvel, 1902a, p. 54;
Growth ridges indicated, but not distinct. 1905, p. 84, fig. 94; Pilsbry, 1907b, p. 60; Nilsson-
"The tergum is large, triangular, with the occludent Cantell, 1955, p. 219; Tarasov and Zevina, 1957,
margin straight. It meets the carinal margin in a very p. 142.
slightly produced little point. The carinal margin is Scalpellum talismani Gruvel, 1902a, p. 86, pl. 2,
straight in the upper, feebly convex in the underpart. figs. 3D, 6 , 7; 1905, p. 86, fig. 96; 1920, p. 23;
Ridges of growth very indistinct. Nilsson-Cantell, 1955, p. 219; Broch, 1953, p. 8, fig. 4.
"The carina is simply and not strongly bowed. The Scalpellum formosurn Pilsbry, 1907b, p. 58, fig. 22;
roof is flat, increasing in width from the apex down- Gruvel, 1920, p. 23; Stubbings, 1936, p. 55, in part;
wards, bordered by ridges. The lateral parts rather Nilsson-Cantell, 1938, p. 21; 1955, p. 219.
narrow and having about the same width over the non Scalpellurn forrnosurn Hoek, 1907b, p. 110, pl. 8,
whole length. figs. 11, l l a .
"The upper latus is large and has a somewhat ir- Scalpellurn bellurn Pilshry, 1908, p. 111 (replace-
regular shape. Its umbo is at the apex which pene- ment name for Scalpellurn formosum Pilsbry, 1907b).
trates into the corner between the scutum and the ter- Diagnosis: Capitular plates ornamented with weak
gum. It has a long scutal margin, slightly convex in radial ridges; rostrum small, narrow, about 1/2 height
the upper and strongly hollowed out in the under part. of and covered by rostral latera in adult, but same
The carinal margin is short; the lines of growth are height as rostral latera in juveniles and always very
rather indistinct. narrow; inframedian latus about 112 height of margins
"The rostrum is very small and triangular; it is of adjoining plates; carinal margins of carinal latera
placed in the corner between the tips of the rostral interdigitating below carina; anterior ramus of cir-
latera and the scuta.
"The rostral latus quadrilateral, rather small, as is
also the case with the other valves of the lower whorl.
"The infra-median latus very narrow, with the umbo
at the apex, very slightly increasing in width down-
wards.
"The carinal latus has the umbo near the base. It
can hardly be said to project beyond the line of the
carina. The upper part is triangular and fits into the
angle between the carina and carinal latus.
"The peduncle is short, broader in the upper part.
The scales are broad and narrow; they are irregularly
distributed over the surface.
"Size. The total length of the animal is 9 mm., that
of the capitulum 7.5 mm.
"This deep-sea form comes nearest to the species I
describe on p. 105 as Sc. gracile. I consider the two Fig. 44. Arcoscalpellum vitreum ( H o e k ) : A, B, left
forms as different not so much because a rostrum is and right sides, respectively, of same female specimen.
present in the one and wanting in the other species, Eltunin, Sta. 18.

ANTAKCTIC CIRRIPEDIA
I'ia. 45. Arcoscalpe/lun~vitl-eum (1Ioek) : A, view of right side of lemale; B, cxtrrrlal view of rostrum; C,
mandible; D, enlarged view of thirti tooth and inferior angle of s~urre mandible; E, lahrum and palps;
I", portion of crest of labmm; (;-I, maxillae I ; J, maxilla I I ; K, mandiblc; L, rnlirrged view of inferior
angle of sanlc mantlihle; M, infr~imedianlutus; N, view of abutment of carinal latc~raand basal portion of
~ . a r i n a . I:'/IUILI'IL,
Sta. 18.
rus I shorter than posterior ramus; caudal i~ppmdape cific Basirl off Hrlarl (Ioast, l<llsuorth Land (6305JfS,
uriiarticulatc, and clothed with 15-18 setire ; nlarldible :;3 " O3'W), 1531 meters, 1 specimen ; E a s t ~ m n l ,Sta.
~ i t h o u small
t teeth on upper slope of secol~(ia11(1third 7617, Hatteras Plain, east of Charlcstos, South Caro-
major teeth, maxilla I with medial 1idc:h be lo\^ which lina (33°5:i.7fN, 75"42'W), 2280 meters, 1 specimen.
are 8-10 spines; maxilla 11 triangular arid trilol~aie; it^^^: USNM cat. N,,. 125269 (sta. 18) ;
dwarf male sack-like, with calcareous plates. 125270 (Sta. 791).
Material: Eltanin, Sta. 18, Labrador Basin off south- Supplementary Descriplion: Female-Capitulum con-
ern tip o l Greenland (5801SrN, 4E03h'W), 340143422 sists of fully calcified, approximate plates; large spcci-
meters, 4 spec:irnens; Sta. 791, south of Southeast Pa- mens with membranous space between tergum and

SYSTEMATIC ACCOIJNT 89
upper latus arltl c.arina; capitulum elongate and oval; or slightly convex; umbo apical, slightly produced,
(:overed with moderately thick hirsute cuticle; occlu- and projecting over the basioccludent corner of the
(lent and carinal margiris convex; shell widest metli- tergum; a rihlet more strorigly developed than the
ally ; plates orliamerited with unequally spaced growth others extends c u r v i l i n e a ~ l from
~ apex to basilateral
ridges, and low, narrow riblets radiating from region angle. Cariria bowed; roof flat, hounded laterally by
of umbo. Tergum triangular; acumirlate at extremi- low to relatively high, narrow to broad ridges; sides
ties; basal and occlutlent margins about 'l,; length o f ~ i t hscalloped margins, strongly sculptured with ir-
caririal margin. Scutum quadrilateral; about twice as regular, noncorltinuous, oblique ribs. Upper latus es-
high as broad; occludent margin corivex; basal and sentially triangular, hut nith 4 or 5 siclrs; umbo api-
tergnl margins slightly concave; lateral margirl straight cal; scutal anti telgal margins straight or slightly con-
Fig. 46. Arcoscalpell~irnvztreum (Hoek) : A, inlrametlian latus; U , lahrum and palps; C, D, enlarged
portions of crcst of ldblunl; L, ~nandlhle; F, enlarged view of inferior anglc of samc mandible;
G, 13, maxillae I ; I, maxilla 11; J, intermediate article of cirrus V I ; K, cautlal appendage. Eltanir~,
Sta. 791.

Fig, 47. Artoscalpellurn v i ~ r r u m (Hock) : A, R, left cirri J; C, D, caudal appcndagrs; E, irltcrmediate

article of cirrus VI. Eltanin, Sta. 18 (figs. A, C-E), and Sta. 791.
vex; tergal and basal margins of equal length; scutal between second and third tooth; inferior angle pecti-
slightly longer. Carinal latus largest of the latera; nate, 11-13 spines on superior margin and tip, none
5-sided, but irregularly quadrate; umbo on carinal along inferior edge (figs. 45D, L; 46F). Cutting edge
margin about 36 of the length of the plate from the of maxilla I relatively straight, notch variable in de-
inferior margin; umhorial region elevated above level gr ee of prominence (figs. 45G, H; 46G. H) ; 2-3 long,
of plate; latera extending behind and below carina; stout, 1 short, stout arid rarely 1 short, thin spine
interdigitatirig below carina. lnframedian latus in above notch; 9-12 long or short and slender spines
form of isosceles triangle (figs. 45M, 46A) ; height- below notch; superior and inferior. Maxilla I1 tri-
width ratio varying with age; width of base ranging angular and trilobate in outline; marginal setae ar-
from more than 14 to about %?( height of plate; height ranged in 3 distinct clusters; maxillary lobe moder-
always greater than width; plate shorter than adjoin- ately short, broad, and apically rounded.
ing latera. Rostra1 latus quadrate; nearly as wide as Cirrus I separated from cirrus 11; y5 to z$<its
high; width of basal margin less than lh scutal mar- length; cirrus I1 not greatly modified. Anterior ramus
gin; occluderlt margin of latera contiguous; apical l , { of cirrus I slightly shorter than posterior ramus (fig.
of lateral margin contiguous with lateral margin of 47A, B) ; intermediate articles of posterior ramus
carinal latus; umho at apex of scuto-occludent angle. about % width anterior ramus; inner lateral face of
Rostrum marrow, rlongate and oval; totally hidden 1 ) ~ hoth rami densely clothed nith long setae. Proximal
rostral latera; plate 1;; to height of occludent mar- 2 or 3 articles of cirri 11-VI either totally or partially
gin of rostral latera; umho apical. fused; articulations along greater curvature of inter-
Peduncle short, allout 'I/;; or less height of capitulum;
mediate articles of cirri 111-VI with 1 or 2 long and
peduncular scales 1)road and ahout 4 in each row;
1 or 2 shorter setae, interartic.ular areas along greater
exposed portions of scales convex.
curvature generally free of statar. Setation ctenopod;
1,abrurn bullate (f~gs.45E, 4613) ; free of setae and
.I-5 pairs on cirri 111-VI; a few, unpaired, very short
bristles ; crest '11 med w ilh 40-45 triangular teeth ;
teeth occnrrir~gin one row in central portion of crest, setae at bases and between bases of major setae.
but separating into 2 rows at lateral extremities. k'alps Caudal appendage extreme1y small; uniarticulate ;
narrowly triangular ; moderately setose; setae a n ayed about 1h to 1h3 height of first segment of pedicel
along Loth margins and face of appendage. Mantlihle of cirrus VI; numerous setae present on anterior and
with 1 teeth irlcludirig inferior angle (figs. 45C, 'JOE) ; superior margins; setae of superior margin nearly as
distan~ebetween first and second tooth greatrr than long or longer than appendage.

Measurements of the 5 specimens: Gruvel (1920, p. 23) drew attention to the fact
that the validity of A . bellr~rn(=A. jormosum Pilsbry)
Total Capitular (:apitular
S~x.c~inrcnHcight Height Widtl~ might at some time he questioned when he stated
" . . . Sc. I'ali~rnar~i
- - -
A. G'ruvel, forme, d u reste, voisir~e
Eltanin, Station 18 1 - 18.6 10.3
et de Laquelle se rupproche beaucoup, si mgme e l k
2 16.0 14.2 7.5
3 8.7 7.0 3.2 n'e.5~pas irientique le Sc. formosum cie Pilsbry." Broch
4, 9.6 8.6 4.0 ( 1953, p. 8 ) synonymized A. bellurn with A. ialismani
stating, "lt is impossible to trace differences of taxon-
omic value in the drawings arid descriptions given
Enstwurd, Station 7617 1 - 18.3 9.1 1)y Gruvel (1902) and Pilsbry (1907)." We concur
with Rroch's decision to synonymize these two species,
Cirral counts for the specimens are as follows:
which in turn we consider synonymous with A.
vilreum.
There are no differences in the morphology of the
Eltanin, Station 18
shell by which A. ialisrnani and A. vitreum can be
Specimen 1 10 20 xx xx 27 27 1
- -- - - - - separated. The only difference noted between the
11 19 xx 24 24 24 present arctic and antarctic specimens, and the speci-
men of A. talismani described and illustrated by Broch
11953, p. 8 ) , is an additional, spurious tooth in the
m~ndibleillustrated. The cirral counts and chaetotaxy
that Broch presented are remarkably similar to those
of the present specimens. No mention or description
is given of the caudal appendage. The bathymetry
of the present specimens is in accord with the pub-
Eltanin, Station 791 lished records of A. vitreurn, A . bellurn. and A. talis-
Specimen1 10 19 22 26 27 27 1 mani; the specimens from arctic waters occurring at
- - .
- - - -
34013422 meters, those from antarctic waters at
12 18 20 23 27 24
4531 meters.
9 18 20 25 23 26 1
- - - - - - The only complete description of A. vitreum was
12 19 20 21 23 28
given by Utinomi (1958, p. 283), but it does not
Eastward, Station 7617 agree well with the illustratio~isand description given
Specimen 1 8 18 21 23 24 27 1 by Hoek. Among the numerous differences, the strong
- - - - - - -
11 17 20 23 24 25 radial sculpture of the capitular plates, the diamond
rather than oval shape of the rostrum, the height of
the inframedian latus being in excess of the rostra1
latus, the deep furrow in the roof of the carina, and
Dwarf male sack-like, elongate and oval (1.05 x 1.25 tFe lack of interdigitations along the carinal margins
mm) ; sack covered with concentric bands of short 01 the carinal latera are very important. Another
spines; 4 calcareous plates. paint is the bathymetry; Utinomi's specimens were
taken at a depth of less than 250 meters whereas
Remarks: The description of A . vitreum by Hoek
A . vitreum occurs in water deeper than 3000 meters
(1883, p. 115) was based on a single specimen col-
lected by the Challenger off Tokyo, Japan (34"37'N, (see A. uiinomii sp. nov. herein).
140032'E) at a depth of 311.29 meters, but there was Arcoscalpellum vitreurn is closely allied to A.
no mention of the arthropodal structures. Four speci- formosum Hoek, from which it be readily sepa-
mens in the present collections dredged off southern rated on the basis of the height of the inframedian
Greenland (Sta. 18) were initially identified as A. l a t the
~ absence of teeth on the upper slope of the
talismani Gruvel, and upon subsequent second and third major teeth, and by the trilobate
proved to be identical with a specimen dredged off maxilla 11. Included in this complex of species are
Ellsworth Land (Sta. 791), which had previously been A . curvatum (Gruvel, 1900b, p. 192) and A. rigidum
considered A. vitreum. (Aurivillius, lS98, 11. 190).

Arcoscnlpel2urn weltneri (Gruvel, 1907) curvature with 1 long seta and 1 extremely s h o ~ t ,
Plate VIII B, Text-fig. 48 stiff bristle; interarticular areas along greater curva-
Scalpellum weltneri Gruvel, 1907b, p. 159; 1909, p. ture free of setae and bristles. as are lateral faces.
205, ~ 123, . figs. 4-5, pl. 26, figs. 8-11. Setation ctenopod; 2 major and 1 minor pair of
setae; at bases of major pairs 1 or 2 short slender
Lliagnosis: Capitular plates separated by narrow,
setae (fig. 48H).
translucent, chitinous interspaces; tergum inclined,
Caudal appendage uniarticulate; laterally com-
length 3 times greatest width; roof of carina without
pressed; about length of first segment of pedicel
medial sulcus ; irif ramediarl latus triangular with fan-
of cirrus V1; anterior and posterior borders of ap-
shaped, subcuticular, apical extension; rostrum when
pendage clothed with extremely short, stiff bristles; no
partially exposed with exposed portion triangular
terminal long setae present.
in outline, but when fully exposed square to slightly
Of the 7 specimens collected only 5 were sufficiently
rectangular, umbo apical; labrum covered completely
well preserved to prokide measurements (in mm) :
with ctenae; mandible with 4 teeth and receding in-
ferior angle; maxilla I with medial notch; maxilla 11 Total Capitular
ovotriangular, but not trilobate; caudal appendage Specimen Hcight Height
--
uniarticulate; dwarf males sack-like, lacking calcare- 1 7.5 5.2
ous plates. 2 7.3 4.9
3 6.6 4.7
Muter i d : E l ~ c n i n , Sta. 10E1, north of Coronation 4 4.4 3.2
Island, South Orkney Islands (60°22'S, 46O5OfW), 5 4.1 2.9
29U403 meters, 7 specimens.
Specimens 1 and 2 above were dissected; counts or
Depository: USNM Cat. No. 125271. the cirri are:
S ~ p p l c r n e n t a r i~l e s c iplion:
~ Female - Labrum bul-
late; short, broad, rounded distally; superior surface
and margins covered with clusters of 3-6 short, fine Specimen I 6 9+ 13 13 14 14 1
- -
bristles; bristles directed towards crest; crest armed 8 13 xx 13 14 14
with about 20-25 very low, broad, M-shaped or comb- 7 10 13 14 14 14 1
like teeth (fig. 48A, B) . Palps moderately long, tri- - - - - - -
8 13 13 14 xx 14
angular; distally acute; superior and proximal inferior
margirls covered with clusters of short, fine bristles
similar to those of the labrum; l o ~ e margin r and ter-
minal portion of appendage armed with stout setae.
Mandihle with 5 teeth including inferior angle (fig.
48C, D ) ; sec,ond tooth well separated irom first. Max-
illa I ~ i t hdetl), promirlerlt notch above center (fig. Dwarf male sack-like; rlongate-oval (0.25 X 0.75 to
4:E) ; above notch 2 long stout and 2 short stout 0.35 x 0.65 m m ) , covered with concentric bands of
spines; helow 11otc.h 5-7 stout spines, 4-5 of which are fine spines; lacking calcareous plates.
long and 1-2 short; superior and inferior margins of Remarks: The present species was first described
appendage clothetl with clusters of long, slender irom off Gaussberg on Wilhelm 11 Coast (approxi-
bristles. Maxilla TI ovotriangular; marginal setae mately 66"S, Z9"E). It was dredged several times by
segregated into 3 clusters; cluster at distal tip brar- the Causs from depths of 350 to 385 meters. The
ing heavirst (,oncentration of setae; maxillary lo1)c.s Eltur~inrecord from a comparable depth extends the
very low, hroad. apex broadly rounded; surface free range of A. weltneri to the other side of Antarctica.
of setae (fig. 48F). To judge from the illustrations of A. wcl~neriand
Anterior ramus of cirrus I about q5length of pos- A. bouvieri (Gruvel, 1900, p. 272) presented by
terior ramus (fig. 48G) ; intermediate articles of Gruvel (1907b, 1909) and subsequent workers, it
posterior ramus slightly narrower than those of might appear that they are one and the same species,
anterior ramus. Rami of cirrus I about 1 h length of and the morphological similarities between them have
rami of cirrus 11. Cirrus I1 normal. Cirri 11-VI been touched upon by Nilsson-Cantell (1930b, p. 244),
esseritially equal in length with rami subequal; each who maintained their distinctiveness. The only notice-
articulatior~ of intermediate articles along greater able difference, as Nilsson-Cantell mentioned, is in

SYSTEMATIC ACCOUNT
Fig. 48. Arcos~al~ellurnweltneri (Gruvel) : A, labrum and right palp; B, portion of crest of labrum en-
larged; C, D, mandibles; E, maxilla I ; F, maxilla 11; G, cirrus I ; H, intermediate article of cirrus VI ;
I-M, view of right side of females. Eltanin, Sta. 1084.
the inframedian latus being keyhole-shaped in A. differences. The slight variations in capitular mor-
zueltneri, and hourglass-shaped in A. bouvieri, with phology are insufficient to settle the matter, but look-
the apical extension flaring to a greater or lesser ing to certain arthropodal structures clearcut differ-
extent. ences can be discerned: the number of teeth in the
In the specimen of A. boz~vierifigured by Nilsson- mandible, excluding the inferior angle, is 4 in A.
Cantell (1926, text-fig. 1F) the inframedian latus is, weltneri but only 3 in A. bouvieri; the anterior cur-
point for point, the same as that figured here for vature of the intermediate articles of the posterior cirri
A. weltneri (pl. VIII B, text-fig. 48K-M), but there support 3 pairs of setae in A. weltneri, but 4 in A.
is some difference in the relative proportions of bouvieri. These generally reliable differences taken
the scutum which is not obvious from a study of together should provide a means for discriminating
Gruvel's figures. Similarities in arthropodal structures between these two closely related species.
can be detected also, especially in maxillae I and 11, KEY TO GENERAALLIEDTO Neoscalpellum
and, taking into account the slightly larger size of
1. Basal margin of scutum with notch ................. 2
NilssOn-Cantell's the cirral counts and the 1. ~~~~l margin of scutum without notch ................ 3
progression in number of segments from cirrus I to 2. Inframedian latus higher than wide, umbo medial.. ....
cirrus VI amear com~arable. .............................. Neoscalpellum Pilsbry
1 A
Although Nilsson-Cantell emphatically professed the

2. Inframedian latus equilateral, umbo apical ..........
.........................Gymnoscalpellum gen. nov.
of these two based up0n a of 2. Inframedian latus wider than high, umbo basal .......
the types, he did not clearly set forth in detail the .........................Abathescalpellum gen. nov.

ANTARCTIC CIRItIPEDIA
A Abafhescolpel/um koreonum ( Hiro) A Neoscalpellum debi/e (Aurivillius)

o Gymnoscolpellum farasovi gen, el sp. nov. Neoscolpellum phonlasma (Pilsbry)
Gymnoscolpellum insigne (Hoek) Neoscalpellum sch~zoplacinumsp. nov
Gymnoscalpe//um lorvale (Pi lsbry) Neoscolpellum elfaninae s p, nov.
0 Gymnoscalpellurn leoni (Zevina) Neoscolpellum marginofum (Hoek)
Chart 6. Distribution of thc rnonirtypic genus Abuthescalprll~~mgen. nov., Gyn~r~osccrlpellrrrngrn. no\. , ~ n d Nroscnlprllu~n.
T h e t r a n s p o l a ~ connectton between tlie species of N~~oscalpe1lurnis suggestetl b c u u s e t l ~ e vale abyssal form-.

SYSTEMATIC ACCOUNT
Litoscolpellum f/ssicorinotum sp, nov. A Mesoscolpellumjovonicum ( H oek )

+ Litoscolpellum simplex sp, nov. o Mesoscolpellum imperfectum (PIIs bry)
A Liloscolpellum wolleni sp, nov. 0 Mesoscolpellum sonctoeborboroe (Pilsbry)
Litoscolpellum discoveryi (GruveI ) Mesoscolpellum cochleorium ( H iro)
8 Litoscolpellum convexum (Nilsson-CantelI )
+ Litoscolpellum aurorae (Bage)
+ Litoscolpellum intermedium (Hoek)
8 Litoscolpe/lurn nipponense (Pi lsbr y)
Litoscolpellum loccodivicum (Annanda le )
Chart 7. Distribution of Litosculprllum gen. nov. and Mesoscalpellr~m. The conncrtion between species of Mesoscalpellurn
across Ce~ntralAmerica is suggested because M . imperfectum occurs in the eastern I'acific (near Galapagos Islands) as well as
the Atlantic. However, likc Neoscalpellum (Chart 5), these are little-known abyssal forms.

3. Inframedian latus triangular, umbo apical .......... groups are not only readily definable taxonomic units
............................Litoscalpellum gen. nov. but that they constitute biogeographical entities.
3. Inframedian latus rectangular or hourglass-shaped, umbo
submedial ....................Mesoscalpellum Hoek
3. Inframedian latus vase-shaped, umbo submedial to basal
Neoscalpellum debile (Aurivillius, 1898)
.............................Annandaleum gen. nov. Text-figs. 49, 50
Genus Neoscalpellum Pilsbry, 1907 Scalpellum debile Aurivillius, 1898, p. 189; Gruvel,
1905, p. 27; 1920, p. 31, pl. 5, figs. 13-15, pl. 7, fig.
Diagnosis: Capitulum of adult female with 14 re- 1 ; ~,11,,, 1959, 2.
duced calcareous plate; tergum in shape of inverted Scalpellum edwar&si Gruvel, 1902a, p. 63, pl. 2,
V; scutum with basal margin notched, with long figs. 3 ~ 16;
, 1905, 28, fig. 27.
a~icOlateralarm % to % length of tergal margin; Scalpellum (Neosca~pe~lum)dicheloplax Pilsbry,
upper latus bifid, with long slender depending arm, 1907b, p. 70, figs. 28a-c; Hock, 1914, p. 4; Gruvel,
umbo apical; carinal latus higher than wide, common- 1920, p. 32, pl. 7, fig. 2; withers, 1926, p. 102.
'Y forked with Or submedia' umbo; ScnlpellUm ( N e o s c a ~ p e ~ ~ ndicheloplax
m) benthophila
inframedian latus slender, higher than wide, com- pilsbry, 1907b, 73, fig. 28d: cruvel, 1920, 7,
monly forked apically, with medial or submedial fig. 9.
umbo; rostral latus higher than wide: with notched Scalpellum alboranense Gruvel, 1920, p. 33, 5,
or forked carinal margin; caudal appendage long, figs. 4-6; Belloc, 1959, p. 4.
4 or more segments. ?Scalpellurn (Arcoscalpellum) dicheloplax bentho-
Type-species: Scalpellurn dicheloplax Pilsbry, 19Q7b phila: weltner, 1922, 67.
( = Scalpellum debile Aurivillius, 1898 = Neoscalpel-
?Scalpellurn dicheloplax benlophila : Nilason-
lum debile (Aurivillius) , 1898). Cantell, 1938, p. 7.
Remarks : Neoscalpellum includes five abyssal species :
Diagnosis: Capitulum of female ovate in outline,
N . debile (Aurivillius) , 1898, p. 189 (syn.: S. ed-
laterally compressed, rostrum large, broad apically,
wardsi Gruvel, 1902a, p. 65; S. dicheloplax Pilsbry,
clearly exposed; roof of carina with deep medial
1907b, p. 70; S . dicheloplax benthophila Pilsbry,
cleft; scutum with apicolateral arm, and basal margin
1907b, p. 73; S. alboranense Gruvel, 1920).
of plate not hollowed out; inframedian latus initially
N . phantasms (Pilsbry) , 1907a, p. 194.
rectangular, becoming wineglass-shaped, the umbo
N. schizoplacinum sp. nov.
medial or higher; crest of labrum armed with about
N. eltaninae sp. nov.
40 small teeth; mandible with 3 teeth and receding
N. marginatum (Hoek), 1883, p. 65 (syn.: S . ova-
serrate inferior angle; maxilla I with 3 or 4 moder-
turn Hoek, 1883, p. 69).
ately short, stout spines above medial notch; maxilla
In proposing the section ~lieoscal~ellum, Pilsbry
11 triangular and slightly trilobate; setation of cirri
(1907b, p. 69) in all likelihood tacitly assumed that
111-VI ctenopod, with 4 or 5 pairs; distal seta on
it would subsequently come to be r e c o ~ i z e dat the
intermediate articles of inner ramus on posterior
genus-group level because he designated Scalpellurn
cirri paired, but on outer ramus hypolasiopod with 4
dicheloplax as type-species. Initially, Neoscalpellum
setae of unequal lengths; caudal appendage multi-
constituted only a section of Scalpellum, but in the
articulate, with 7-8 segments longer than pedicel of
following year Pilsbry (1908, p. 109) adopted Hoek's
cirrus VI.
(1907b, p. 59) section Arcoscalpellum, elevated it to
subgeneric rank, and included Neoscalpellum within Material: Albatross, Sta. 2711, type-locality for Scal-
its confines, thus clearly delimiting its affinities. pellum dicheloplax dicheloplax, east of Atlantic City,
Withers initially employed Arcoscalpellum and Neo- New Jersey (3S059'N, 70°07'W), 2823 meters, 3 speci-
scalpellurn as subgenera (Withers, 1926), but later as mens; Sta. 2222, east of Atlantic City, New Jersey
full genera (Withers, 1953). (39"03'15"N, 70" 50'45"W) , 2810 meters, 1 speci-
Since it is apparent that Neoscalpellum embodies men; Sta. 2221, east of Atlantic City, New Jersey
a group of scalpellids that in many ways differ from (39O05'30"N, 70°44'30"W), 2788 meters, 3 speci-
other scalpellids, we too have employed it at full gen- mens; Sta. 204.2, type-locality for Scalpellum dichelo-
eric rank, and at the same time segregated several of plax benthophila, on continental slope south of Nan-
the species into other distinct groups at a coordinate tucket Island (3g033'N, 6S027'W), 2843 meters, 1
level. It is now apparent that these morphological specimen; Chain 50, Sta. 81, northwest of Bermuda

SYSTEMATIC ACCOUNT
Fig. 49. Neoscalpellz~m debile (Aurivillius) : A, mandible; B, enlarged view of third tooth and
inferior angle of same mandible; C, maxilla I; D, maxilla 11; E, palp; F, intermediate article
of cirrus VI; G, caudal appendage. Albatross, Sta. 2221.
(34"41'N, 66"28'W), 5042 meters, 1 specimen; At- described by Pilsbry. Those specimens not described
lantis I I , Sta. 93, northwest of Bermuda (34"39'N, by him offer little supplementary information, other
66'26'W), 5007 meters, 1specimen. than that all have a clearly defined rostrum. One of
Depository: USNM Cat. No. 11951 (holotype of the dissected paratypes, from Albatross, Sta. 2221
Scalpellum dicheloplax dicheloplax; Sta. 2711) ; 32862 (USNM Cat. No. 8633), measured as follows: total
(paratype of S. dicheloplax dicheloplax; Sta. 2222) ; height 36.4 mm, capitular height 24.3 mm, capitular
8030, 8631, 8633 (paratypes of S. dicheloplax dichelo- width 15.1 mm. Measurements of the specimen from
plax; Sta. 2221) ; 32878 (holotype of S. dicheloplax Chain 50, Sta. 81 are as follows: total height 26.8 mm,
benthophila; Sta. 2042). capitular height 22.1 mm, capitular width 11.7 mm.
Supplementary Description: The shell structure of Those for the specimen from Atlantis 11, Sta. 93 are:
the majority of the specimens we studied has been total height 21.8 mm, capitular height 17.6 mm, capit-

Fig. 50. Neoscalpellum debile (Aurivillius) : A, mandible; B, enlarged view of third tooth
and inferior angle of same mandible; C, D, maxillae I ; E, maxilla 11; F, palp; G , intermediate
article of cirrus VI; H, caudal appendage. Albatross, Sta. 2M2.

SYSTE&IATIC ACCOUNT 99
ular width 8.8 mm. Cirral counts for this and three a depth of 2959 meters. This record is in need of
other specimens are presented below : confirmation for it appears questionable that N. debile
occurs in the Indian Ocean and Weltner generally had
difficulty in distinguishing between species.
Albatross, Station 2221 9
- 21
- 26
- 28
- 30
- 30
- Neoscalpellum debile occurs on both sides of the
(USNM 8633) 14 24 26 27 27 29 Atlantic Ocean in the northern hemisphere, from
2788 to over 5000 meters.
Neoscalpellum phantasma (Pilsbry, 1907)
Albatross, Station 2042 9
-
18
-
23
-
27
-
29
-
27
-
Text-fig. 51
(USNM 32878) 12 22 24 25 26 30
Scal~ellum~ h a n t a s m aPilsbry, 1907a, p. 194, pl. 6,
10
-
18
-
23
-
27
-
28
-
28
-
8 fig. 1.
12 22 23 26 27 27 Scalpellum (Neoscalpellum) phantasma: Pilsbry,
1907b, p. 73.
Chain 50, Station 81 8 16 23 28 32 32 8
- - - - - - Scalpellum japonicum : Tarasov and Zevina, 1957,
11 21 24 27 31 29
p. 1444, figs. 46, 47.
Diagnosis: Capitulum ovate in outline, laterally com-
pressed, supporting reduced plates in the adult;
Atlantis 11, Station 93
-
8 16 24 28 28 31
- - - - -
8 rostrum small, keyhole-shaped in outline, but dome-
12 20 24 25 27 28 shaped in section, umbo above middle (fig. 51A, B) ;
roof of carina bordered by high, rounded ribs, not
deeply cleft; scutum with apicolateral arm, and a
deeply bifid basal margin; inframedian latus initially
Remarks: Neoscalpellum debile was briefly described rectangular, becoming wineglass-shaped with medial
by Aurivillius (1898, p. 189), from material dredged umbo; mandible elongate, with 3 teeth and receding
from 4400-5005 meters off the Azores. Gruvel (1920) serrate inferior angle (fig. 51C) ; maxilla I with or
subsequently separated specimens from two of the without medial notch, when present with 3 stout spines
three localities Aurivillius cited for this species, and above notch; maxilla I1 trilobate; setation of cirri
described them as a new species, Scalpellum albora- 111-VI ctenopod, 5 pairs (fig. 51H) ; distal setae
nense. However, these are young stages of N. debile paired on inner ramus of posterior cirri, but hypo-
and are included in the present synonymy. lasiopod on outer ramus; caudal appendage 8-9
Pilsbry (1907b, p. 70), working without the bene- se,pents, nearly twice length of pedicel of cirrus
fit of illustrations of N. debile, described Scalpell~~mVI (fig. 511).
dicheloplax. The illustrations and supplementary des- Material: Albatross, Sta. 4397, type-locality, off Santa
cription of N. debiCe by Gruvel (1920) make it read- Catalina, California (33O 10r15"N, 121°42r15'rW),
ily apparent that Pilsbry's species is the same as 4014+4073 meters, 1specimen.
that described by Aurivillius, as Gruvel noted. At the
Depository: USNM Cat. No. 32421 (holotype) .
same time Pilsbry described S. dicheloplax, he des-
cribed S. dicheloplax benthophila, dredged by the Supplementary Description: Aside from pointing out
Albatross from essentially the same region and depth the presence of the rostrum in N. phantasma, nothing
as the nominate subspecies. Study of the types of the need be added to the original description. The
two forms leads us to conclude that Pilsbry fell into present species has 3 pairs of well-defined spines
the same error as Gruvel; that is, S. dicheloplax ben- present on the dorsum of the thorax, comparable to
thophila is simply a young stage of S. dicheloplax di- those seen in other species of the genus. Cirral counts
cheloplax. of the holotype are as follows:
Hoek (1914, p. 4 ) noted some differences between
the specimens he studied, and those figured by Pilsbry,
but only stated that they appeared to be intermediate
growth stages.
Weltner (1922, p. 67) reported S. dicheloplax
benthophila from off Dar es Salaam, Tanzania, from

Fig. 51. Neoscalpellum phantasma (Pilsbry) : A, B, front and sidc views of rostrum, respectively;
C, mandible; D, enlargrd view of third tooth and inferior angle of same mandible; E, maxilla I;
F, maxilla 11; G, palp; H, intermediate article of cirrus V I ; I, caudal appendage. Albatross,
Sta. 4397 (holotype) .
Remarks: In many respects N. phantasma is similar pedicel of cirrus VI. The caudal appendage figured
to the northern Atlantic species N. debile. It is readily by Tarasov and Zevina (1957, fig. 64) does not ap-
separable from N. debile by possessing a basal notch pear to be as long as that of the holotype.
in the scutum, a keyhole-shaped rostrum, lacking a The specimens Tarasov and Zevina (1957) des-
cleft in the roof of the tergum, and by the caudal cribed and illustrated as Scalpellum japonicum are
appendage, which is nearly twice the height of the tentatively referred to the present species. Notable

differences 11etwc:en S. japonicurn, anti their specimens ting edge; maxilla 11 triangular in outline; dwarf male
become readily apparent when one studies the shape sack-like, lacking rudimentary c,alcareous plates.
arid form of ~ l l ccapitular plates, especially the rostra1 Matc7rial: Eltanrn, Sta. 1150, type-localit), southwest-
latus, inIramedian latus, and the upper latus. ern edge of Southeast Pacil~c Hasin (6So37'S,
Neoscnlpellum schizoplacinz~msp. nov. 121 "Ob'W), .375::-180 1. meters, 2 specimens.
Plate X A, Text-fig. 52 Drpository: U S N M Cat. No. 125272 (holotype) ;
Diagnosis: Capitulum of female with reduced calci- 125273 (paratype).
fied plates; both arms of upper latus forked terminal- l)esr.riptiorr: Femalc Capitulum elongate-oval, higher
ly; hasal margin of scutum with deep, broad notch; than broad; both margins convex, carinal more so
rostrum small, triangular, apically acute with umho than occludent; plates marked by growth ridges only.
at apex; roof of carina lacking median sulcus and Tergum forked; lateral arm extending to carina at
bordering ribs; mandible with 4 teeth including re- level below midpoint of scutum. Scutum with slender
ceding inferior angle; maxilla I without notch in cut- al)icolaieral arm; basal margin of plate deeply notched.
Fig. 52. Neoscalpellurr~ schizoplatin~~n~ sp. nov.; A, view of lcft side of thorax showing dorsal thoracic
spines; B, mandible; C cnlargcd vicw of inferior anglc of same mandible; D, maxilla 11; E, niaxilla I;
F, palp; G, intermediate articles of cirrus VI; H, caudal appendage. Eltnnin, Sta. 1150 (holotype, fig.
B, C only).

102 ANTARCTIC
Upper latus forked; lateral arm broader and shorter intermediate articles with about 22 setae arranged in
than depending arm; both arms forked terminally. 4 irregular rows. Cirri 111-VI equal; each articulation
Carinal latus forked; plate about Ijc, higher than infra- along greater curvature of intermediate articles of
median latus; apicocarinal arm slender, acuminate; cirri JI-VI with 1 long and 1 or 2 short setae. Greater
basal segment broad; on paratype a small prong bis- curvature of the intermediate articles of the inner
sects plate angle. Inframedian latus large, hour~lass- ramus of cirrus 11 with 1-2 spines, cirri 111-VI with
shaped, apically forked. Rostra1 latus quadrangular; none; outer ramus of cirrus I1 with 2-3, cirri 111-VI
lateral margin higher than occludent; plate same with only one or none. Setation ctenopod (fig 52G) ;
height as inframedian latus; lateral margin deeply cirri 111-VI with 4 pairs; cirrus 11 with 6 pairs on
excavated. Rostrum small, triangular, well hidden by outer ramus, 5 pairs on inner ramus; cirrus I11 of
rostral latera; plate about lh height of occludent mar- paratype with 5 pairs, and 4 pairs on cirri IV-VI.
gin of rostral latera. Cirral counts for both types are as follows:
Peduncle about v$or less height of capitulum;
about 11 rows of closely packed scales, 8 scales per
row; scales thin, arcuate, somewhat irregular. Holotype x xx 28 32 33 34 9
- - - - - -
x xx 26 28 xx 32
Total Capitular Capitular x 20 xx xx xx xx 9
- - - - - -
Height Height Width x 25 xx xx xx xx
- - -
IIolotype 33.2 22.4 13.7

I'aratype 9 14 25 25 27 27 7
Paratype 21.1 16.3 8.4 - - -- - - -
10 17 21 23 24 26
Labrum partially destroyed in paralype, and lack- 10 17 21 23 26 28 8
ing in holotype; apparently bullate, simple, no teeth 11 17 21 22 24 25
visible. I'alp small, conical, tubular; sparsely covered
with stout spines terminally and on superior margin; Caudal appendage with 7 to 9 articles (fig. 52H) ;
spines not serrate or plumose. Mandible with four appendage longer than pedicel of cirrus V I 3 long
teeth, including receding inferior angle; lateral face terminal spines, longest nearly as long as distal 3
with stiff setae; inferior margin hirsute; first tooth articles; each article sparsely setose.
separated from second; third tooth overhanging in- Dwarf males sack-like, globose, ovate in outline
ferior angle; inferior angle serrate, 8 teeth on holo- (0.95 x 1.35 mm) ; surface covered with concentric
type (fig. 52C). Maxilla I small, lateral face sparsely bands of spines, lacking calcareous plates.
covered with bristles; 2 notches along cutting edge; Remarks: The forked or cleft development of the
2 long stout and 1 short stout spine above superior capitular plates places this species within the genus
notch; 3 s p i 1 1 ~in~ middle clusier somewhat shorter Neoscalpellum. Of the species included in the genus,
than rpines in superior cluster; 5 spinc%sin basal N. schizoplacinum mt st closely resembles A'. debile
cluster, 4 short and 1 long; long spines same length and N. phan~asma. A major diiference useful in
as long spines in middle cluster. Maxilla I1 roughly separating it from this group is the small rostrum
triangular; margins moderately covered with stout hidden by the rostral latera. The basal margin of
spines; spines not clearly segregated into distinct the scutum in N. dehile is either entire or the basi-
clusters; maxillary lobe large, stout, apically trun- lateral angle is slightly rounded off, never deeply
cate; free of setae (fig. 52D). notched as it is in N. phnntasrr~a and N. schizoplaci-
Dorsolateral surface of thorax with 3 pairs of hook- num. The terminal bifurcation of both arms of the
like projections (fig. 52A) ; short, distally acuminate, upper latus also serves to separate N. schizoplacinum
on segments supporting cirri 111-V. from the previously described species.
Cirrus I somewhat separated from cirrus 11; ahout The holotype of N. schizoplacir~umhas been bored
36 length of cirrus 11; anterior ramus slightly shorter by a gastropod, the small hole in the scutum being
than posterior ramus; intermediate segments protuber- near the insertion of the adductor muscle. However,
ant, about broader than segments of posterior the hole had been repaired and the animal, although
ramus; both rami densely clothed with long, raridomly it lost nearly all of the cirri on the left side, re-
scattered, nonplumose setae. Inner ramus of cirrus I1 generated moderately large, uniarticulate anterior
somewhat longer than outer ramus; inner face of cirri.

The specific name is derived from the Greek, and 11 ovotriangular in outline; dwarf male sack-like,
refers to the nature of the capitular plates, which are lacking rudimentary calcareous plates.
split or forked. Material: Eltunin, Sta. 43, type-locality, west of San
Neoscalpellun~eltaninae sp. nov. Vicente de Cafiete, Peru (1301gfS, 78°04/W), 5 2 3 4
Plate X B, Text-fig. 53 5314 meters, 1 specimen.
Diagnosis: Capitulum of female with reduced calci- Depository: USNM Cat. No. 125274 (holotype) .
fied plates; carinal arm of upper latus short and Ilescription: Female - Capitulum narrow, trapezi-
broad, depending arm long and very slendcr; inframe- form, apically acuminate. Scutum. tergum and upper
dian latus key-hole shaped; lateral margin of roslral latus notched, remaining valves unmodified. Lateral
latus not notched; rostrum extremely narrow, acicular, fork of tergum broad, long, extending almost to
exposed; mandible with 3 uniform, widespaced teeth carina; lateral arm of scutum ?13of tergal margin,
and a receding inferior angle; maxilla 1 with numer- narrow. Upper latus with long, narrow, depending
ous stout spines, no notch in cutting edge; maxilla spur. Carina simple, bowed. InIramedian latus sim-
Fig. 53. Nroscalpellurra eltaninae sp. nov.; external view of roFtrun1, adjoining rc~stral latera ant1
sc:uta; B, inframcdian latus; C, labrurn and right palp; D, palp; E, mandible; F, enlargcd view of third
tooth ant1 inferior angle of same mandible; G, maxilla 1; H, maxilla 11; I, intermediate article of
inner rarnus of right cirrus VI; J, distal articlcs of same cirrus VI; K, caudal appendage, Eltanin,
Sta. 43 (Ilolutype).

ple, long and narrow, with apical, spatulate, subcuticu- Caudal appendages G to 9 articles; proximal 5 as
lar extension (fig. 53E). Carinal latus apically acumi- long as pedicel of cirrus VI; terminal article sup-
nate, twice the hcight of the inframedian latus. Rostral porting 5 distal setae, the longest being nearly as
latus quadrangular, narrowing slightly toward occlu- long as the 3 terminal articles (fig. 53K).
dent margin. Rostrum extremely narrow, nearly as Dwarf male sack-like, globose, elongate (0.95 X 1.55
high as rostral latus (hg. 53A). All valves with mm) , covered with concentric bands of spines; lacking
growth lines, but no longitudinal striae. calcareous plates; no appendages observed.
Peduncle approximately 1,k to ihe height of the Remat ks: The present species is tentatively assigned
capitulum; heavily armored with broad. imbricating to the genus Neoscalpellum because of the lack of a
plates in E-9 rows. notch in the lateral margin of the rostral latus, the
The unique specimen measures as follows: total abscnce of the bifurcating apical border of the in-
height 22.0 mm, capitular height 16.9 mm, capitular framedian latus resulting in the typical hourglafs or
width 8.1 mm. wineglass shape, and the extremely broad lateral arm
Labrum slightly bullatc.; anterior portion smooth; of the upper latus. However, since the specimen may
crest with about 25 short, peg-like teeth arranged in not have reached that point in its development where
2 staggered rows. Palp tubular, conical, with short the plates assume the condition found in other rnem-
plurnose selae on both superior and inferior margins; bers of this genus this assignment is considered ac-
long plumose setae on distal portion. Mandible with ceptable on the basis of other similarities. Of the
1 teeth including inferior angle; teeth more or less many similarities to lhis group the apicolateral arm
equidistantly spaced (fig. 53E) ; inferior angle re- of the upper latus, a rostral latus that is as high as
ceding and serrate, with 1 0 teeth (fig. 53F) ; lower or nearly as high as wide, a male lacking calcareous
$5 of mandibular surface clothed with clusters of setae; plates, and the receding inferior angle of the man-
2 to 3 setae per cluster. Maxilla I with 11 spines; dible, serve as uniting chararteristics. The shape of
superior pair long, stout, prominent; cutting edge the rostrum in N. cltaninae, in addition to the above
without notch (fig. 53G). Maxilla 11 subquadrangu- mentioned characters, serves to separate this species
lar, clothed with 2 clusters of plumose setar, on from other Neoscalpellum.
supe~ior,upper frontal and lower frontal margins;
lower cluster with f e ~ e setae
r than remaining clusters; Abnthescalpellum Fen. nov.
maxillary lobe large, erect and free of setae. Diagnosis: Capitulum of adult female with 14 modi-
Cirrus I separated from cirrus 11; l e ~ sthan I,$ fied o r reduced calcified plates; tergum not forked
ler~gthof cirrus 11; rami subequal; anterior about 1A basally; scutum with short apicolateral arm, about
broader than posterior. Posterior rami of cirrus I1 I,<; length of tergal margin, and with broad and
slightly longer than anterior; arrangement of setae shallow notch in basal margin; upper latus bifid, with
of intermcdiatc articles ctenopod (fig. 531), 4 pairs long and broad depending arm; carinal latus higher
on face of lesser curvature, 1 or 2 on greater curva- than wide and umbo at basicaririal angle; inframedian
ture; I major arid 1 or 2 lesser spines at each articu- latus wider than high, slightly hollowed out apically,
lation along greater curvature of intermediate articles; and umbo basal; rostral latus about as high as wide;
inner lateral face densely setose, setae randomly ar- caudal appendage short, with less than 3 fused sep-
ranged. Cirri 111-VI subequal; setation at each arti- mmts.
culation and on lersrr curvature same as cirrus 11; Type-species: Scalpellurn ko~pclncln~Hiro, 1933,
setae on irlner lateral face of intermediate articles of p. 36.
cirrus 111 less setose than 11, arranged in 2 staggered E ~ y m o l o g y : The genrric name, derived from the
rows; those of 1V iri one row of 4, on V in orie Greek, alludes to the shallou-watcr occurrence of this
row of 5, ant1 on V1 one row of 2-3. Cirral courits scalpellid (Gr. abathes, shallow).
for the holotype arc as follows: Remarks: This genus includes a single species, and
has its closest affinities with Neoscalpellum. The
normal shape of the tergum, the broad, swollen arms
of the upper latus and the extremely broad inframed-
ian latus in Abathescalpellum readily serve to separate
the two. Moreover, Abathescalpell~~rn is geographical-
ly and bathymetrically isolated from Neoscalpellum.

S \ STEMATIC ACCOUNT 105
(;ymnoscalpellum gen. nov. pendage multiar ticulate, 5-6 segments; dwarf male
sack-like, bcaring 4 rudimentary calcareous plates.
Diagr~osi.5: Capitulum of adult female with 14 re-
duced calcifietL plates; tergum in shape of inverted Material: Eltanin, Sta. 11:3, southeast of O'Brien
V; scutum either lacking or possessing a minute, ex- Island, South Shetland Islands (62°07'S, 55'58'W),
ceedingly narrow, apicolateral arm '/I? or less length 1113-1153 meters, 2 specimens; Sta. '130, type-local-
of tergal margin. and a deeply notched basal margin; ity, southwest of King George Island, South Shetland
upper latus either banana-shaped or bifid, when lrifid Islands (62"38'S, 59O37'W), 681-1409 meters, 4
with long depending arm; carinal latus higher than specimens; Sta. 091, north of Bridgeman Island,
wide, umbo at basicaririal angle; inframedian latus South Shetland Islands (00"57'S, 56O52'W), 2072-
small, triangular, with concave basal margin, umbo 3020 meters, 1 specimen; Sta. 095, south of Aspland
apical; rostral latus wider than high; caudal ap- Island, South Shetland Islands (6l057'S, 55'53'W),
pendage long, 4 or more segments. 2119-2502 mete~s, 1 specimen; Sta. 997, south of
Aspland Islantl, South Sheiland Islands (61°44'S,
Type-species: Gymr~oscalpell~~rntarasovi sp. nov.
55" 56'W) , 760 meters, 2 specimens.
Etymology: The generic name is derived from the
Greek, arid draws attention to the reduced armament
of the capitulum.
types; Sta. 413) ; 125278 (paratype; Sta. 991) ;
Remarks: Gymnoscalpellurn is proposed for the re- 125279 (paratype; Sta. 995) ; 125280 (paratypes;
ception of four species, namely: Sta. 997).
G. tarasovi sp. nov.
Description: Female-Capituluni elongate and oval,
G. larvale (Pilsbry) , 1907a, p. 194.
higher than wide; both margins slightly convex; more
G. insigne (EIoek) , 1883, p. 68. apically acuminate in small specimens than in large
G. leoni (Zevina) , 1968, p. 90. ones; plates ornamented with growth lines only.
Differences between the two foregoing genera and Lateral arm of tergum extending to and abutting
the present one rest largely on the virtual absence in against carina in small specimens, but not in large.
Gynanoscalpellum of an apicolateral arm on the Scutum with small apicolateral spur; spur not be-
scutum; the broader than high rostral latus; the coming significantly larger with subsequent growth;
triangular inframedian latus with the nmbo apical,
basal margin entire in young stages, but subsequently
rather than medial as in Neoscalpellum or basal as in
developing a deep notch in later stages. Carina bowed;
A1:wthesculpellum; and the extremely deep basal notch
in the scutuni approaching 1," the total height of the roof sulcate and bounded by prominent ridges. Upper
plate, rather than less than y5in Abathescalpellum or latus subrectangular without basal notch in young;
less than 1k3 in Neosca/pellum. notch gradually deepening, producing a plate with 2
Assignment of Scalpellum insigne to this genus is arms; subcuticular apical extension of plate large,
predicated on the fact it appears to be in many fea- spatulate. Rostra1 latus about twice height of infra-
tures identical with certain stages of S. larvale. In median latus; subquadrangular in small specimens;
all likelihood, the specimen I-Ioek figured represents apical half narrowing and becoming acuminate in
an ontogenetic stage similar to that shown in plate large specimens; basal width remaining proportion-
XI B herein. ately constant. Inframedian latus small, triangular
(fig. 54K, L) ; apical portion gradually narrowing and
Gymnoscalpellum tarasovi sp. nov. becoming acuminate in subsequent stages. Carinal
Plate XI, Text-fig. 54 latus rectangular, broader than high; height not ex-
ceeding that of infraniedian latus. Rostrum small,
Diagnosis: Upper latus bifid, with long apical, sub-
subtriangular, basally acute; umbo subapical (fig.
cuticular extension; inframedian latus essentially tri-
angular and lower than adjoining margin of rostral 545).
latus; rostrum subtriangular, acute basally, with umbo Peduncle about y2 height of capitulum; plates thin,
subapical; maxilla I with deep medial notch in cutting irregular and closely packed in small specimens, few
edge; maxilla 11 ovotriangular in outline; caudal ap- and widely separated in adults.

Fig. 54. Gymnoscnlpc~llr~ntnrnsoui gen. e t sp. nov.: A, mantiihlc; B, enlarged view of inferior angle of
same mandible; C, ~ n l a r g e dview of inferior angle of diffcrrnt rr~andible; U, nlaxilla I; E, rnaxilla 11; F,
palp; G, intermediate article of outer ramus of left cirrus VI; 11, ter~nirlal articles of same cirrus V I ;
I, caudal appendagc; J, rostrum and adjoining portions of rostra1 latera; K, L, inframedian latcra. Eltenin,
Sta. 430 (holotype, figs. A, B, D-I, K only) and Sla. 41.3.

Measurements of the ten specimens of this species Cirral counts for the holotype and a paratype from
are presented below : Station 430, and a paratype from Station 413 are
Total Capitular Capitular given in order below:
Station Specimen Height Height Width
I I1 111 IV V VI Ca
430 Holotype 28.2
430 2 24.3 Station 430
430 3 15.9 Holotype 9 17 23 24 xx 29 6
- - - - - -
430 4 10.5 12 20 24 xx 25 xx
413 1 32.5
413 2 12.5
991 1 40.4
995 1 30.4
997 1 Specimen 2 10 13 19 22 24 25 5
17.6 - - - - - -
997 2 19.1 11 16 20 24 24 24
Labrum bullate; anterior portion smooth; crest 7 13 19 22 25 26 5
- - - - - -
armed with about 50 very small, low, triangular teeth 10 17 21 22 24 25
in large, 20-25 teeth in small specimens. Palp tubu-
Station 413
lar, conical, sparsely setose; setae slightly plumose.
Specimen 1 9 19 22 26 27 28 6
Mandible with 4 evenly spaced teeth, including in- - - - - - -
12 21 23 25 27 27
ferior angle (fig. 54A) ; tooth 3 overhanging inferior
angle of 4 spines. Maxilla I with deep medial notch
(,fig. 54D) ; 2 large, stout spines and 2-3 short spines
above notch; 7-9 spines below notch. Maxilla I1 sub- Caudal appendage moderately long, 6 articles (fig.
quadrate; plumose setae more or less segregated into 541) ; proximal 4 segments as long as first segment
3 clusters along cutting edge; maxillary lobe large, of pedicel of cirrus VI; complete appendage shorter
moderately short, broad, apically truncate, free of than both articles of pedicel; terminal article with 2
setae (fig. 54E). long and 3 short setae on crown; long setae approx-
Cirrus I well separated from and approximately imately twice length of terminal article.
1/2 length of cirrus 11; anterior ramus slightly shorter
Dwarf male sack-like, elongate-oval (0.45 X 0.90
than posterior ramus with intermediate articles about
mm to 0.55 X 1.05 mm) ; external surface covered
1/4 broader than posterior; both rami clothed with
with concentric bands of fine spines; calcareous plates
long, nonplumose setae, randomly scattered on inner
present; size of plates on an individual differ con-
lateral face, absent on narrow zone both above and
siderably, the larger pair being nearly twice the size
below each articulation. Inner ramus of cirrus I1
of the smaller pair. Male cyprids very long and
significantly longer than outer ramus. Each articula-
slender (0.35 X 1.10 mm to 0.50 X 1.25 mm).
tion along greater curvature of intermediate articles
of cirri 11-VI with one long and 1 or 2 short setae; Remarks: Both of the previously described species
on greater curvature of cirrus I1 intermediate articles here assigned to Gymnoscalpellum occur in the north-
with 6-8 setae. Cirri 111-V with 3 to 4 unpaired ern hemisphere. Of these, G. tarasovi is closely
setae; cirrus VI with 2-3 unpaired setae. Distal clus- related to G. larvale (Pilsbry, 1907a, p. 194), dredged
ter of setae on intermediate articles of outer ramus by the Albatross from 1168 meters off San Diego, -
of posterior cirri hypolasiopod; inner ramus normal. California. The banana-shaped upper latus, total ab-
Setation along lesser curvature ctenopod (fig. 54G). sence of an apicolateral scutal arm, and the distinctly
Number of pairs of setae per intermediate article of notched basal margin of the inframedian latus dis-
right cirri are as follows: tinguish G. larvale from G. tarasovi. Moreover, the
I1 I11 IV V VI caudal appendage of Pilsbry's species is shorter, has
Outer ramus fewer segments, and the terminal cluster of setae on
Holotype (Sta. 430) 8-9 5 5 5 5 segment 4 are markedly longer than found in this new
Paratype (Sta. 430) 5 4 4 4 4 species. Gymnoscalpellum insigne, dredged by the
Paratype (Sta. 413) 8-9 5 5 4 4 Challenger from 2788 meters off the Azores (36'
Inner ramus
23'N, 11°18'W), apparently had not reached that
Holotype (Sta. 430) 5-6 5 5 5 5
Paratype (Sta. 430) 5 4 4 4 4 stage in its development where the basal margin of
Paratype (Sta. 413) 5 5 5 4 4 the scutum bifurcates, and in general the plate appears

108 ANTARCTIC
to be sigrlihcantly broader than is the case in G. I,. aurorae (Gage), 19S& p. 6.

larvale and G. tarasovi. L. interrrzeiliurn t Hock), 1883, p. 70.
The rostrum of G. insigne was apparcntly over- L. nipponense (Pilsbry) , 190Tb, p. 73.
looked by Hoek, as it was in G. larvale by Pilsbry. 1,. laccadivzcurn (Annanclale) , 1906, p. 393 (syn.:
The rostrum of G. larvale is small, triangular, acute Scalpellum lmcadivicum var. investigatoris An-
basally and totally hidden by the rostral latera, and nandale, 1006, p. 395).
thus similar to that of G. tarasovi. L. korotkevitshae (Zevina ) , 1968, p. 33.
This new species is named in honor of the late L i ~ o ~ c a l ~ e l l is
u mproposed for a large number of
Russian cirripedologist, Dr. Nicolas lvanovitch Ta- species with relatively unspecialized capitular armor,
rasov, 1909-1965. holding a superficial resemblance to Arcoscalpellum.
Zevina's (1962) paper was received after submit- While relatively unsprcialized plates can serve to dis-
ting this manuscript for publication. The holotype tinguish the two genera, certain arthropodal struc-
of Scalpellurr~ leoni (Zevina 1968, fig. 4a) is appar- tures aid considerably in separating them. The multi-
ently a distinct species, whereas the paratype (Gg. articulate caudal appendage is generally useful, but
4b) is clearly referrable to G. tarasovi sp. nov. The it is occasionally multiarticulate in Arcoscalpellun~,
morphological characters of S. leoni (=Cymnoscal- especially in the group of A. michelottianum. Like-
pellum leoni) that separate it from G. tarasovi are wise, the spines along the curvature of the intermedi-
the prominent apicolateral arm and deeply incised ate articles of the posterior cirri is apparently a
carinal margin of the scutum, banana-shaped upper characteristic of Li~oscalp~~llrrrr~ rarrly found in Arco-
latus, irrrgular and compressed rostral latus, shorter scalpellzcm.
carinal arm of the tergum which does not extend to
the carina, and greater disparity in number of articles Litoscalpellum fissicarinatum sp. nov.
between the anterior and posterior rami of cirri 111- Plate X C, Text-fig. 55
VI. Diagnosis: Roof of carina with deep, narrow
Litoscalpellum gen. nov. cleft running complete length of plate; inframedian
latus small, lower than adjoining margin of rostral
Diagnosis: Capitulum of female armed with 14
latus, and broadly triangular; rostrum T-shaped, but
mostly unreduced calcareous plates; tergum normal
essentially triangular in outline; mandible with 4
or with broad shallow notch in basal margin; scutum
teeth and a serrate, produced inferior angle; maxilla
without, or with slight apicolateral arm less than l/i-,
I with 2 notches in cutting edge; maxilla I1 triangular
width of narrow to moderately broad tergal margin,
in outline and slightly trilobate; caudal appendage
and hasal margin entire, never hollowetl out; upprr
multiarticulate and longer than pedicel of cirrus
latus triangular or elongate, commonly with slightly
VI; dwarf male sack-like, with 4 rudimentary cal-
hollowed out hasal margin, but rarely with a deep
careous plates.
notch; carinal latus as high as wide or slightly higher
than wide; inframedian latus triangular, narrow, Material: Eltanin, Sta. 1084, type-locality, between
umbo apical; rostral latus wider than high, with Larsen and Inaccessihlr Islands, South Orkney lslands
height diminishing from lateral to rostral border; (6O022'S, 46"50'W), 293-103 meters, about 30 speci-
caudal appendage moderately short, with I to 9 srg- mens.
mrnts. Depository: ITSNM Cat. No. 125221 (holotype) ;
Type-species: L i t o ~ c a l ~ e l l u jissicarinat~crr~
m sp. nov. 1252::2 (paratypes).
Etymology: The generic name, derived from the Description: Female-Capitulum elongate and oval;
Greek, alludes to the relatively unspecialized or simple nearly twice as high as broad; occludent less convex
nature of the capitular plates. than carinal margin; capitular plates with growth
Remarks: The following species are included in this lines only. Carina houed; roof deeply cleft from
genus : base to apex, not bounded by angles (fig. 55B).
L. jissicar inatum sp. nov. Tergum crescent-shaped, projecting apically, acu-
L. simplex sp. nov. minate at both extremities. Scutum with short apico-
I,. walleni sp. nov. lateral arm; superior margin of arm with narrow,
L. discoveryi (Gruvel), 1906, p. 271. subcuticular extension; hasal margin of plate entire.
I,. convexum (Nilsson-Cantell) , 1921, p. 103. Upper lalus quadrangular; later stages with short

SYSTEMATIC ACCOUNT
Fig. 55. Litoscalpellum fissicarinatum gen. et sp. nov.: A, inframedian latus and adjoining latera; R,
basal portion of carina with transverse section of same to the right, and contiguous portions of carinal
latera; C, rostrum and adjoining portions of rostra1 latera; D, mandible; E-G, inferior angles of man-
dibles; H, maxilla I ; I, maxilla 11; J, palp; K, intermediate article of inner ramus of left cirrus VI;
L, caudal appendage. Eltanin, Sta. 1034 (holotype, figs. D, F, H-L only).
carinal arm and long depending

- - scutal arm; with Carinal latus subquadrate to triangular; broader than
long, apical, triangular, subcuticular extension. In- high in holotype; equidimensional in small para-
framedian latus small (fig. 55A), approximately 3 types; apically acuminate in holotype, but less so in
times as broad as high in large paratypes and holo- paratypes. Rostra1 latus quadrangular; broader than
type, nearly equidimensional in specimen 8.2 mm in high; narrowing towards occludent margin. Rostrum
capitular height, and higher than broad in smaller triangular to T-shaped (fig. 55C) ; umbo central;
specimens; higher than rostra1 latus in small para- apical margin broad and slightly convex; acute
types, but shorter in holotype and large paratypes. basally.

Measurements of the holotype and 9 paratypes are followed by a summary of cirral counts for the holo-
summarized below : type and two of the dis~ectedparatypes:
Height Hcight Width
Holotype 36.3 26.1 15.7
R
- 6.5-36.3 5.2-26.1 2.8-15.7
X 16.75 12.36 7.08
Peduncle in large specimens $4 or less height of

capitulum; peduncular scales small, sparsely scat-
tered and primarily subcuticular.
Labrum bullate; crest supporting 2 or more rows
of short, triangular or peg-like teeth; teeth number
about 30 in holotype, and decrease in number in
smaller specimens; central region of crest free of
teeth; soft setae absent. Mandible with 4 teeth in-
cluding inferior angle (fig. 55D) ; superior 3 teeth
equidistantly spaced; third tooth not overhanging
inferior angle which consists of numerous imbricating
teeth; lower T$of lateral side covered with clusters Cauclal appendage moderately long, 6-7 segments
of 2-3 stiff setae. Maxilla I with 2 notches along (fig. 55L) ; appendage taller than pedicel of cirrus
cutting edge (fig. 55H) ; 4 spines above first notch VI; terminal article with 3 long and 1 short spine on
on 3 specimens dissected; 4-7 teeth between notches; apex; long spines somewhat longer than terminal
4 8 teeth below inferior notch. Maxilla I1 triangular article of appendage.
in outline; stout, nonplumose setae segregated into Dwarf male sack-like, globose, ranging in size from
3 distinct clusters along margin; numerous stout 0.40 X 0.75 mm to 0.75 x 1.20 mm: surface of sack
setae on lateral face; maxillary lobe long, broadly covered with concentric bands of spines, and bearing
rounded apically, free of setae (fig. 551). four rudimentary calcareous plates; 3-5 males in
Cirrus 1 separated from and about q$length of each scutal pouch.
cirrus 11; anterior ramus slightly shorter than poste- Remarks: This species is characterized by the broad-
rior; intermediate articles protuberant; about lh ly T-shaped rostrum and the cleft roof of the carina,
broader than segments of posterior ramus; both rami which readily separates it from such new species as
densely clothed with long, nonplumose, randomly ar- L. simplex and L. walleni, described below. The
ranged setae. Rami of cirrus I1 subequal. Cirri young stages of these species will likely prove difficult
111-VI subequal; each articulation along greater cur- to separate, but in L. fissicarinatum the cleft in the
vature of intermediate articles with l long spine and carina appears when the specimens reach a total
W2 short spines. Number of setae between articula- height of about 7-10 mm. In the young stages, the
tions, along greater curvature of intermediate articles, rostrum is extremely large and triangular, and rather
of cirri 11-VI on left side are as follows: dissimilar to the adult form.
The specific name, derived from the Latin, draws
I1 I11 IV v vI attention to the cleft in the carina.
Ilolotype
Scalpellurn korotkeuilshae Zevina (1968, p. 89)
anterior 4-5 3 2-3 2-3 2-3
posterior 3 4 2-3 2-3 1-2 1-2 is here assigned to Litoscalpellum Sen. nov., and is
Paratype 1 apparently related to L. fissicarinaturn sp. nov., since
anterior 1-2 1-2 1 1-2 1-2 it too has a narrow cleft in the roof of the carina.
posterior 1-2 1 1 1 1 However, it differs from L. fissicarirzatum in possessing
Paralype 2 a minute key-hole shaped rostrum, a labrum lacking
antprior 2-3 1-2 1-2 1 1-2 teeth, a bifid inferior angle of the mandible, rami of
posterior 1-2 1 0-1 CLl &1
cirri 111-VI equal in length, and in having 5 pairs of
Setation ctenopod (fig. 55K) ; 4 pairs on cirri IV- setae per article on the lesser curvature of the rami
V1. Cirral counts of the holotype are presented below of cirrus VI. Future studies should serve to confirm

or negate the apparent differences between these two rostrum in one specimen is small and rounded, in the
species. other minute. The lower lateral plate is triangular
as also are the carino-laterals which do not meet
Litosca2pellum aurorae (Bage, 1938)
below the carina.
Text-fig. 56
"Two specimens with the capitulum measuring 20
Scalpellurn (Arcoscalpellurn j aurorae Bage, 1938, mm. and 22 mm. respectively.
p. 6, text-fig. 1. "The peduncle is very short, in both specimens,
Diagnosis: Roof of carina evenly rounded, heavy, slightly less than half the length of the capitulum. It
extremely wide in apical half; tergum sigmoid; in- narrows very much immediately below the base of the
framedian latus triangular, umbo apical; rostrum capitulum, and its scales are very narrow, elongated
small and rounded; carinolaterals not interdigitating horizontally, and though arranged regularly, far
below carina. apart" (Bage, 1935, p. 6-7).
Material: Aurora, Sta. 3, north of Addie Coast Litoscalpellum convexum (Nilsson-Cantell, 1921)
(66'325, 141°39'E), 287 meters, 2 specimens.
" 57
Text-fie.
Original Description: "Capitulum with 14 plates
Scalpellurn convexum Nilsson-Cantell, 1921, p. 194,
with narrow uncalcified strips, covered by a hairy
text-fig. 29; 1930b, p. 244, fig. 9.
cuticle, between them. Carina continuously curved
more sharply at the anterior end, and at its posterior Material: Swedish South Polar Expedition 1901-
end it reaches the capitulum. The lower border of 1903, off Cumberland Bay, South Georgia (54'115,
the tergum is slightly convex, the carinal border 36" l S f W ) , type-locality, 252-310 meters, 3 specimens.
straight until beyond the carina where it is definitely Supplementary Description: "The first stage in this
concave forming a recurved point, less than one third collection is a pupa with the first primordial valves
of its border projecting beyond the carina. The oc- developed. Of the other valves a very small upper
cludent border is slightly convex. The scutum also latus can be traced. Broch, 1912 and 1924, states
has a distinct upcurving point near its anterior oc- that the upper latus is first developed after the five
cludent border where the umbo is situated. Its lateral primordial plates in species of Scalpellurn. No pe-
border is convex, posterior border definitely concave duncle scales are to be seen as in the corresponding
and into it fits the long narrow rostro-lateral. The stage of S. gibberurn, C. W. Aurivillius, 1892.
upper lateral plate has a curved projection which lies "In the second stage all the valves of the capitulum
in the angle between the tergum and scutum. The are formed. As no intermediate stages are repre-
~ e n t e dwe cannot indicate the order of the appearance
of the plates. We see that the rostrum is originally
of a triangular shape with the umbo at the base. On
the peduncle the first four large scales are developed.
The shape of the inframedian latus is more rounded
than in the later, especially mature, stages. The pro-
cesses, mentioned by Hoek and other authors in
young Scalpellurn at the apex of the capitulum, are
very distinct.
"In the third stage the valves of the capitulum are
somewhat larger. The new scales of the peduncle ap-
pear in the region between the capitulum and the
peduncle, according to Broch. The primordial plates
as in the previous stage.
"In the fourth stage the inframedian latus is more
elongated by the upward growth of the sides which
forms a small accessory part above the strong umbo.
The scales on the peduncle are more developed in
Fig. 56. Litosculpellum aurorae ( B a g e ) : Individual this stage.
viewed from right side (redrawn f r o m Bage, 1938, "Of the fifth stage three plates are figured. The
P. 7 ) . inframedian latus has now reached its definite shape,

1"ig. 57. Litoscalpellum convexurn (Nileson-Cantell) : .A, young individual front

left side, and rostrum with rostra1 margin of rostrolatcral plates; B, full grown
spccirnen f r o n ~left side, from rostra1 side, and carina and carinolatcrals; C,
carina, inframedim latus and rostral latus of young specimen; U, young speci-
rnen from Icft sidc, and rostrum with rostral margins of rostrolateral platrs
(rcdrawn from Nilsson-C'lntell, 1930c, fig. 9, C-1%).
and in the carina the umbo is removed from the apex. now found again in Antarctic regions. There are
"In [the fully grown] stage all the plates are fully some species from the same parts of the ocean nearly
calcified, but separated by chitinous interspaces. The lelated to this, namely S. bouvieri, Gruvel, 1906, and
limits in the chitin show the same shape as the orig- S. weltneri, Gruvel, 1907b. I first thought that all
inal plates, as in the first stages. The calcified por- three were the same species. but since I have com-
tioris of the plates are, in some respects, different pared the types with the present material I must affirm
flom the iridividual figured by me in 1921. Thus the that thcy are all distinct species, though the differ-
tergum has the basal margin incompletely calcified. cwces are not very great. As the figures of both spe-
The other plates are smaller, and the interspaces cies, S. weltneri and S . bouvieri, do not give an exact
wider, than in the young stages. This may be due to impression of the shape of the plates, I must here
the fact that the capitulum and the upper part of the note that the species a l e very closely related in ex-
peduncle are much swollen. ternal characters. Both have the carina concave
"By tlissrcting the specinlens I found a large riurn- dorsally, more in the latter species than in the former.
ber of Cypris stages, as the first development takes S. convexzbm differs from both distinctly in the strong
place in the capitulum. The upper part of the pe- dorsal convexity of the carina. Other differences in
duncle is much cxpanded and, in consecIuerice of this, the plates also exist. S. weltneri arid S. bouvieri are
it is without scales. Such an interspace is not to be externally rather similar, but differences are to bc
fourid in younger specimens. fourrd in the carinolaterals and the irlframedian later-
"In thc: original description the rostrum is said to als. The Grier structure of the plates which I have de-
be long. with the uml)o at the apex. From this ma- termined I)y studying the typcBsmakes the differences
terial wc (:an study the shape more precisely. First, greater than it is possible to see from figures only.
it is triarigular anrl I~ecomesmore and more elonptcd. S. weltneri has a very hairy cuticle and S. bouvieri
The visilde part is often narrow and long but the real a cuticle without hairs. They are thus all found to
shape is morc nc.arly triangular. The umbo must 1)e be good species" I Nilsson-Cantell, 1930b, pp. 244-
situated at the: hase. to judge from the young speci- 2471.
mens as the plate in the upper part is often broader.
The plate seems to bc more anti more reduced during Litoscalpellum nipponense (Pilsbry, 1907)
ontogenp. Text-fig. 58
''r 1
Illis species, first desc:ril)ed hy me in 1021, is Scalpellum nipponense I'ilsbry, 1907h, p. 73, fig. 29;

Broch, 1931, p. 24, fig. 9 ; Hiro, 1933, p. 25, text-fig. 5, and Nilsson-Cantell (1921) have described the capitu-
pl. 1, figs. 7-9b. lar morphology of this species in full. The arthropodal
Scalpellurn intermedium: Nilsson-Cantell, 1921, structures were described and figured by Nilsson-
p. 208, fig. 33. Cantell, Hiro (1933) and Broch (1931). All that
Diagnosis: Tergum bifid; upper latus forked basally; need be pointed out at this time is the arrangement of
roof of carina flat or slightly sulcate, and with border- setae along the anterior curvature of the intermediate
ing ribs; lateral margin of rostra1 latus and carinal articles of the posterior cirri. On the inner ramus of
latus indented; crest of labrum armed with about 50, cirrus VI the setae are arrayed in the normal ctenopod
simple, peg-like or triangular teeth; mandible with 4 manner, but the distal cluster on the outer ramus has
teeth including serrate inferior angle; max- 4 setae of unequal lengths.
illa I with 4 spines above a slight to moderately deep Cirral counts of the holotype are as f o l l o ~ ~ s :
and broad medial notch; maxilla I1 ovotriangular, but
not trilobate; caudal appendage with 3 4 segments,
shorter than first segment of pedicel of cirrus VI.
Material: Albatross, Sta. 3697, type-locality, off Man-
azuru-misaki, Sagami-wan, Japan, 219-302 meters, 1
specimen.
Depository: USNM Cat. No. 32909 (holotype) . Remarks: There are several notable differences be-
Supplementary Description: Both Pilsbry (1907b) tween the holotype and specimens described and illus-
Fig. 58. Litoscalpellum nipporzense (Pilsbry) : A, mandible; B, enlarged view of third tooth and
inferior angle of same mandibIe; C, same, but different mandible; D, maxilla I ; E, maxilla 11; F,
palp; G, intermediate article of cirrus V I ; H, caudal appendage. Albatross, Sta. 3697 (holotype).

trated by previous workers. To judge from the de- (1933, p. 26). The cirral counts are also bracketed by
scriptions of Nilsson-Cantell and Hiro, in the early the counts given by Hiro and other workers.
stages this species has a roof bounded by prominent
Litoscalpellum simplex sp. nov.
ribs. Evidently, these ribs are lost during subsequent
Plate X D, Text-fig. 59
ontogenetic stages since the holotype does not have
bordering ribs. Of the arthropodal structures differ- Diagnosis: Roof of carina flat, without medial cleft;
ences exist in the chaetotaxes of the posterior cirri, inframedian latus small, broadly triangular; rostrum
with the number of pairs ranging from 4 to 5. broad, subquadrate, partially covered by rostral latera,
Spination along the posterior border is also variable umbo apical; mandible with 3 teeth clothed in soft
with 1 to 3 spines present on the intermediate articles. bristles and a weakly serrate inferior angle; maxilla I
On the specimens figured by Nilsson-Cantell, Broch with slight subapical notch in cutting edge and highly
and Hiro, maxilla I has a deeply incised broad notch plumose spines; maxilla I1 quadrate in outline, and
in the cutting edge. The holotype does not exhibit this with plumose setae; caudal appendage multiarticulate,
development; that is, the notch is extremely small and and longer than pedicel of cirrus VI; dwarf male sack-
shallow. The number of segments of the caudal ap- like, lacking rudimentary calcareous plates.
pendage of the holotype is 3 on the left side and 4 on Material: Eltanin, Sta. 410, type-locality, between
the right, agreeing with the range reported by Hiro Elephant Island and Aspland Island, South Shetland
Fig. 59. Litoscalpellz~m simplex gen. et sp. nov.: A, rostrum and adjoining portions of rostral lat-
era; B, inframedian latus; C, mandible; L), enlarged view of inferior angle of mandible; E, maxilla
I ; F, maxilla 11; G, palp; H, intermediate article of inner ramus of right cirrus VI; I, terminal
segments of same cirrus VI; J, left caudal appendage. Eltanin, Sta. 410 (holotype).

Islands (61 1 8 5 , 5fi009'W), 220-240 meters, 2 speci- maxillary lobe large, erect, tubular, somewhat apically
mens. rounded (fig. 59F).
Depository: USNM Cat. No. 125283 (holotype) ; Pedicels of all cirri squamous. Cirrus I approxi-
125284 (paratype). mately q3length of and well separated from cirrus 11;
Description: Female-Capitulum elongate and oval, rami subequal; intermediate articles of anterior ramus
higher than broad; occludent less convex than carinal about 1/4 broader than posterior; setae of proximal
articles of anterior ramus highly plumose. Rami of
margin; ornamented with growth lines only. Carina
simple, bowed; roof flat. Tergum crescent-shaped, pro- cirrus I1 subequal; spination on greater curvature of
jecting apically; pointed at both extremities. Scutum each intermediate article of outer ramus with 4 or 5
spines, inner ramus 1 or 2 ; each articulation along
with short apicolateral arm; basal margin entire. Up-
greater curvature with 1 or 2 minor spines and 1
per latus simple, broad and subquadrangular in small
major spine; setation ctenopod, 4-6 pairs; inner lat-
specimen with short depending occludent arm; signifi-
eral face of intermediate articles setose, setae appear
cantly narrower in larger specimen and roughly sickle-
shaped. Inframedian latus extremely small, triangular scattered randomly. Cirri 111-VI subequal; each ar-
(fig. 59B) ; plate of small specimen much broader ticulation of intermediate articles on greater curvature
relative to large specimen. Rostra1 latus triangular to same as cirrus 11; along each segment 1 or 2 spines;
quadrate, broader than high, narrowing towards occlu- setation ctenopod (fig. 59H), 4 or 5 pairs.
dent margin. Carinal latus triangular; lateral margin Caudal appendage of 5 segments (fig. 59J) ; longer
about 3 times higher than inframedian latus. Rostrum than both segments of pedicel; distal end of terminal
small, partially exposed, subquadrate in outline; basal segment with 2 long setae and several short ones; long
margin broadly rounded; umbo apical (fig. 59A). setae slightly longer than distal segment.
Peduncle approximately I/" height of capitulum; Cirral counts are as follows:
plates small, randomly scattered, primarily subcuticu- I I1 I11 IV V VI Ca
lar on upper % of peduncle. Holotype 7
-
13
-
17
-
19
- -
19 20
-
5
Measurements of the two types are as follows: 9 13 17 19 19 20
Total Capitular Capitular 6 12 17 16 20 23 5
- - - - - -
Height Height Width 6 12 17 17 20 23
Holotype 32.1 20.1 12.1 5
Paratype 7 14 16 19 20 20
Paratype 24.3 16.5 9.7 - - - - - -
9 15 17 xx 18 18
Labrum bullate; crestal region densely covered with
soft setae; one row of about 20 teeth along crest;
teeth closely spaced, short, triangular, peglike when
worn. Palp slender; armed with plumose setae on Dwarf male sack-like, globose, measuring about
superior margin and proximally with soft setae only; 0.65 x 0.80 mm; surface covered with concentric
inferior margin free of setae. Mandible with 4 teeth, bands of fine spines; no calcareous plates present.
including inferior angle (fig. 59C) ; teeth widely sepa- Remarks: The present species is unusual in that the
rated, superior margins of lower teeth densely covered trophic appendages bear soft and densely plumose
with soft setae; left mandible of holotype wikh 4 setae. This feature is not known to occur in any of
major teeth as on right, but with 1 minor tooth be- the other species presently assigned to Litoscalpellum,
tween first and second; similar development of denti- nor has it been observed in Neoscalpellum, Gymno-
tion occurs on left mandible of paratype; proximal scalpellum, or Mesoscalpellurn. Aside from general di-
superior margin of mandible covered with soft setae. agnostic differences in capitular morphology, and the
Maxilla I with small subapical notch (fig. 59E) ; 3 setation of the mouth parts just mentioned L. simplex
stout, long, major spines and 3 4 lesser spines above, and L. fissicarinatum can be separated on the basis
17-20 lesser spines below; spines densely plumose as of the shape of the upper latus and tergum. In L. sim-
are those of distal inferior lateral region. Maxilla I1 plex the tergum is narrower, the basicarinal angle
quadrate in outline; plumose setae arranged in 3 ma- more acicular and the upper latus much narrower than
jor clusters, proximal superior, superior and upper found in L. fissicarinatum.
frontal margin and lower frontal margin; lower fron- The specific name, simplex, alludes to the unmodi-
tal cluster not well separated from upper cluster; fied simple nature of the capitular plates.

Litoscalpellum walleni sp. nov. lacking prominent apicolateral arm; mandible with 4
Plate X E, Text-,fig. 60 teeth including a sparsely serrate, produced inferior
Diagnosis: Inframedian latus essentially triangular, angle; maxilla I with slight medial notch, teeth diverg-
extremely small, narrow, shorter than adjoining mar- ing laterally from cutting edge; maxilla I1 subtri-
gins of latera, umbonal region elevated above general angular and slightly trilobate; caudal appendage multi-
surface of capitulum; carina with flat roof bounded articulate and slightly shorter than first segment of
by angles; rostrum minute, T-shaped, about height pedicel of cirrus VI; dwarf male sack-like, and bearing
of rostral margin of rostral latus, umbo apical; scutum 4 rudimentary chitinous plates.
Fig. 60. Litoscalpellurn walleni gen. et sp. nov.: A, rostrum and adjoining p o r t i ~ n sof rostral latera; E,
inframedian latus and portions of adjoining latera and scuta; C, mandible; D, enlarged view of inferior
angle of same mandible; E, maxilla I ; F, maxilla 11; G, palp; H, intermediate article of inner ramus
of left cirrus VI; I, caudal appendage. Eltanin, Sta. 948 (halotype).

Material: Eitanin, Sta. 948, type-locality, northeast of setae on greater curvature of each intermediate article
Peter I Island, southeast Pacific Ocean (65050rS, of inner ramus 7-9, on outer ramus 3 4 . No spines
88' 56'W) , 4502 meters, 1 specimen. present on greater curvature on inner ramus of each
Depository: USNM Cat. No. 125285 (holotype) . intermediate article of cirri 111-VI; on outer ramus
there are 3 on cirrus 111, 1-2 on cirri IV and V, and
Description: Female-Capitulum elongate and oval;
only one on cirrus VI. At each articulation along
higher than broad; both margins slightly convex;
greater curvature of cirri 11-VI one long spine and
plates ornamented with growth lines only. Carina
1 or 2 spinules. Setation ctenopod (fig. 6OH) ; 5 pairs
simple, bowed; roof slightly convex, bounded by an-
on intermediate articles of cirri 111-V; 4 pairs on
gles. Tergum triangular, projecting apically; pointed
cirrus VI. Inner face of intermediate articles of
at both extremities. Scutum lacking definitive apico-
inner and outer rami of cirrus IV with 4-5 spinules
lateral arm; lateral margin concave, basal margin en-
in one row; cirrus V with 1-3 on inner ramus and
tire. Upper latus simple, roughly quadrangular; with
3 4 on outer; cirrus VI with 1-2 on inner ramus and
large, apical, triangular, subcuticular extension. Cari-
2-3 on outer ramus, in both cases in 1 row.
nal latus higher than wide; lateral margin parallel to
Cirral counts for the holotype are as follows:
carinal margin; 2 prominent ridges extending from
I I1 I11 IV V VI Ca
umbo to basal margin. Inframedian latus small, tri-
angular, extremely narrow (fig. 60B) ; umbo apical; - 8 13 18 21 22 23 4
- - - -
plate slightly more than y2 height of lateral margin 10 16 20 22 22 23
of carinolateral. Rostra1 latus quadrangular; slightly 7 13 19 21 23 23 4
- - - -
broader than high; occludent margin higher than lat- 10 17 21 22 23 22
eral margin; one prominent ridge extends from apex
to basal margin. Rostrum minute, narrow, essentially Caudal appendage moderately short; 4 articles;
rectangular (fig. 58A), expanded apically; umbo api- proximal article nearly as long as distal 3 ; slightly
cal; plate about 1/4 height of occludent margin of ros- shorter than first article of pedicel of cirrus VI (fig.
tral latera. 601) ; terminal article bears 4 extremely long, pinnate
Peduncle about Ij3 height of capitulum; heavily ar- spines, and a few short spines at apex.
mored with low, broad, imbricating scales; 19 rows of Dwarf male sack-like, elongate and oval, measur-
scales with 4-5 scales in each row. ing from 0.75 X 1.05 mm to 0.60 X 1.25 mm;
covered with concentric bands of fine spines and 4
Measurements of the holotype: total height 24.3 mm,
chitinous plates.
capitular height 19.4 mm, capitular width 11.5 mm.
Remarks: Litoscalpellum walleni sp. nov. is most close-
Labrum bullate; crest lacking dentition and soft
ly allied to the two foregoing new species, but it differs
setae. Palp long, spatulate, rounded distally ; sparsely
covered with few stout spines along superior border. in several essential features. The roof of the carina
Mandible with 4 teeth including inferior angle (fig. in L. fissicarinatum is deeply cleft whereas in the pres-
60C) ; first 3 teeth more or less equidistantly spaced; ent species and L. simplex the roof is flat or slightly
inferior angle serrate along basal margin, of 4 spines. convex. The inframedian latus of the present form is
Maxilla I with inconspicuous, medial notch (fig. 60E) ; significantly narrower and the rostrum extremely
arrangement of spines along cutting edge unusual in smaller than in either L. simplex or L. fissicarinatum.
that bases of spines occur in 2 well-separated parallel Maxilla I in L. simplex is quadrate in outline, but in
rows of 5 and 8 on right appendage and 4 and 7 on L. walleni and L. fissicarinatum it is triangular or sub-
left. Maxilla I1 subtriangular in outline; pinnate triangular. In both L. simplex and L. fissicarinatum
spines arranged in 3 distinct clusters along margin; the caudal appendage is longer than both articles of
maxillary lobe long, broad, somewhat apically trun- the pedicel of cirrus VI, but in L. walleni it is shorter
cate, free of setae. than the first article.
This new species is named in honor of Dr. I. E.
Cirrus I separated from and about Ys length of
Wallen, Head, Office of Environmental Sciences,
cirrus 11; anterior slightly shorter than posterior
ramus; intermediate articles about l h broader than Smithsonian Institution, Washington, D.C.
posterior ramus; segments of both rami protuberant, Litoscalpellum discoveryi (Gruvel, 1906)
densely covered with long setae. Inner ramus of cir- Plate X F, Text-fig. 6 1
rus I1 slightly longer than outer ramus; number of Scalpellum discoveryi Gruvel, 1906, p. 271; 1907a,

L l 0 rnm
BL0.l
L0.05
D O .O 5
Fig. 61. Litoscalpellum discoveryi (Cruvei) : A, view of right side of juvenile female; B. labrum
and palps; C, mandible; D, enlarged view of third tooth and inferior angle of mandible; E,
maxilla I ; F, maxilla 11. Eltanin, Sta. 439.
p. 2, figs. 4-6; Nilsson-Cantell, 1928, p. 11, fig. 5; Remarks: Because of the negative allometry of the
1931, p. 7 ; 1939, p. 230, fig. 2. plates during ontogeny some confusion has surrounded
ScalpelEum (Arcoscalpellum) discoueryi: Borradaile, the identity of this species. Moreover, the absence of
1916, p. 128, fig. 1 ; Bage, 1938, p. 5. succinct trophic and cirral descriptions of these ap-
Scalpellurn (Arcoscalpellum) nymphonis Borradaile, pendages has also served to compound this problem.
1916, p. 129, fig. 2. The specimens Borradaile (1916, p. 129) described, as
Diagnosis: Roof of carina evenly rounded, nonsul- Scalpellum nymphonis from McMurdo Sound, belong
cate; inframedian latus narrowly triangular, higher to the present species. Although the inframedian latus
than adjoining plates in juvenile stage but lower in of the present specimens appears to be somewhat dif-
adult; rostrum in juvenile triangular, basally acute ferent from that described for this species, it can be
with umbo apical, in adult slender, triangular with demonstrated that the difference is ontogenetic, one of
acute apical umbo, always higher than adjoining mar- the many forms the plate passes through before reach-
gins of rostra1 latera; mandible with 4 teeth including ing the level of development illustrated by Gruvel
slightly receding serrate inferior angle; maxilla I with (1907a, fig. 4 ) .
deep, prominent medial notch in cutting edge; maxilla The caudal appendage is atypical for this genus in
I1 slightly trilobate; caudal appendage uniarticulate being uniarticulate and extremely small. The append-
and shorter than first segment of pedicel of cirrus VI; age does not bear large setae, and because it is later-
dwarf male sack-like, with 4 calcareous plates. ally compressed, it is relatively difficult to locate at the
base of cirrus VI, as in A. uitreum (fig. 47C).
Material: Eltanin, Sta. 439, north of Brabant Island,
Litoscalpellum discoueryi has been reported from
Palmer Archipelago (63'515, 62 O38'W), 128-165
several localities around the Ross Sea and South Geor-
meters, 1 specimen.
gia. The present record from the Palmer Archipelago
Depository: USNM Cat. No. 125286.
reduces the apparent gap in its distribution along
Supplementary Description: The single juvenile speci- the Pacific perimeter of Antarctica. The distribution
men measures: total height 7.5 mm, capitular height of L. discoueryi may in large part depend upon the
4.6 mm, capitular width 2.7 mm. Cirral counts for occurrence of large pycnogonids, on which it com-
this specimen are given below: monly occurs.
Genus Mesoscalpellurn Hoek, 1907
Diagnosis: Capitulum of adult female with 14 re-
duced calcareous plates; tergum shaped like an in-
verted V ; scutum with basal margin entire, never hol-
lowed out, with diminutive to very short apicolateral

S Y S T E M A T I C ACCOUNT 119
arm which may exceed !/I2 length of extremely narrow 2-3 rows of short, stiff bristles; setation of cirri III-
tergal margin; upper latus triangular, with long apical VI ctenopod, with 4 pairs of setae (distal setae paired
extension, never with depending basal arm; carinal on inner ramus, but a cluster of 4 unequal setae on
latus horn-shaped, umbo at basicarinal angle; infra- outer ramus) ; caudal appendage multiarticulate and
median latus narrow, rectangular or hourglass-shaped shorter than pedicel of cirrus VI.
with umbo submedial; rostra1 latus wider than high, Material: Albatross, Sta. 2731, type-locality, north-
carinal margin less than twice the height of inframe- east of Cape Hatteras, North Carolina (36'45'N,
dian latus; caudal appendage moderately long, 5 or 7E028'W), 1427 meters, 2 specimens; Sta. 2741, off
more segments. Chesapeake Bay, Maryland (37 '44'N, 73 '57'W), 1557
Type-species : Scalpellum javanicum Hoek, 1907b ; meters, 1 specimen; Sta. 2196, south of Nantucket Is-
by subsequent designation, Pilsbry, 1908, p. 110. land (3!1°35'N, 69'44'W), 2228 meters, 1 specimen.
Remarks: The genus as presently defined contains the Depository: USNM Cat. No. 32914 (holotype and
following described species: paratype; Sta. 2731) ; 12891 (paratype; Sta. 2741) ;
M. javanicum (Hoek) , 1907b, p. 78. 9007 (paratype; Sta. 2196).
M. imperfectum (Pilsbry), 1907b, p. 75. Supplementary Description: Female-A dissected
M . sanctaebarbarae (Pilsbry) , 1907b, p. 77. specimen from the same locality as the holotype meas-
M. cochlearium (Hiro) , 1933, p. 42. ured: total height 25.4 mm, capitular height 19.1 mm,
Since the section Mesoscalpellum was described by capitular width 12.0 mm. Cirral counts for this speci-
Hoek in 1907b, it has not been widely regarded as hav- men are:
ing even subgeneric status. Mesoscalpellum contains a
cluster of species having an intermediate degree of
reduction of the capitular armament, between Litoscal-
pellum and Neoscalpellum. In attempting to subdivide
the Scalpellidae into more manageable groups, we find
it both convenient and biogeographically illuminating
to consider Mesoscalpellurn a genus (chart 7 ) . It may
be distinguished by the relatively unspecialized nature Remarks: This species is widely distributed, having
of the capitular plates, in contrast to Gyrnnoscalpellum been reported from the eastern coast of the United
and Neoscalpellum, and the greater specialization of States (Pilsbry, 1907b, p. 75), south of Iceland
the plates than found in Litoscalpellum. (Broch, 1953, p. 9 ) , off the Cape of Good Hope,
Africa (Barnard, 1924, p. 47), off Equatorial Guinea
Mesoscalpellurn imperfecturn (Pilsbry, 1907)
(Spanish Guinea before December 20, 1963), Africa
Text-fig. 62
(Stubbings, 1961, p. 1 2 ) , and off the Galapagos Islands
Scalpellurn imperfecturn Pilsbry, 1907b, p. 75, fig. (MacDonald, 1929, p. 537). This last record is in
30, pl. 4, fig. 15-18; Annandale, 1913, p. 233; Bar- need of confirmation since the specimen reported
nard, 1924, p. 47; MacDonald, 1929, p. 537; Broch, might be either a new species or M. sanctaebarbarae.
1953, p. 9; Stubbings, 1961, p. 11, text-fig. 2; 1967, The bathymetric range of M. imperfectum is from
p. 234. less than 600 meters to more than 2400 meters, but
Scalpellum (Mesoscalpellurn) imperfecturn: Pilsbry, most commonly around 1500 meters.
1908, p. 110.
Mesoscalpellum sanctaebarbarae (Pilsbry, 1907)
Diagnosis: Roof of carina flat, bounded by angles
Text-fig. 63
or low ribs; tergum bifid; scutum with short apico-
lateral extension; upper latus pyriform; inframedian Scalpellum sanctaebarbarae Pilsbry, 1907b, p. 77,
latus higher than adjoining margins of latera, rectan- fig. 31; Annandale, 1913, p. 233; Barnard, 1924,
gular with submedial umbo somewhat elevated above p. 47; MacDonald, 1929, p. 534; Stubbings, 1961,
general surface of capitulum; crest of labrum armed p. 12.
with about 80 teeth; mandible with 4 teeth including Scalpellurn (Mesoscalpellum) sanctaebarbarae: Pils-
slightly receding, serrate, inferior angle; maxilla I bry, 1908, p. 110.
with slight medial notch; maxilla I1 triangular and Scalpellum bigelovii MacDonald, 1929, p. 533, pl. 2,
trilobate; posterior border of cirri 11-VI armed with fig. 2; Stubbings, 1961, p. 12.

Fig. 62. MeaoscaZpellum imperjectum (Pilsbry) : A, mandible; B, enlarged view of third tooth
and infrrior angle of same mandible; C, maxilla I ; D, maxilla 11; E, palp; F, intermedidte article
of cirrus VI; G, caudal appendagr. Albatross, Sta. 2731 (paratype).
Diagnosis: Roof of carina flat, with bordering an- tles; distal setal arrangement on outer ramus of inter-
gles; tergum bifid; scutum with large apicolateral mediate articles of posterior cirri normal; caudal
arm; upper latus 5-sided, but essentially triangular appendage multiarticulate, taller than pedicel of cirrus
with long apical extension; inframedian latus long, VI.
narrow, umbo suprabasal, and plate higher than ad- Material: Albatross, Sta. 2839, type-locality, off Sarita
joining margins of latera; rostrum small, triangular; Barbara Island, California (28°08'N, 118°40'W),
crest of labrum armed with about 45 teeth; mandible 757 meters, 8 specimens; Sta. 2980, northwest of
with 4 teeth, including receding serrate inferior angle; Santa Barbara Island, California (33O49'45''N, 119'
maxilla I without notch in cutting edge; maxilla I1 24'30"W), 1102 meters, 1specimen.
subquadrate in outline; posterior curvature of cirri Depository: USNM Cat. No. 32915 (holotype and
11-VI armed ~ i t hnumerous rows of short, stiff bris- paratypes; Sta. 2839) ; 40288 (paratype; Sta. 2980).

Supplementary Description: A dissected paratype relationship of Pilsbry's species, M. sanctaebarbarae

from the type-locality measured: total height 27.7 mm, and M. imperfectum, and have suggested they are
capitular height 20.6 mm, and capitular width 14.2 synonymous. Annandale (19131, who had the oppor-
mm. Cirral counts are given below: tunity to study one of the paratypes of M. sanctaebar-
I I1 111 IV V VI Ca barae, did not clarify its affinities with his species,
Scalpellurn gruvelii, but he implied it was a senior
7 17 22 24 27 27 7
- - - - synonym of both of Pilsbry's species. However, there
11 21 23 24 24 26
are several gross morphological differences which tend
8 17 22 27 26 26 7 to distinguish Pilsbry's two forms. S. gruvelii is also
- - - - -
11 19 25 25 28 28 distinct and fits within the definition of the new genus

Remarks: Several workers have speculated on the Annandaleum described herein.
Fig. 63. Mesoscalpellurn sanctaebarbarae (Pilsbry) : A, right mandible; B, third tooth and inferior
angle of same right mandible; C, enlarged view of third tooth and inferior angle of left mandible;
D, maxilla I ; E, maxilla 11; F, palp; G, intermediate article of cirrus VI; H, caudal appendage.
Albatross, Sta. 2839 (paratype).

Comparison of the Atlantic and eastern Pacific wide, vase-shaped, umbo submedial to basal; rostral
species indicates that the arthropodal structures are latus wider than high, and commonly less than twice
similar, and in all likelihood they are closely related. the height of inframedian latus; caudal appendage
Both species possess short, stiff bristles found along relatively long, with 5-11 segments.
the posterior curvature of cirri 11-VI, but in M. im- Type-species. Scalpellum subflavum Annandale,
perfectum these are limited to only a few rows, whereas 1906.
in M. sanctaebarbarae there are numerous rows. The Etymology: This genus name honors Thomas Nelson
distal cluster of setae on the intermediate articles of Annandale, 1876-1924, founder and Director of the
the outer rami of the posterior cirri of M. sanctaebar- Zoological Survey of India, Superintendent of the In-
barae is normal, but it is hypolasiopod in M. imper- dian Museum, and long time student of the Cirripedia.
fectum. In the latter, the caudal appendage has 4-6 Remarks: The genus Annandaleum as defined, in-
segments rather than 7, and it is shorter rather than
cludes the following species:
longer than the pedicel of cirrus VI. The cirral counts
A. subflavum (Annandale), 1906, p. 397 (syn.:
of these two species do not vary sufficiently to be of
Scalpellum polymorphum Hoek, 1907b, p. 80; S . mol-
diagnostic value at this time. Perhaps an additional
liculum Pilsbry, 1911, p. 68; S . japonicum biramosum
point of difference between these two species is that
Pilsbry, 1911, p. 68).
the crest of the typically bullate labrum in M. sanctae-
barbarae is armed with about 45 teeth, about y2 the A. japonicum (Hoek), 1883, p. 67 (syn.: Scalpellum
number found in M. imperfectum. japonicum metapleurum Pilsbry, 1907c, p. 360; S .
curiosum Hoek, 1907b, p. 79; S . chitinosum Hoek,
Stubbings (1961, p. 1 2 ) , in his discussion of the
validity of M. imperfectum and M. sanctaebarbarae, 1907b, p. 73).
contended that, "If these species are eventually merged A . lambda (Annandale), 1910a, p. 115 (syn.: Scal-
into the synonymy of Sc. gruveli [sic] Annandale pellum longius Annandale, 1913, p. 234).
(1906), then Sc. bigelovii MacDonald (1929) also A . gruvelii (Annandale) , 1906, p. 390 (Scalpellum
must be so merged for in many points it bridges the gruvelii var. quadratum Annandale, 1906, p. 391).
gap between Sc. imperfectum and Sc. sanctae-bar- Annandaleum exhibits highly specialized capitular
barae." Although there is no compelling reason to armor, the most diagnostic feature of which is the vase-
synonymize Pilsbry's species, there are reasons for shaped and extremely large inframedian latus. By
placing Scalpellum bigelovii in synonymy with M. this characteristic Annandaleum can be distinguished
sanctaebarbarae. froni all other scalpellids. Annandaleum differs from
Scalpellum bigelovii was dredged by the Albatross Litoscalpellum in the capitulum being less heavily
from 1643 meters off Isla de Malpelo, west of Buena- armored, in possessing an inverted V-shaped tergum,
ventura, Colombia (43O03'N, 81'31'W). It is similar in the scutum having a shorter apicolateral arm, and
to M. sanctaebarbarae in the shape of certain capitu- the upper latus always lacking a depending arm.
lar plates, in particular the scutum, and the upper, Mesoscalpellum also possesses an upper latus that al-
inframedian and rostral latera. In each of these there ways lacks a depending arm, but the scutum bears a
is a point for point correspondence with the same very short apicolateral arm.
plate in M. sanctaebarbarae, in addition to the overall Annandaleum holds its greatest similarities with
facies similarity and the position of the umbo of the Litoscalpellum and Mesoscalpellum, but the relation-
carina. At the present time we can find no compelling ships remain obscure. Perhaps with further refine-
reason not to merge these two taxa. ment and reorganization of this complex assemblage,
it will be possible to decide between convergent and
Annnndaleum gen. nov. phyletic evolution. Presently there is too little data
Diagnosis: Capitulum of female with 14 partially cal- on this deep-water complex, as is obvious when one
careous plates; tergum in shape of inverted V; scutum considers that all material, of which we have seen but
with moderately long apicolateral arm about 1/4 to I,$ a fraction, constitutes about one specimen per million
length of tergal margin; basal margin of scutum en- square miles of open ocean.
tire; upper latus pentagonal to triangular to subrec-
tangular, with or without short depending arm; carinal Genus Scalpellum Leach, 1817
latus as high as or slightly higher than wide, umbo Diagnosis: Capitulum of females armed with 14 fully
at basicarinal angle; inframedian latus higher than or partially calcified plates; carina angularly flexed

A Annondaleum subflavurn (Annandale) Annondaleum lambda (Annandale)

Annondaleum /aponl'curn (Hoek) + Annandaleum gruvell'i (Annandale)
Chart 8. Distribu~ionof Annnndalrr~mgcn. nov.
at umbo, or rarely simply bowed with umbo subapical capitular plates; carirla angularly flexed, umbo at
to apical; tergum normal or forked, with umbo apical; angle; tergum commonly in form of inverted V; umbo
scutum with umbo apical, or removed fram apex along of scutum subapical, on occludent ma1 gin ; rostral latus
occludent margin ; caudal appendages common1y uni- nider than high; inframedian latus small, pentagonal,
articulate; dwarf males sack-like, not divided into umbo medial or submedial; mandible with more than
capitulum and peduncle, with or without 4 rudimen- 5 simple teeth; male sack-like, with calcareous plates.
tary calcareous plates. Species included in group:
Type-species: Lepas scalpellurn Linnaeus, 1767 (= S. scalpellum (Linnaeus, 1767)
Scalpellum vulgare Leach, 1 G 1 8 = Scalpellzcrn scal- S. stearnsi Pilsbry, 1890
pdlurn (Linnaeus) ,1767). S. stearnsi inerme Annandale, 1905
Remarks: It has long been recognized that there are S. pfeifleri Weithoier, 1E87
several species groups within the genus Scalpellurn Group of Scalpellurn gibburn
(cf. Pilsbry, 1907b; Broch, 1924, 1947; Withers, Diagnosis: Small scalpellids with capitulum com-
1953), which may eventually be recognized as genera. pletely covered by calcareous plates; carina angularly
These groups are defined after the following key: flexed, umbo at angle; tergum normal, never in form
of inverted V; umbo of scutum subapical, on occludent
margin; rostral latus er~uiclimensional; inframedian
1. ~ l m b oof scutum apical ............................. 2
1. llmbo of scutum subapical, on occludcnt margin ...... 3 latus large, subpentagonal to ovate umbo supramedial;
2. Rostral latus higher than witlc; umbo of carina medial mandible with less than 5 serrate teeth; male sack-like,
................................Group of S. stroemii without calcareous plates.
2. Rostral latus wider than high; umbo of carina not
Species included in group :
medial ......................... .Group of S. formae
3. Infrarncdian latus horn-shapcd, umho basal.. .......... S. gibburn Pilsbry, 1907b
.............................. .Group of S. magnum Group of Scalpellurn stroemii
3. Inframedian latus pentagonal, umbo medial ........... 4
4. Unrbo of carina not rnedial ...... .Group of S. anglicum
Diagnosis: Small scalpellids with capitulum fully or
4. llmbo of carina mcdial ............................. 5 partially covered by calcareous plates; carina angu-
5. Mandible with more than 5 simple tccth; males with larly flexed, with umbo at angle; tergum normal, never
calcareous plates ............Group of S. scalpellum in form of inverted V; umbo of scutum apical; rostral
5. Mandible w i ~ hless than 5 serrate teeth; males without
latus higher than wide; inframedian latus large, penta-
calcareous plates .............. .Group of S. gibbum
gonal to rectangular, umbo submedial to basal; man-
Group of Scalpellurn scalpellurn dible with less than 5 simple teeth; males sack-like,
Diagnosis: Large scalpellids u ith reduced calcareous u ithout calcareous plates.

Scolpe//urn vonhoffeni Gruvel Brochio bulofo gens et sp. nov.

+ Sco/pe/lurn gibberurn Aurivi l l ius o Srni/iurn ocufurn (Hoek)
A Ausfrolsco/pel/urn schizomofoplocinum gen, ef sp. nov A Verruco gibboso Hoek
Chart 9. Distribution of antarctic and subantarctic Scalpellum, nlonotypic Aust~alscalpell~~mgen. nov., monotqpic subant-
arctic Brochia gen. nov., Smilium acutum and Verruca gibbosa.
Species included in group : Group of Sealpellum magnum

S. stroernii Sars, 1859 Diagnosis: Large extinct scalpellids with capitulum
S. stroemii substroemii Pilsbry, 1907 fully or partially covered by plates; carina angularly
S. stroernii latirostrum Pilsbry, 1907 flexed, with umbo at angle; tergum normal or with
S. gibberum Aurivillius, 1892 hollowed-out basal margin, never in form of inverted
S. vanhoffeni Gruvel, 1906 V; umbo of scutum subapical on occludent margin;
S. ornatum (Gray, 1848) rostral latus wider than high; inframedian latus large,
S. faurei Barnard, 19241 horn-shaped, umbo basal.
Group of Scalpellurn anglicum Species included in group :
Giagnosis: Small to moderately large, extinct scal- S. magnum Darwin, 1851
pellids uith capitulum fully or partially covered by S. sigmoideurn Withers, 1953
calcareous plates; carina simply bowed, not angularly S. moraviense Withers, 1953
flexed, with umbo subapical to apical; tergum normal, S. studeri Tieche, 1905
never in form of inverted V; umbo of scutum sub- S. burdigalense Des Moulins, 1875
apical, on occludent margin; rostral latus wider than S. moliniar~urnSeguenza, 1876
high; inframedian latus large, pentagonal, umbo S. lovisatoi De Alessandri, 1895
central. S. dalpiazi Withers, 1953
Species included in group : S. nettlebladti Noelting, 1886
S. anglicum Withers, 1953
Group of Scalpellurn formae
S. fischeri Bertrand, 1891
S. fischeri costatum Withers, 1953 Diagnosis: Moderately large, extinct scalpellids with
S. bartonianurn Withers, 1953 capitulum fully or partially covered by calcareous
S. robusturn Reuss, 1864 plates; carina simply bowed, not angularly flexed, with
S. hungaricum Szorenyi, 1934 umbo subapical; tergum normal, never in form of
S. nauckanum Reuss, 1862 inverted V; scutum with umbo apical to subapical;

rostra1 latus wider than high; iriframedian latus large, [reness, Patagonia (.52"45'S, 60°34'W), 92 meters, 3
0
essentially pentagonal, umbo supramedial. specimens; Sta. 339, south of East Falkland, Falklarid
Species included in group : Islands (53"05'S, 5g031'W I , 512-586 meters, 2 speci-
S. formae De Alessandri, 1895 mens; Sta. 377, western edge of Falkland Trough,
S. avenionense Joleaud and Joleaud, I01 b southeast of Falkland Islands (53"57'S, 55"54'W),
The genus Scalpelll~rnhas long been considered of 1879-1:lttO meters, 7 specimens; Sta. 969, southeast of
more recent origin than Arcoscalpellum, from mhich Kio Grande, Tierra del Fuego (54"56'N, 65"03'W),
it no doubt evolved. It is riot surprising, therefore, to 220-265 meters, 1 specimen; Sta. 981, east of Punta
note the presence of an evenly 1)owed carirla in the de Arenas, Tierra del Fuego (52'44'S, 6704ZrW),
Eocene forms (cf. Withers, 1953) corrtained iri the 4 0 4 9 meters, 12 specimens.
group of Scalpellurrr anglicurn and S. formuc,. It could Deposi/ory: USNM Cat. No. 125287 (Sta. 217) ;
he argued that Iry the shape of the carina these forms 125288 (Sta. 222) ; 125289 (Sta. 337) ; 125200 (Sta.
might better be included in the genus Arcoscalpellzcm. 330) ; 125291 (Sta. 377) ; 125292 (Sta. 969) ; 125203
However, the position of the umbo of the scutum and r Sta. 981 i.
shape of the plates in the lower whorl are more like
Supplementary Ilescription: The gross morphology of
those of the recent species. In Arcoscalpellum, as well
this species has been described in some detail by
as the partially armored gl oups (M~~soscalpellum, Neo-
Nilsson-Cantell (1921, 1930b), and nothing further
scalpellurn, Gymnoscalpellunr, etc.), the umbo of the
need be added at this time. The arthropodal struc-
scutum is never subapical, and this alone should serve
tures, however, are described below. Female-Labrum
to separate them.
bullate; broadly triangular; periphery and superior
Scnlpellum gibberum Aurivillius, 1892 surface covered with short, stiff setae (fig. 64G) ; crest
Plates XI1 A, X111; Text-hg. 61 lacking dentition. Palps cylindrical, narrow, elongate;
Scalpellurn gibberun~Aurivillius, 1892, p. 123; 1894, superior margin free of setae; inferior margin with
11. 50, pl. 4, figs. 3, 4; Weltner, 1Ll97, p. 248; 189811, a few long setae; extremity with a strong cluster of
p. 4; 1900, p. 305; Gruvel, 1902b, p. 244; 1905, p. 40, long setae. Mandible with 3, rarely 4, teeth, excluding
fig. 40; Pilsbry, 1907b, p. 18; Nilsson-Cantell, 1921, inferior angle; second tooth well separated from first;
11. 178, figs. 24, 25; 1930b, p. 228, figs. 1, 2 ; 1939, third tooth equidistant between second tooth and in-
p. 227; Kriiger, 1940, p. 262, fig. 258; Nilsson-Cantell, ferior angle, number of teeth of inferior angle 1 to 7
1957, p. 14; Zevina, 19644, p. 252. (fig. 641, J ) . Maxilla I with extremely deep, very
Scalpellum calcaratum Aurivillius, 1U94, p. 48, pl. 4, shallow notch, or no notch (fig. 64K). Maxilla I1
figs. 5, 6 ; Weltner, 1897, p. 247; Gruvel, 1902b, ovotriangular; marginal and lateral setae with setules;
p. 244; 1905, p. 41, fig. 41; Pilsbry, 1907b, p. 14. superior margin free of setae; maxillary lobe short,
Scalpellum patagonicum Gruvel, 1900a, p. 188; broad, free of setae (fig. 64JA).
1O02h, p. 236, pl. 12, figs. IA, 16; 1905, p. 45, fig. Cirrus I about 11'~ to 36 length of cirrus 11; anterior
48; Pilsbry, 1907b, p. 18. ramus slightly shorter than posterior ramus; inter-
Diagnosis: Capitulum of adult partially covered with mediate articles strongly protuberant; faces of articles
calcareous plates, without prominent ornamenlation; clothed with stout setae. Cirrus I1 not greatly modi-
inframedian latus high and rectangular to low ant1 Led, approximating cirri 111-VI. Cirri 11-VI essen-
square or ovate, umbo basal to suprabasal; rostrum tially equal in length and with equal or nearly equal
large, rectangular, clearly exposed, umbo medial; man- rami. Setation ctenopod, 3 pairs; 1 or 2 short setae
dible with 3 teeth and a serrate inferior angle; maxilla at bases of each major pair. Each articulation along
I with deep subapical notch in cutting edge; 2 pairs greater curvature of intermediate articles of cirri 11-VI
of filamentary appendages on prosoma; peniform fila- with 1-2 long and 1-2 significantly shorter setae;
mentary appendage dorsal to anal opening; dwarf interarticular areas along greater curvature free of
male sack-like, lacking calcareous plates. setae; inner lateral faces of proximal and intermediate
Material: Eltanin, Sta. 217, Estrecho de Le Maire, articles with centrally aligned setae.
Tierra del Fuego (54'22'S, 64O42'W), 106-110 Cirral counts for each of the specimens dissected
meters, 125 specimens; Sta. 222, east of Punta de from 5 different stations are given below. The counts
Arenas, Tierra del Fuego (53O 1 5 5 , 66' 5 I f W ), 79-80 are preceded by measurements (in mm) of the dis-
meters, 175 specimens; Sta. 337, east of Punta Dun- sected specimen :

Station 217 Station 337

Total Capitular Capitular Total Capitular Capitular
Specimen Height Height Width Specimen Height Height Width
1 27.3 17.6 11.6 20.4 13.5
2 31.1 19.5 13.0
3 19.8 15.7 10.7
Specimen 1 6 13 15 15 15
- - - - -
8 15 15 16 16
Specimen 2 6 13 16 16 18
- - - - -
10 15 16 xx 18
Station 339
Specimen 3 6 13 15 15 16
- - - - -
8 14 15 16 16 Total Capitular Capitular
Height Height Width
--
7.6 4.7
Specimen 1
Station 222
Specimen Height Height Width
1 32.7 18.1
2 18.8 11.1
3 28.2 16.6
Station 377
Specimen 1 6 11 14 15 Specimen Height Height Width
-- --
- - - -
7 12 14 14 1 14.1 9.9
2 12.2 7.9
Specimen 2 6 12 13 16 Specimen 1 5 10 13 13 13
- - - - -
9 13 16 xx 7 11 13 xx 13
Specimen 3 6 13 14 xx Specimen 2 5 10 13 13 13
- - - -
8 13 15 15 7 12 13 13 14

Sl STEItlATIC ACCOUNT
Fig. 64. Scalpellurn gibberurn Aurivillius: A-F, carinal and right lateral views of thrce different size
classes of females; 6, lahrum; H, palp; I, enlarged view of third tooth and inferior angle of mandible;
J, same mandible; K, maxilla I ; L, maxilla 11; M, cirrus I; N, intermediate articles of both rami of
cirrus V I ; 0,pedicel of cirrus VI showing relationship of caudal and post-anal filamcntary appendages; P,
front view of postanal filamentary appendage. Eltanin, Sta. 217.

Caudal appendage ovoconical in section, laterally meters, and is thus the deepest k n o ~ nspecies of
compressed; anterior and posterior borders armed with Scalpellun~.
very short setae; inner and outer faces bear long, stout Nilsson-Cantell (1921) noted the presence of 2 pairs
setae; appendage slightly shorter than pedicel of cirrus of filamentary appendages on the prosoma. These are
VI (fig. 640). commonly found interwoven betneen the brooding em-
Peniform filamentary appendage dorsal to anal open- bryos in the mantle cavity, and thus appear to function
ing faintly annulated, with short stiff sparsely scattered in reproduction rather than as respiratory organs of
bristles (fig. 640, P ) ; appendage varying in length the adult. In addition to these. there is a small append-
from about the length of the first srgment of the age, previously seen in no other lepadomorphan, situ-
pedicel of cirrus VI to about 254 times the lcv~gthof ated postanally on the thorax, and it is of an unknown
the first segment. function.
Two filamentary appendages on each side of pro-
soma; longer than rami of cirrus VI, very thin, and Scalpellum vanhoffeni Gruvel, 190%
interwoven between larvae iri mantle cavity. Plate XTI C, Text-fig. 65
Remarks: Scalpellum gibberurn is perhaps the best Scalpellurn vanhoffeni Gruvel, 1907b, p. 158; 1900,
known species in the genus. It is limited to the south- p. 202, pl. 23, figs. 1-3, pl. 25, figs. 3-9.
ern tip of South America and adjoining islands, at Diagnosis: Capitular plates approximate, ornamenled
depths of less than 50 meters to more than 1800 with prominent radial ridges; inframedian latus essen-
Fig. 65. Scalpellr~mvanhoffeni Gruvel: A, view of right side of female; 8, view of rostrum, ad-
joining rostra1 latcra and scuta; C, lahrum and right palp; D, mandible; E, maxilla I ; F, n~axilla
11; G , interrxiediate article of cirrus VI; H, caudal appendage. Eltanin, Sta. 1084.

tially quadrate, with medial umbo elevated well above armored with imbricating plates about twice as wide
general surface; rostrum clearly exposed, with apical as high, exposed margins gently convex and slightly
umbo; peduncle covered with tightly packed, imbricat- irregular, about 10-12 rows, 8-10 plates each.
ing scales; mandible with 5 teeth including simple in- Measurements of the single specimen are as follow7s:
ferior angle; maxilla I with 4 spines above subapical total height 8.2 mm, capitular height 5.4 mm, capi-
notch ; maxilla I1 triangular in outline ; caudal append- tular width 3.4 mm.
age uniarticulate. Labrum bullate; periphery and anterior portion with
Material: Eltanin, Sta. 1084, north of Coronation ctenoid scales (fig. 65C) ; crest apparently lacking
Island, South Orkney Islands (60°22'S, 46'50'W), dentition; palps elongate-triangular; superior margin
298-403 meters, 1specimen. with a few short, acicular spinules; 2 or 3 long, stout,
distal setae. Mandible with 5 short, sharp teeth in-
Depository: USNM Cat. No. 125294. cluding inferior angle (fig. 65D) ; first and second
Supplementary Description: Female-Capitulum cov- teeth well separated; third equidistant between second
ered with calcified plates, rectangular in outline, cuticle and fourth; inferior angle short, overhung by fourth
very thin and not noticeably hirsute; plates orna- tooth. Maxilla I with 2 long, stout and 1-2 short, stout
mented with unequally spaced, low growth lines and spines above deep notch; 9 or 10 spines below notch
radiating ridges; occludent and carinal margins (fig. 65E). Maxilla I1 roughly triangular in outline;
slightly convex (pl. XI1 C ) . Tergum somewhat tri- marginal setae sparse (fig. 65F).
angular ; distal extremity acuminate; occludent and Rami of cirrus I essentially equal in length; inter-
basal margins convex; carinal margin concave; umbo mediate articles not noticeably protuberant; inner
apical; faint ridges radiate from umbonal region. Scu- faces lightly clothed with long setae on distal ys of
tum 34 higher than broad; basal and occludent mar- each segment. Cirrus I1 separated from cirrus I and
gins slightly convex; tergal and lateral margins slightly resembling more cirri 111 and IV. Each articulation
concave; umbo apical; thin, unequally spaced, in- along greater curvature of intermediate articles of cirri
clined ridges emanate from umbonal region. Carina 11-VI with 1 long seta and 1 extremely short spine:
typical with strong, angular flexure above middle of interarticular areas along greater curvature without
plate; 2 parallel ridges on each side of plate emanating setae; inner lateral faces of intermediate articles ran-
from umbonal region, setting off tectum from parietes domly setose. Setation ctenopod (fig. 65G) ; 3 major
and parietes from intraparietes. Upper latus of 5 sides; and 1 minor pair on cirri 11-VI; 1 or 2 short setae
scutal, tergal and carinal margins essentially of equal situated near bases of major setae. The cirral counts
length and straight; umbo apical; radiating ridges thin were as follows :
and low. Carinal latus essentially triangular, but with
5 unequal sides; carinal margin concave; basal margin
convex; apex projecting beyond carinal border; umbo
at extremity of projection; apices of carinal latera ap-
proximate below basal margin of carina; radiating
ridges moderately broad, rounded, unequally spaced.
Inframedian latus with 5 sides, hut essentially rectang-
ular in outline; basal margin twice width of upper Caudal appendage uniarticulate (fig. 65H) ; ap-
margin; lateral margins parallel and nearly of equal proximately 3i length of both segments of pedicel;
length; plate extremely thick and high, i.e., elevated supporting an apical tuft of about 3-5 short, strong
above level of rest of capitulum; umbo above center of setae; anterior and posterior margins clothed with
plate; ridges radiating from umbonal region high, short acicular setules.
broad, irregular. Rostra1 latus 4-sided, but roughly Remarks: Scalpellum vanhofeni was dredged several
triangular in outline; scutal and lateral margins of times at the Gauss winter station off Gaussberg on
equal length; rostral margins about 1/3 length of basal Wilhelm I1 Coast (approximately 66OS, 89OE), and
margin; umbo on rostral edge of plate; surface ridges briefly described by Gruvel in 1907b, and subsequently
irregular, low, moderately broad. Rostrum small; par- in detail by the same author in 1909. I t is one of the
tially hidden by rostral latera and scuta; broadly tri- small species in the genus, and is closely allied to the
angular; umbo apical (fig. 65B). group of S. stroemii Sars. This group is readily char-
Peduncle about or less height of capitulum; well acterized by the radially sculptured capitular plates,

large rectangular inframedian latus with the umbo rate Australscalpellurn from other ScalpelIidae is the
elevated well above the general level of the capitulum, unusual placement of the inframedian latus. In the
umbo of the scutum apical, and the peduncle heavily adult form it is inserted between the upper latus and
armored. These characteristics and differences in the the scutum. This is compensated for by the lower por-
trophic structures will probably require species of this tion of the upper latus depending between the infra-
group to be distinguished as a subgenus of Scalpellum. median latus and the basilateral margin of the scutum.
Aside from the unusual position of the inframedian
Australscalpellurn gen. nov. latus, the unique V shape of this plate as well as the
hourglass shape of the upper latus should serve to
Diagnosis: Capitulum of female with 13 calcified, the genus from other scalpe1lidae.
approximate ~ l a t e s ,but varying degrees of ~ - e ~ - m a - The loss of the rostrum in the present genus is in-
ment evident; scutum and tergum cleft, umbos apical; teresting. It could be correlated with rearrangement
carina strongly flexed at umbo; rostrum lacking; UP- of the capitular plates resulting from rearmament, but
Per latus large, higher than with umbO at Or the rostrum in Arcoscalpellum is often reduced with no
above center; inframedian latus same as or slightly evidence for de- or rearmament.
wider than high and less than 34 height of capitulum; The unusual ontogenetic pattern of scutal and tergal
umbo of carinal latus effectively basal; rostra1 latus rearmament in A~~~~~~~~~~~~~~~~is not
higher than wide; dwarf males sack-like, without rudi- known to occur in scalpellidae other than the new
mentary calcareous plates. genus Brochia. It is found in the unrelated poecilas-
Type-species: Australscalpellurn schizmatoplacinum matids Octolusrnis and Trilusrnis (cf. Newman, 1961b,
sp. nov. pp. 104-106). In Neoscalpellum the scutum and ter-
Etymology: The generic name, derived from the Latin, gum, as well as other c a ~ i t u l a rplates are not initially
alludes to the affinities and geographic distribution. forked or cleft in juveniles. The interesting difference
between the present genus and Neoscalpellum is that
RtTLarks: The genus Austrazscaz~ezzum is proposed once unarmored the latter are not known to undergo
include one 'pecies of Sca1Pe11idae7 which rearmament. In Scalpellum and related spe-
differs from other 'pecies in the cies, the tergum is forked, but as with NeoscalPellnm
Unique morphological features of the genus are the it never rearmors. In Arcoscalpellum fissum, described
cleft or split found along the basal margin of both by Hoek (1907b, p. 116) from the region of the Moluc-
the scutum and tergum (through ontogenetic stages) 7
cas, neither the tergum nor the scutum is cleft, but
and the total lack of a rostrum (cf. fig. 66). instead the upper latus is deeply cleft.
In very small individuals the scutum and tergum It would appear that at some time in the history of
may lack clefts, but in a specimen with a ca~itular Australscalpellum selection for full capitular armament
height of 0.85 mm the notches have begun to appear. was relaxed and that exploitation of more exposed
Succeeding ontogenetic stages, as determined from a environments at a later period in its history, where
series of specimens, show a deepening and slight fuller armament would be selected for, would then
broadening of the notches in both plates. However, account for the rearmament. Predation is suggest-
after a certain point the enlarging process is arrested ed as the primary factor because it has been ob-
and a rearmament ~ h a s einitiated. The cleft in the served during the course of this study that borings
scutum gradually closes, proceeding from the lateral of rapacious gastropods frequently occur in the cal-
to the occludent margin, and is pinched off and iso- careous portions of capitular plates rather than through
lated as an elongated slit. This slit, in turn, is ~ r a d u - the chitinous interspaces (cf. Broch, 1922, p. 237).
ally closed. In all of the larger specimens thus far This suggests that these gastropods select carbonate
examined, the cleft is reduced to a short, linear rudi- areas to bore, and such selection would favor the re-
ment and, although no specimen showed complete duction by the barnacle of the area enveloped by a
obliteration, it may ultimately disappear. The basal calcareous covering concomitant with a proportionate
margins of the cleft in the tergum, on the other hand7 increase in the exposed chitinous area. This explana-
apparently do not reunite, subsequently pinch off, and tion may apply to the naked, deep-water heteralepadids
isolate the cleft, but rather apparently remain open wherein specimens have not been reported with holes
throughout the life of the individual. bored through the chitinous capitulum. Should the
Another morphological feature that serves to sepa- habitat or the predator change before a species lost

its plates completely, presumably the rearmament high as wide in large specimens but slightly wider
process would ensue. than high in juveniles; c.arina1 margin concave and
longest, followed by lateral, basal and apical; umbo
Australscnlpellum schizmatoplacinum sp. nov. situated at basicarinal angle; latera extend around and
Plate XI1 D, E ; Text-fig. 66 approximate below basal margin of carina. Irifra-
Diagnosis: lnframedian latus broadly V-shaped, with median latus higher than hide; V-shaped in adults;
occludent arm{{,j longer than caririal arm; upper latus occluderit arm inserted between scutum and upper
shaped like hourglass; region immediately below crest latus; umbonal region elevated and situated at junc-
of labrum aimed with numerous rows of M-shaped or tion of arms of V; arms widest distally, region im-
comb-like teeth; mandible with 3 teeth excluding in- mediately distal to umbo depressed. Rostra1 latus
ferior angle, superior slope of third, but rarely srcond quadrate; wider than high in juveniles, but higher
tooth serrate; maxilla 1 with deep medial notch in than wide in adults; scutal ant1 basal mar,'c r ~ nconcave;
s
cutting edge; caudal appendage with less than 2 short, lateral margin convex; occludent margin straight;
termirial setae. umho situated at middle of occludent margin.
Material: Eltnr~ir~,Sta. 410, type-locality, between Measurements (in m m ) of 25 females from the type-
Elephant Island and Aspland Island, South Shetland locality range as follows: total height 1.0-5.9, capitu-
Islands (6I018'S, 56°09'W), 220-240 meters, 25 speci- lar height 0.85-3.6, capitular width 0.60-2.7.
mens; Sta. 993, between O'Rrien Island and Bridge- Labrum bullate; long, broadly rounded distally;
man Island, South Shetland Islands (6I025'S, 56' superior surface, except for narrow zone anterior to
30'W'l, 300 meters, 1 specimen; Sta. 1003, north of crest, covered with ctenoid scales; crest and region
Joinville Island, Antarctic Peninsula (62O4IrS, 54' immediately below with numerous rows of teeth, either
43'W), 210-220 meters, 107 specimens. single or in clusters of 2 to 4 (fig. 66P). Palps mod-
Depository: USNM Cat. No. 125295 (holotype; Sta. erately long and slender; superior and inferior mar-
410) ; 125296 (paratypes; Sta. 410) ; 125297 (para- gins free of setae; distal end margins covered with
type; Sta. 993) ; 125295 (paratypes Sta. 1003). ctenoid scales. Mandible with 4 teeth including in-
Description: Capitulum with fully calcified plates, ferior angle; dislance between first and second tooth
laterally compressed, essentially quadrate in outline slightly greater than betneen second and third; infe-
(fig. 66A-L) ; height slightly greater than width; rior angle variably toothed, of 1-2 long slender or
slight membranous space between carina and adjoining 6-12 coarsely serrate teeth; superior slope of third
plates; plates marked by low, close-spaced growth tooth serrate (fig. 6 6 s ) . Maxilla I with deep, promi-
ridges only. Tergum elongate; triangular; occludent nent medial notch (fig. 66T). Maxilla I1 ovate in
and basal margins convex, carinal margin concave; outline; marginal setae not distinctly segregated into
cleft arising from basal margin; fissure widest in small clusters (fig. 66U) ; maxillary lobe low, broadly
specimens, narrowing and trending towards closure in rounded apically.
large specimens; umbo apical. Scutum essentially tri- Ramus of cirrus I subequal; intermediate articles
angular in outline; occludent margin slightly convex, equal in width, not protuberant; articles with a few
about twice length of slightly concave tergal margin; ctenoid scales parallel to distal articulations. Rami of
basilateral margin sinous; cleft arising from lateral cirrus I about length of rami of cirrus 11; cirrus
margin open in small specimens, but closed and par- I1 about % length of cirrus 111. Cirri 111-VI essen-
tially obscured in larger specimens; no specimens hav- tially equal in length with equal or slightly subequal
ing totally obliterated suture; umbo apical, slightly rami; each articulation along posterior margin of
produced and projecting over basioccludent angle of intermediate articles with 1 long, slender seta and ?
tergum. Carina strongly bent at umbo about 120°, as extremely short, stiff bristle; interarticular areas along
in Scalpellurn (sensu stricto) ; 2 high, broad ribs posterior curvature, as well as inner and outer lateral
emanating from umbonal region, bounding the flat faces free of setae and bristles, but lightly clothed
roof and extending to basal margin of plate. Upper with ctenoid scales. Setation ctenopod; 3 major and
latus in small specimens essentially 5-sided; plate 1 minor pair of setae, each major pair with 1 short,
shaped like hourglass in large specimens; scutal and slender seta at base (fig. 66V).
basicarinal margins concave and marked with depres- Cirral counts for the holotype and 3 paratypes are
sions extending nearly to the somew hat central umbo. given below. Two of the paratypes are from localities
Carinal latus essentially 4-sided; more than twice as other than the type.

Fig. 66. Australscalpellurn schizmatoplncinum gen. e t sp. nov.: A-L, right lateral view of females showing
various stages in the development of tlie scutal and tergal cleft; ill, view of upl:cr portion of capitulum
showing male cyprid entering pouch in scutum; a small portion of a second male can be seen immediately
above the nearly fully exposed male; N, 0, male cyprid and antenna I of same, respectively; P, labrum
and palps; Q, portion of crest of labrum; K, palp; S, mandible; T, maxilla I ; U, maxilla 11; V, intermedi-
ate article of cirrus VI; W, caudal appendage. Eltanin, Sta. 410 (holotype, figs P-U; paratypes, figs. H-0)
and Sta. 1003 (paratypes, A-G, V, W ) .

S\ISTEMKI'I(: AC(:OUNT 133
I 11 JIJ IV V VI Ca Uroch of Oslo, Norway, for his iririumerable coritribu-

tions to cirripedology during nearly half a century.
Bltanrn, Srdtio~r410
Holotype 5 x 9 9 1 0 1 0 1 Ren~arks: Hrochia and Az~stralscalpc~llum are remark-
- - - -
6 8 9 9 1 0 1 0 able among the Scalpellidae in having uridergorie what

is termed a rearmamerit process. While reduction in
armament is k r l o ~ r lin other groups of the family, the
reversal of this process has so far I~eerirecognized only
in these two genera. Rrochia differs from Au~/rnlscal-
pellurn in having retained rather than lost the rostral
plate. having an essentially oval rather than V-shaped
inframedian latus, and an oval rather than hourglass-
shaped upper latus. I h ~ s erhalacteristics suggest af-
finities u ith the largely Ijoreo-Arctic group of Scal-
p l l u m strorrnii Sars.
Rroch~aand ,4ustralscalpc~llurnare similar in having
cleft scuta and terga. In the former, complete fusion
and obliteration of the scutal cleft occurs ontogeneti-
Eltanin, Station 1003 cally. A u ~ ~ r a l s c a l ~ e l lhowever,
um attains a larger size
Paratype 4 8 8 8 9 1 0 1 than Brochia, yet of the more than 100 specimens ex-
5 8 8 9 9 9 amined, none were ohserved uith a totally obliterated
scutal cleft.
The depths inhabited by Brochia are far in excess
of that recorded for Australscnlpellum; the former
Caudal appendage short, broadly triangular, later- apparent1 y inhabiting the continental slope, the latter
ally compressed; consisting of a single segment about the continental shelf.
?$ to height of first article of pedicel o l cirrus
VI; apex supporting either a single or no short, ter- Brochia bulata sp. nov.
minal setae; complete appendage covered with ctenoid Plate XI1 B, Text-fig. 67
scales (fig. GGW) .
Diagnosis: Rostrum large, rectangular, umbo sub-
Male cyprids elongate and oval (fig. 66N), measur-
medial, clearly exposed; inframedian latus largest of
ing 0.25 X 0.65 mm; one cyprid found in each of the
latera, 5-sided but essentially quadrate in outline; up-
two scutal pouches (fig. 66M). Dwarf male sack-like,
per latus elongate-oval; crest of labrum armed with a
measuring about 0.20 X 0.4'5 mm, surface covered with
single row of simple conical teeth; maxilla I without
concentric bands of small spines, lacking calcareous
medial notch in cutting edge; caudal appendage with
plates.
2 or more short, terminal setae.
The specific epithet is derived from the Greek, and
denotes the split nature of the tergal and scutal l,lates. Material: Eltanin, Sta. 557, type-locality, east of
Choiseul Sound, East Falkland, Falkland Islands
Brochia gen. nov. (51°5fi'S, 50"39'W), Z55-866 meters, 2 specimens.
Diagnosis: Capitulum of female comprised of 1-1 Depository: U S N M Cat. No. 125299 (holotype) ;
calcified, approximate plates, but varying degrees of 125300 ( paratype).
rearmament evident; scutum and tergum clelt, umbos Description: Female-Capitulum subquadrate in out-
apical; carina angular; rostrum present, clearly dis- line, height nearly twice the width. Capitular plates
cernible externally; upper latus large, higher than more or less fully calcified, sculptured with unequally
mide, with umbo more or less central; inframedian spaced growth ridges and low, approximate, fine,
latus twice as high as wide and more than y-j height rounded radial ridges emanating from umbonal re-
of capitulum; carinal latus with marginal, submedial gions. Tergum narrow, triangular, basicarinal angle
umbo; rostral latus higher than wide; dwarf males acute; occludent and basal margins convex; carinal
sack-like, without rudimentary calcareous plates. margin concave ; cleft arising from basal margin ;
Type-species: Brochia bulata sp. nov. umbo apical. Scutum triangular; tergal and occludent
Etymology: The genus Brochia honors Dr. Hjalmar margins essentially straight; lateral margirl scalloped;

Fig. 67. Brochia bulata gen. et sp. nov.: A, view of right side of female; B, internal view of left
scutum; C, internal view of right scutum showing pouch for dwarf male (dotted line) ; D, external view
of upper latus; E, rostrum; F, dwarf male; G, antenna I of same male; H, crest of labrum; I, J, man-
dibles; K, maxilla I ; L, palp; M, cirrus I ; N, intermediate article of cirrus V I ; 0,caudal appendage;
P, enlarged view of caudal appendage. Eltanin, Sta. 557 (holotype, figs. A, F, G, I-N, P ) .
cleft arising from lateral margin; no evidence of split latus subpentagonal, higher than wide, umbonal region
and subsequent fusion in holotype, but in paratype slit occurring about K the height from the basal margin,
still visible both externally and internally; adductor projecting beyond margin of capitulum and laterally
muscle slightly triangular; undercut ridge below basal margin of carina. Inframedian latus larg-
parallel to tergal margin present for reception of male
est of latera, 5-sided; width y2 of height; umbo sub-
(fig. 67C). Carina angularly flexed at umbo; roof
central; apical portion inserted between scutum and
evenly rounded; no ridges separating roof from sides.
upper latus with 5 sides, but essentially ovate in out- upper latus. Rostra1 latus essentially rectangular; twice
line (fig. 67D) ; higher than wide; carinal margin as high as wide; umbo slightly below midpoint of ros-
longest; umbo more or less centrally located. Carinal tral margin. Rostrum rectangular in outline; exposed

portion hourglass-shaped, height about 3 tirnes uidth; Caudal apt)endage short, laterally compressed, sub-
umbo slight1y sub cent^ al. triangular, broad or acutely rounded; a single article
Peduncle ahout "% height of c.,~pitulum; 2-4 times about height of first segment of pet1ic:el of cirrus
u ider than high; fully armored; scales not imbricating. V I ; apex supporting 3 terminal setae, longest scta
Measuremerrts of holotjl~e: total height 5.1 mm, niore than height; (:overed c\ ilh c:~enoitlscales (fig.
capitular height 3.9 mm, capitular width 2.2 mm. 670 .
The paratype n as partiall) ciestroj erl. Dwarf male sack-like, ~ i t h o u ttraces o f calcareous
Lal-)rum hullate; extreme11 lung and rial r O M . br oatlly plates; covered uiih concentric har~tlsof short bristles;
rounded tlistally; superior surfare and margins cov- one male in each pouch (fig. 07F .
r 7
ered with ctenoid scales; crest armed with a single lhc: specific: name is deriveti from the J,atin, ant1
ron of ahout 60-65 simple, conical teeth (fig. 67H). draws attenlion to the extremely large irilramediari
T'alps model ately long, slender ; distal end at utel y latus.
rouritlecl, and supportirig a feu short setae; s u p e ~ i o r
arid inferior margins free of setae; lateral face and
margins covered nith ctenoid scales (fig. 67L). Man- Diagr~osis: Asymmetrical hox-like sessile cirriprtls
dil~lewith 4 teeth including inferior angle; distance with wall consisting of 4 plates in Receiit forms (ca-
betnren second and third tooth about I,$ distance be- r i m , rostrum, scutum and tergumi, and 6 plates in
tneen first and second; inferior angle serrate, 7 short, extinct forms (aforegoing plus carinal and rostra1
triangular. arid nariow tezth; lateral face of appendage latera) ; operculum of remaining scutum and tergum
covered M itli short, stiff setae existing singly or in forming movable lid-like top; basis meml~rarious or
clusters of 2-3 (fig. 671, J ). Maxilla I without notch calcareous.
in cutting edge; loner portion of cutting edge in- Family VEKKUCIDAE
Daru in, 1254
clinrd proximally (fig. 6719). Maxilla 11 broadly oval
Verrucidae: Withers, 1935, p. 323, s j nonymy.
in outline; marginal setation very sparse and in 2
distinc~clusters. Genus Verruea Schumacher, 1817
Anterior ramus of cirrus I about $6 lrngth of poste- Verruca: Withers, 1935, p. 340, synonymy.
rior ramus (fig. 67M) ; intermediate arlicles of both
Remarks: The genus Verruca is presently divided
rami s l i ~ h t l yprotuberant arid about equal in uidth;
into 5 subgenera, a key to which is given below.
distal end of each article supporting a few ctenoid
scales. Posterior ramus of cirrus I about I/r, length of
posterior ramus and "; length oI anterior ramus of 1. Oprrcular plates nearly parallel to base .............. 2
c i ~ r u sllT; cirri 111-VI essentially equal in length M ith 1. Oprrc:ular p l a t ~ snearly prrpendit:ular to base ........ 4
slightly suld~qualrami. Articulatioris along posterior 2. Fixetl scutum with aclductor ridge forming inyophore
tuivaLure of intermediate ar~iclesof cirri 11-VI nith
........................... Metauerruca Pilsbry, 1916
2,. Fixed scuturn with adductor ridge not forming myophore 3
1 lorig seia; interarticular areas along posterior mar- 3. Apcx of r o s ~ r u m apical .... Verruca Schumacher, 1817
gins and lateral faces free of setae; distal end of each 3. Apex of rostrum subapical. .Rostratoverruca Broch, 1922
article with few ctenoid scales. Setation ctenopod; 3 4. Fixcd scutum without adductor ridge ................
major and 1 minor pair of setae; at base of each .............................Altiverruca Pilsbry, 1916
4,. Fixctl scutum with adductor ridge forming myophore
major pair, 1-2 short, slender setae (fig. 67N).
........................ Carneraverruca Pilsbry, 1916
Cirral counts for the types are given below:
Subgenus Altiverruca Pilsbry, 1916
Altiverruca Pilsbry, 1916, p. 40.
Holotype 5 8 8 9 9 1 0 1
- - - - - - Diagnosis: Wall composed of 4 plates; top nearly
7 8 9 x x x x perpendicular to base, fixed scutum without adductor
ridge; no internal recesses of the general body cavity.
Type-species: Verruca hoeki Pilsbry, 1907b, p. 113.
Paratype Verruca (Altiverruca) gibbosa Hoek, 18G3
Plate XIV, Text-fig. 68
Verruca gibbosa Hoek, 1883, p. 134, 131. 6, figs. 17,
18, pl. 11, figs. 5-9, pl. 12, figs. 1-5; Gruvel, 1905,

Fig. 68. Yerruca gibbora Hoek: A, crest of labrum; E, palp; C, abnornial mandihlr; D, maxilla 11;
E, enlarged view of third tooth and inferior angle of mandible; F, saine mandible; G, enlarged view of
lower portion of maxilla I ; H, same maxilla I ; I, penis and basal portion of caudal appendage; J, caudal
appendage; K, light cirrus I ; L, left cirrus 11; M, left cirrus 111. Eltanin, Sta. 1067.
p. 178, fig. 195; Pilsbry, 1916, pp. 41, 44; Nilsson- Verruca bicornuta Pilsbry, 1916, p. 43, pl. 7, figs.
Cantell, 1928, p. 25, figs. 12, 13; 1929a, p. 470; Broch, 1-lc, ~ 1 8,
. figs. 3-3b, pl. 9, fig. 1 ; Nilsson-Cantell,
1931, p. 45; Nilsson-Cantell, 1938, p. 11; 1950, p. 219; 1955.13. 219.
2 L
Tarasov and Zevina, 1957, p. 154, figs. 51-53. Verruca gibbosa somaliensis Nilsson-Cantell, 1929a,
V t ~ r u c asulcata Hock, l883. P 139, p 1 figs- 1 9 7 p. 470, fig. 4; Broth, 1931, p. 45; Nilsson-Cantell,
20; Gruvel, 1905, p. 179, fig. 196; P i l s b r ~ 1916,
, p. 41.
1938, p. ll.
Verruca rnitra Hoek, 1907a, p. 5, figs. 14.
Verruca darwini Pilsbry, 1907b, p. 111, pl. 10, figs. Diagnosis: Labrum bullate; crest armed with about
4-8; 1916, p. 45; Nilsson-Cantell, 1955, p. 219. 40 simple teeth, superior surface covered with a nar-
verruca rathbuniana pilsbry, 1916, p. 41, pl. 7, row short zone of acicular spines anterior to crest and
figs. 2 4 , ; Nilsson-Cantell, 1927, p. 776, fig. 15; 1950, in turn followed by a broad zone of ctenoid scales;
p. 219. anterior ramus of cirrus I slightly longer than poste-

rior ramus; posterior ramus oI cirrus I1 much longer Cirral counts for this spec,imen are:
than auterior ramus; cirrus I11 normal.
Muterial: Ellanin, Sta. 1007, on Scotia Ridge, south
of South Georgia (59'5ifS, 34'4IfW), 1098-1153
meters. 1 specimen.
Drposi~ory: ITSNM (:at. No. 125r301
S u p p l r ~ l r ~ eyn ~ l)escrip/ion
(~~ : 1lermapl.lrodite-The
h'ernnrks: Nilsson-Cantvll (1928, 1929a) studied the
single specimen measured: basal Irr~gth E.5 meters, shell and arth~opodalmorphology of this species, as
basal \+itiih 2.0 mm, distarice l)et~$.lct.n apices of carina well as seve~alclosely related ones. As a result, lie
arid rostrum 0.9 mm, height of fixed tergum 0.:: mm, plac.ed several H ide-ranging species in synonymy u ith
height of fixed scutum 5.9 mm ( pl. XIV ) . V. gihbosa, giving the present species a worldwide
1,ab1 um bullate; short, brtrad ; superior surface sup- distribution, from depths of 500 to \$ell over 3000
porting a small, narrow zone of short acicular spines meters. The present study suggests that this synonymy
surrounded by ctenoitl scales (fig. 08A) ; crest armed should be reviewed.
with single row of about $0 simple, conical teeth. Palps
long, slender, acute1y rounded distally; superior mar- Suborder BALANOMORYHA Pilsbry, 1916
gin and distal extremity bear a few short arid long Diagnosis: Sessile cirripeds with symmetrical wall
setae; outer proximal half covered with cterioid scales. formed of 8, 6, 4 or 1 plate and an o ~ e r c u l u m(except
Mandible ~ i t h4 teeth including inferior angle; teeth in Xenobalanus) formed by the paired terga and scuta.
short, acute; fir st tooth eiiher close to or %ell separated
from second tooth; inferior angle coarsely serrate,
1. Innrr rarr~i of thirtl cirri unmotlifirtl, rescnrbling the
about 22 short to long. slender subspatulate teeth (fig. posterior rather than the anterior pairs, outer ranius
O:K, F) . Maxilla I slender, nlocler ately short (fig. sometirries antenniform; labrum bullate, crest without
08H) ; deep, broad notch occupying middle region of notch or nretlian incision; penis wittlout basitlorsal
cutting edge; above notch 1-2 long, stout and 1 short, poirrt, caudal appendages some ti arc.^ present; wall
always solid (none known from Antarctica) ..........
stout spine; below notch 10-11 short and long. stout
......................... .Chthamalidac Darwin, 1854
spines ~ h i c hmay he separated into an upper cluster 1. 1:oth rami OIthird cirri motlilic~tl, arrtenr~ilor~n and/or
of 5-7 antl a lone1 cluster of 4-6 (fig. 6:iG). Maxilla rc.scmblir~g morr the anterior than t l ~ eposterior pairs;
I1 tall, lrilolred; marginal setae sparse, segre~atedinto lahrurn not bullate, crest with or witlrout mtdian
3 clusters. ~ ~ o t c tor
i incision; pctris will1 or wittiout basitlorsal
point; caudal appcntlayes a h s c n ~ ; wall solid or per-
Anterior ramus of cirrus I slightly longer than pos- nlcatctl by 1 or more rows crt longitudinal canals.. . . 2
terior ramus (fig. 6 l K ) ; intermediate articles hardly 2. 1,abrurn with median notch or incision; ranli of thirtl
modified; both rami clo~hedmith long setae. Anterior cirri usually not antenniforrri; per~isnsually with basi-
ramus of cirrus I1 ahout :) i of posterior ramus
length dorsal point; Ualanidae Leach 1817 (no Recent
(fig. 68L) ; posterior ramus about same length as pos- antarctic: rcprescntatives) .......................... 4
terior ramus of cirrus I arid :x length of posterior 2. Lahrum without median incision; rami of third cirrus
somctimcs antenniform; penis without basidorsal
ramus of cirrus 111. Posterior 3 pairs of cirri on both point ............................................ 3
sicles except fifth pair, badly damaged. Each articula- 3. Crest of labrum without notch; 1 or both rami of second
tion along greater curvature of intermediate articles and thirtl cirri sometimes antenniform; wall always
of cirrus V with 1 long antl 1 short, thin spine; inter- without radii, solid or pvrmeatcd by a single row of
articular areas and lateral faces free of setae. Setation chitin-fillet1 longitntlinal carials ....................
ctenopod, 2 major and 1 minor pair. ......................... .Bathyllasmatidae fam. nov.
3. Labrum with notch more or less developed; rami of
Caudal appendage long, slender, of 9 articles (fig. third cirrus seldom antenniform; wall permeated hy 1
68J), probably same length as, or longer than, pedi- or morc rows of separate or confluent longitudinal
cel of cirrus VI; each articulation encircled by setae; ca~rals often partially or completely fillcd with cal-
terminal segment with 11-13 setae. carcous and rlritinous material.. ......... .Tetracliiidae
(=Tetraclitinae Gruvel, 1903 nom. transl. Ross, 1968;
Penis short, stout, slightly longer than caudal ap- none known from Antarctica)
pendage; distal end slightly bulbous; surface faintly 4. Cirri oftcn with simple hooks o r spincs, never providcd
annulated and sparsely setose (fig. 681). with grapples; rami of third cirrus nevrr antenniform;

wall solid or with 1 or more rows of longitudinal Darwin had said that it was always notched in this
tubes, never filled with chitinous material; basis genus.
usually calcareous; penis with basidorsal point. . . . . . In 1913, Hoek described two more deep-sea species,
. . .
. . . . . .. . . .. . . . . .. . .. . .. . .Balanidae, ex. Elminius
taken by the Siboga in the Indo-West Pacific, which,
4. Cirri without hooks and spines but often provided with
like those of the Challenger report, had the peculiar
bipectinate grapples; third cirrus sometimes antenni-
form; wall solid, but sometimes filled with chitinous unnotched labrum. He further noted that the longer
material; basis membranous; penis with or without ramus of the third cirrus in these species, in contrast
basidorsal point ( n o representatives south o f the to the situation in Balanus, in many respects resem-
Antarctic Convergence) . . . . . . . . . . . . . . . . . . . .Elminius bled the structure of the fourth more than the second.
In consideration of this contrast, the solid wall, and the
BATHYLASMATIDAE fam. nov. lack of radii, he proposed a new genus, Hexelasma,
ex. Balanidae, Chthamalidae, in part to which he added the aforementioned, B. callisto-
derma Pilsbry from Japan, and B. hoekianus Pilsbry
Lliagnosis: Hermaphroditic Balanomorpha with wall
from the North Pacific. However, while enumerating
of 6 or 4 plates, solid or with a single row of chitin-
the differences between the new genus and Balanus, he
filled longitudinal tubes, sheath weakly or strongly
retained the new genus in the family Balanidae.
developed but not overhanging. Rostra1 plate: pre-
In 1916, Pilsbry reviewed the status of the group
sumed compound but without vestigial sutures indicat-
and concluded that "The form of the labrum and of
ing fusion of rostrum with rostrolaterals; overlapping
the third cirrus shows at once that Hexelasma belongs
adjacent plates but without development of radii (figs.
to the Chthamalidae, and not to the Balanidae. The
69, 70). Basis membranous, or when calcareous not
armature of the mandible and maxillae, the form of
permeated by tubes. Labrum thick, without notch.
the lower edge of the sheath and the texture are also
Mandible usually with 4 teeth; inferior angle with few
characters pointing in the same direction." He com-
short simple or complex spines, never a comb (pec-
pared Hexelasma with the unquestionable chthamalid
tinate), a cluster of numerous spines, or molariform.
Pachylasma, pointing out the usual presence of caudal
Cirrus I11 intermediate in structure between cirrus I1
appendages in the latter, not to mention the clearly tri-
and IV; with inner, or inner and outer rami elongate
partite nature of the rostra1 plate, that separated the
and antenniform, or normal, even in same species.
two genera, but he otherwise considered them very
Without caudal appendages. Penis without basidorsal
similar. Hexelasma hoekianus he returned to Balanus,
point. All oceans; deep water to greater than 2,006)
as the type of a new subgenus, Metabalanus. In the
meters; Lower Oligocene to Recent.
same paper, he described an additional western At-
Type-genus : Bathylasma gen. nov.
lantic species, H. americanum, pointing out that it was
Etymology: With reference to great depths at which very closely related to his previously described H. cal-
members of the family are found. listoderma from Japan. The genus thus came to en-
compass the following species :
Discussion: In recent monographs (Pilsbry, 1916;
Kriiger, 1940; Withers, 1953) the suborder Balano- H. corolliforme (Hoek)
morpha is divided into two families; the Chthamalidae H. hirsutum (Hoek)
and the Balanidae. The chthamalids are structurally H. callistoderma (Pilsbry)
more generalized (cf. below), and appear in the fossil H. aucklandicum (Hector)
record before the close of the Mesozoic. The balanids H. velutinum Hoek
on the other hand are generally more advanced and H. arafurae Hoek
do not appear until early in the Tertiary. H. americanum Pilsbry
In 1883, Hoek described two deep-sea species and The relationship of the genus to other genera of
assigned them to Darwin's Section G of the genus the Chthamalidae was summarized by Pilsbry (1916),
Balanus (radii wanting, basis membranous, wall as indicated in table 3.
solid), as B. corolliformis taken by the Challenger near Through Pilsbry then, the genus came to reside in
the Kerguelen Islands, and B. hirsutus taken by the the Chthamalidae, being placed phylogenetically close
Triton in the "Faeroe Channel." In his discussion he to Pachylasma primarily on the basis of the compa-
was more or less reluctant to assign the two to Balanus, rable arrangement of the wall plates and nonbalanid
noting that the labrum was not notched at all; whereas, nature of the appendages, and this system has been

Tahle 3. Sunlmary of Relationsllips among the Chthamalidae (adopted f r o n ~l'ilsbry, 1916, p. 291; gencra not given or subse-
quently described included in parentheses) .
8 or 6 compartments 8 6 4
with occessorv plates compartments comportments comportments
fBathybulonusj
Hexelasmu- --- -~Tessure/usmaj
Catophragmus Pachylasmo
8 comportments
CHTHAMALID - - - - - - -.- Octomeris
SERIES
lChione/usmus)
6 compartments
I- - - - - - - - - - - - - - - Chthamalus - - - - - -.- Chomaes~pho
-- - fTetrochthamaIusj
followed since (Kriiger, 1940; Withers, 1953; Utinomi, three new species described herein, we found the wall
1965). However, Bage (1938) remarked, "From the not to be solid at all, but to be permeated by longitudi-
examination of the soft parts it is apparent that the nal tubes that were secondarily filled with a translu-
reference of the genus (Hexelasma) to the Balanidae cent, yellow, chitinous material. Thus, the construc-
or Chthamalidae discussed by Hoek (1913) and Pils- tion of the wall in these five species clearly indicated
bry (1916), remains unsettled. This species (H. ant- that either the existing definitions for the families
arcticurn) seems to have characteristics of both fami- Chthamalidae and/or Balanidae must be broadly
lies.. . . It appears that the genus is somewhere be- emended or that a new higher taxon was needed to
tween the two families." While Utinomi (1965) does accommodate this divergent group. In an attempt to
not agree with Bage's statement concerning the inter- further our understanding- of the situation, we studied
mediate nature of the genus, he says ". . . as far as the other lower or enigmatic Balanomorpha (e.g., Cato-
examination of the internal body of this cirriped is phragmus, Pachylasma, Chthamalus, Bathybalanus,
concerned, the mouth-parts agree well with those of Chelonibia, Metabalanus, Austrobalanus, Solidobala-
other members of Chthamalidae, while the cirri re- nus, Chirona, Elminius and Te~raclita),and concluded
semble, unexpectedly much more those of Balanidae that inclusion of the so-called hexelasmoids in either
than those of Chthamalus." He concludes that its the Chthamalidae or the Balanidae resulted in an un-
affinities are with the Chthamalidae, not the Balanidae. natural and confusing situation. The hexelasmoids,
It is important to recognize that Bage (1938) inter- presently included in the Chthamalidae, are in many
preted the labrum as having a distinct notch and the respects more similar to certain solid walled balanids.
third cirrus as resembling the second more than the We feel then, it is best to suggest the designation of a
fourth-just the reverse of uhat Utinomi reported he new taxon at the familial level, and propose the name
observed. The problem seems to be that the parts in Bathylasmatidae, in reference to the deep station the
question are really neither strictly chthamalid nor members occupy. The family can be distinguished
balanid; hence, the definitions for one or the other from the Chthamalidae, Balanidae and Tetraclitidae
family do not apply closely. as indicated in the following comparison:
In preliminary work on the antarctic materials we
Families of the BALANOMQRPIIA
encountered Henelasma and thus became inadvertently
Comparative Aspects
embroiled in this problem. Of initial importance to us
was simply resolving specific differences, and for this
purpose we went about acquiring specimens referrable 1) Wall of 8, 6 or 4 plates, or totally fuscd; always solid,
to the genus from other parts of the world. In the without cuticular filling or regular internal ribs.
search for additional characteristics that might be used 2) Opercular valves deeply articulated; tergum w i t h ~ u tpro-
nouncetl spur.
in discriminating between populations, we ground thin
3 ) Overlapping plates with or without radii.
sections the s u ~ ~ o s e "lid
dl~ Our 4) l(oslrumwith alac, or discernibly or indisccrrlihly fused
ment, in two previously described species, H. callisto- with rostrolatcral plates and thus overlapping ratlicr
derma (Pilsbry) and H. americanum Pilsbry, and in than underlapping adjacent plates.

r Bafhylosmo hirsufum (Hoek) 4 Hexelosmo spp,

+ Bofhylosma corol/iforme (Hoek) Aoplolasmo gen. nov., spp.
A tBathy/osmo oucklandicum (Hector) + t Jessorelosma pilsbryi Wit hers
Tefrachoelosmo southward gen, et sp. nov.
Chart 10. Distrihutio~nof Bathylasmatidac farn. nov.
5) Basis membranous or calcarrous; when calcareous, solid structure, tcriding to rrsrmble cirrus I1 morc than IV;
and not forming intricate interdigitations with basal posterior (inner) ramus either normal or antcnniform,
margin of wall, often fusing with wall so a s to ter- often in the same species.
minate diamctric growth. 7) Cirri without specialized setae.
6) Cirrus 111 resembling cirrus 1V more than 11; antcrior 8) without caudal
(outer) ramus normal or antennif(~rm,even in same g) penis without hasidorsal Doint.
species. 10) Mandiblc halanoid; inferior nrargin commonly pointed with
7) Cirri I and I1 commonly armed with bipinnate and few srnall spincs, rarely blunt, never molariform as in
other complex sctac, distributrd trrminally. most balaniils, nor pectinated or densely serrated as
8) With or without caudal appentlagrs. in chthamalids.
9) Penis without basidorsal point.
11 ) Labrum chthanraloid; thick (effectively bullate), without
10) Inferior margin of maudible finrly pectinate or coarsely
notch or incision.
serrate.
11) Labrum thick (bullatc) ; crest without notch or incision.
RATHYLASMATIDAE 1 ) Wall of 8, 6 or 4 plates, or totally concrescent; solid or
per~neatrdby longitudinal tubes, usually with internal
1) Wall of 6 or 4 platcs; solid arid without regular internal regular ribs (pl. XXXIV A, B; fig. 69) ; when tul~iferous,
ribs, or with chitin-filled longitutlinal tubes arrangcd -- -
i n a single row and more or less regular internal ribs ' Tlie
austral genus Elminius prescnts certain problems re-
a t base (pl. XLIII A, C ; fig. 6 9 ) . garding its position among the Balanitlae. I t is prcscntly as-
2) Opcrcrilar valves essentially balanoitl; although deeply signed to the subramily Ralaninac, but the characteristics in-
ar~iculatedin some species as in chtlramalids; spur of dicated in thc aforegoirrg kry, separating it from the fanlily in
tergrim as wcll devclolrcd as in most bnlanids. general, make the assignment c~uestiorrable.In fact, the notched
3 ) Overlapping plates without radii. labrnnl ant1 l~asitlorsal point on tlic penis found in some
4) Rostrurri c.omptrund, i~rtliscrrnibl~ fusctl with rostrolateral sprcics apprar to be the only truly halanoid characteristics.
Otlerwisc. E1n1ir1iu.s in many rcspccts stands closer to the
plates.
Tetraclitidac. The wall being of four plates does not in it-
5) Basis membranous or calcart.ous; whvn calcareous not self take on great importance,, hut the platrs being permcatcd
permeated by tubes nor forming coml)lcx interdigita- 1:y thick cuticular chitin (pl. XXXIV C. U ) and the nunlcr-
tions with basal margin of wall, hut often fusing with ous similarities betwccn tlrc oprrcular valves and appendages
wall so a s to terminate diametric growth. suggest a closer relationship to the Tctraclitidac than the
0 ) Cirrus 111, although in some rcspccts in~crmediatr in Balanidar. 1;or a s o l u t i o ~to~ this problcm see Ross, 19701, p. 9.

corrlaining living tissue, ~)artitioned off aricl empty, or 7) Cirri without spec,ialixrd srtat: (rxctapt ill E l l i ~ i r ~ i art-
~~~),
srcon~larilyfillet1 with ral(:arrorrs n~atcrial,or chitinous iclcs of ir~tcrrrrrdiatcpairs corr~monlyarmed with spines,
material iri Elminir~sspp. (111. XXXJLT (1, I) I . trc.tlr or hooks.
2) Oprrcular valvrs not d r r i ~ l y articulatrtl, trrgal spur 8 ) Without c:audal a ~ ~ p ~ n c l a g r s .
usually well devcloprd. 9 ) I'mis with basidorsal point (cxc:rpt in E l n ~ i n i u sspp.).
3 ) Overlapping plates usually with radii. 1 0 ) Ttrfcrior margin of rr1ari~li1)lrtcrrrrinating i n a point of a
f r w small spines or l)luntly rr~olariforn~(pectinate i n
4) Rostrum ronrpoarid; indiscernibly fused with rostrolatcral
platrs (except in Chelonibitr) or Yrcplaccd by direct El~ninirrs).
fusion of roslrolateral plates.
11) Labrum not bullatc:; crest with shallow to derp notch,
or incisiorr.
5) Rasis mcm1)ranous or calcareous; when calcareons, solid
or tubiferous and forming corr~plexinterdigitations with TETRACLITIDAE
margin of wall, usually not fusirrg wit11 wall so a s to 1) Wall of 4 plales, wllcn fusetl remainirrg separable along
tcrniirratc tlianrctric grclwth. suturcs, permeated by one or more rows of separate
6 ) Cirrus TIT somewhat irrterrr~rdiate i n structure between or conHuent longitudinal canals containing living tissue
cirri 11 and I V in some solitl-wallet1 forms, commonly or secondarily filled with calcareous and chitinous ma-
r ~ s e m h l i n g cirrus I1 rrrore than 1 V ; rarni never terial (pl. XXXIV E X , fig. 6 9 ) .
anterrniform (except i n Eln~iniris). 2) Opcrcular valves balanoid.
Aaptolasmo
Ch/homa/us Hexelasma
Pachylasma Chionelasmus Balhy/asmo
Pachylosma
Octomerfs
Cu/ophragmus
Chthamalidqe Bathylasrnatidae Tetraclitidae Balanidae
BALANOMORPHA
Fig. 69. Wall conslruction in the llalanonlorpha ( p l a n views) : C, carina; CL, carinolateral; L. lateral; RL, rtrs~rolateral;
R, rostrum. Arrrong the Chthan~aliciae, Catophmgmus has several rows of imbricate plates a n d Cl~ionc~lusn~us a single row
around the hasal margin. Thrse are indicated by dashed lines on the left side of their wall plans. T h e other genera having
thc same plan lack the imbricate plates (modified from Newman, 1967).

3 ) Overlapping platcs ~ v i hradii; radii reduced or lost in 4. Lcsser c:urvalure of intermctliale articles of postcrior
fuscd forms. cirri s ~ ~ y p o r l i4~ or
~ g morc pairs of sctac per scg~nent
4 ) Kosiruni coinpound. ............................... Bathylasma g m . nov.
5 ) Basis mcrnhranous or cal(::rrcous; when calcarcous solitl, 4 . Lesser curvature of intrmmdiale articlcs of posterior
not forming intcrtligitalions wit11 rnargin nor fusing cirri supl~orting 2 or 3 pairs of sclae pcr srgmrnt
with wall. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hexelasma 1loc:li
6 ) Cirrus I11 rcsernl-)ling I1 more than IV; inncr or outcr
ramus commonly antenniforrn. Bulanus (Me~aOnlnr~us)hoekirc~~z~s TPilsbry (=R.
7 ) Cirri TI atrd JTI conimonly armed with hipinnatc and (Chirona) everrnanrzi Pilshry) has not been included
other complex selae tlis~rihuicd along lesser curvature, in the Bathylasmatidae because the labrum and the
as in Elminius spp.
inferior angle of the mandible are wholly balanoid,
8 ) Wiitlout caudal appendages.
9) Penis wilhout basidorsal point.
as Pilsbry (1916) stated. Furthermore, the pmis bears
10) Inferior margin of mandible pectinate or serrale, never a basidorsal point. The labrum of the type specimen,
molariform. however, had not heen described and is therefore
11) Labrum not bullate; crest %it11 or without sliallow notch, figured here (fig. U3C).
never incised.
Bathyhalar~uspcntacrini Hoek, considered a chtha-
I n the Eathylasmatidae as it presently stands, we malid by Pilsbry (1916), would 1,e another possible
can recognize 3 groups of species and 2 monotypic candidate for inclusion in this new family because of
taxa that constitute distinct genera, as indicated below. its supposed similarities to Hexelasma (Pilshry, 1916) .
These can be separated in the following key and are However, the development of complex radii of the
further characterized in the generic diagnoses herein. balanoid type, the regular ribs on the inlerior of the
1 ) Bathylasma gen. nov. wall interlocltirig with comparable marginal teeth in
the basis, the rather well-defined although shallow
a ) B. corolliforrne (Hoek) : Type species
b ) B. hirsutum (Hoek) notch in the labrum, the bilobed second maxillae and
c ) R. aucklandicum (Hector) the basidorsal point on the penis are characters that
definitely exclude Balhyhalanus from both the Bathy-
2) l'essarelasma Withers
lasmatidae and the Chtliamalidae, and place it back
T . pilsbryi Withers: Type-species
in the Balanidae as a subgenus of Balanus.
3) Tetrachaelasma gen. riov.
Thus, very sharp lines are drawn between the Bathy-
T . soz~thwardigen. et sp. riov. : Type-species
lasmatidae and the Balanidae, as well as the Balanidae
4) H a e l a s m a Hoek and the Chthamalidae, and we are left in the position
a ) H. uelutinuna Hoek: Type-species of not having to vigorously justify familial rather than
b) H. arafurae Hoek subfamilial status for this new taxon. However, super-
c ) H. fosteri sp. nov. ficially the bathylasmatids might be suspected of stand-
d ) Hexelasma sp. Withers (1913, p. 847) ing much at the level of the existing subfamilies of
5) Aaptolasma gen. nov. Ihlanidae (Balaninae, Chelonibiinae, Coronulinae,
a ) A. callistoderma (Pilshry) : Type-species and Emersoniinae). This is not the case, for these fit
b ) A. americanum (Pilshry) the existing definition of the Balariidae while the bathy-
c ) A . triderma gen. et sp. nov. lasmatids do not. To include the Bathylasmatidae as
d ) A . brintoni gen. et sp. nov. a subfamily of Halanidae would entail an extensive
e) A . leptoderma gen. et sp. nov. emendation of the familial definition, an emendation
that in good part \bould require a revision of the larg-
est subfamily, the Balariinae. This highly complex and
1. Wall of 4 plates ..................................2 diversified taxon is in itself greatly in need of revision.
1. Wall of 6 plates ................................... 3
The bulk of the species fall hithin the gerius Balanus,
2. Rasis c a l ( ~ a r ~ o a sforming
, inf1vctt.d margin of wall;
scutunr with tobrrc.ulate callus above atltluctor pit. ... which, over the past 50 years, has itself been divided
............................... Tessarslasma Withers into numerous subgenera, the status of many of which
2. Basis membranous; scuturii witlront callus ............ is presently in a state of flux. Particular dificulties
........................... Tetraclmslasma gcn. nov. have arisen in certain of the solid u~alledforms (cf.
3. Gasis membranoiis; wall not pcrnrcattd with chitin-
Henry and McLaughlin, 1967), and it is superficially
filled tuhcs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
3. Basis calcareous; wall permeated with chitin-filled tubes near these and related forms that the bathylasmatids
................................Aaptolasma gen. nov, stand. It is beyorid the scope and means of the present

study to ur~tlertakcthe needcd revision of these insufi-

ciently kr~o\\rrl~alarlines,hut we are r~onetl~eless com-
1)ellcd to gi\#e th(: liathylasma~itlaeir~tlt.~c:ntlenl
rank.
Despite the marked dilfercncc.~alrearly enurnerated,
B e want to emphasize the apparrnt balanoitl rather
than chthamaloid afinities of the group. As a group
they stand distir~ctlyapart from the c:htharnali:ls ar~tl,
from our vantage point, it seems impossible to supl~ly
the facts supporting the long standing view lhat il is
from the chthamalids that the balar~oirlbarnac-les have
descended. It rather appears that thcir antec,eclents
rnay well be Iound among the sc:alpellitl Lepadomorpha
rather than among themselves; that is, the Balano- Fig. 70. I'la~r ~lotnrnt~laturc~:Scl~rmatic:left lateral
morpha could well be a polyl~hyletic group, as sug- t.r carinolatrrnl wall plate (1)trunded by hcavy solitl
gested in the accompanying diagram. and I)rtrl<en lines, viewctl from within), in rclation-
ship to adjilccnl plutcs 1, total hciglrt; 2, height hc-
Balanidae low shratti; 3, height of entirc shcath; 4, height of
/ shcath c:rossed by growth lines fornied through re-
Bathylasrnatidae
\ / elrninioids"
cession of operrular mcni1)runr; 5, width of basal
margin; 6 , width tlf ala; 7 inirrior alar margin; 8,
Chtharnalidae alar angle; 9, secontlary dldr increment or "wclting"
\ (see pl. XXIV C , XXV E lor exlrrior a q p r ~ t ;) 10,
superior alar margin; 11, apex (uqually truncatcrl
by erosion) ; 12, portion of wall overlapping adja-
cent plate (in this case without radii, a radius being
.............. LEPADOMORPHA the exterior counterpart of thr wclting (9) and
I..poll@clpoid Scolpellidqe
.............
............ j scrving much the same functirm).
Bathylasma gen. nov. 1. External growth lines relalively fine, those on alar rr-
Balanus, in part, Hoek, 1883, p. 155; Pilsbry, 1911, moved from and not closely paralleling infrrior
margin ........................................... 2
p. 7G; Hexelasma, in part, Hoek, 1913, p. 244; Pilsbry,
2. Upturned alar margin of latera wide and flat, f~ rrr~ing
1916, p. 329; Bage, 193G, p. 10; Kriiger, 1940, p. 30;
welting as much as t w ~ c ethe widlh of alar; platcs
Withers, 1953, p. 99; Utinomi, 1965, p. 13. morc than several centimeters hi&. relativelv thin
Diagnosis: Six solid wall plates; basis membranous. (usually 2 mm or less), opercular parts unIcnown
-
Spur of t e r ~ u mbroadlv rounded. essentiallv confluent ..........................B. aucklandicum (Hector)
with basal margin and continuous with articular mar- 2. IJpturncd ajar m a g i n of l a m a very narrow; forming
a braded welting; individuals more than 2 centimeters
gin; articular margin straight or concave, not sinusoid
ill hcight with platcs relatively thick (usually more
as in Tetrachaelasma. Inner ramus of cirrus 111 nor- than 2 mm) .................. .B. hirsutum (Hock)
ma1 or antenniform (in same species). Intermediate
articles of cirrus VI bearing 4 or more pairs of major Rathylasma corolliforme (Hoek, 1EG3)
setae. Plates XV-XXII, XLIII A: Text-figs. 71. 72
u
Distribution: North Atlantic and oceans bordering

Balanus corolliformis Hoek, 1883, p. 155, pl. VI,
Antarctica: ?Lower Oligocene, Miocene of New Zea-
figs. 21,22, pl. XIII, figs. 1-7.
land, Lower Miocene of Burma, Pleistocene to Recent
Hexelasma corollijorme: Hoek, 1913, p. 245; Pils-
in Antarctica.
bry, 1916, p. 330; Nilsson-Cantell, 1930b, p. 252.
Type-species: B&nus corolliformis Hoek, 1383,
Hexelasma coralliforme [sic] : Zevina, 1968, p. 94,
p. 155.
text-fig. 6 a-i.
Etymology: As for the family. non H. hirsutum: cf. Southward and Southward,
KEY T O SPECIESOF Bathylasma 1958, p. 635.
Hexelasma antarcticum Borradaile, 1916, p. 132;
1. External growth lines relatively coarse, those on the
alae running along and paralleling closely the inferior Withers 1924, p. 22; Bage, 1938, p. 1-E; Speden,
alar margin ................ B. corolliforme (IIoek) 1962, p. 746; Weisbord, 1965, p. 1015 in part;

Fig. 71. Brrthylasnrn corollijorme (Hock) : -4-D, right cirri ILIV, respectively; E, intermediate
articlc of cirrus VI; F and G, left and right mandibles, respectively; H and J, right and left
mandihlvs, respectively; I and K, inferior angles enlarged; L anti M, right and left maxillae I,
respectively; N, right rnaxilla I1 (A-C and M: Aurora, Sta. 111 l Australian Mus. No. P7206 1
and I-I-L and N: Umitake-Maru, Sta. 4 ) .
Utinomi, 1965, p. 13; Weisbold, 1907, p. 51 in part, son Land (66'4SfS, 71°24'E), 456 meters, 1 living
pls. I, 11, figs. 1-6; Zevina, 1968, fig. 1. specimen; Umitaka-Maru, Sta. 4, off Prince Harald
Diagnosis: Wall white; plates flaring u idely in young Coast (67O4S.53, 33 '42'E), 790 meters, 1 living
individuals, although less so in older (large) indi- specimen; Umitaka-Maru, Sta. 6 (07'51.5'S, 33'13.5'
viduals, so that aperture is generally as large or larger E ) , 630-680 meters, shells; SGya, Sta. 4 (68'29.7'S,
than the base; young specimens covered with persistent 32°02.7'E), 590 meters, shells; SGya, Sta. 6 (6Q010.0'
yellow epicuticle and numerous hair-like chitinous bris- S, 34'04.0'E), 620 meters, shells; SGya, Sta. 7 (68O
tles; caririolateral plates narrow, I!, to width of 09.7'S, 34'34'E), 830 meters, shells; Geol. Survey
basal margin of lateral plates. External alar growth (N.Z.) GS-7512, McMurdo Sourld (approx. 7g004'S,
lines closely paralleling inferior alar margin, not di- l67O5OfE), 100-200 ft above sea level, 7 plate frag-
verging abruptly near alar angle (111s. XVI U, XVIII ments; GS-7505 (approx. 78'28's. 165"5OrE), 3-10 ft
A ) . Scutum without apical boundary of adductor mus- above sea level, 29 plates and fragments; GS-7506
cle pit sharply delimited. Tergum with articular margin (approx. '7::"2:ifS, 165°50'E) , 0-10 ft above sea level,
straight. Both rami of cirrus 111 sorneu hat elongated ; 7 plates, 1 scutum; Endeavour, Sta. A-463, off Victoria
inner ramus slightly narrouer. sorrietimes very much Idand,Ross Sea (7Z020'S, 174'50fE), 465-468 meters,
elongate and antenniform. numerous living; Ellanin, Sta. 1003, off Antarctic
Material: Challenger, Sta. 150, off Kerguelen Islands Peninsula (O2"4lrS, 54"43'W), 210-23E meters, 2
(52"4'S, 71°22'E), 274 meters, I plate of type; Terra living; Sta. 10.54, off South Sandwich Islands (GO0
Nova, Terra Nova Bay, Victoria Land ( a p ~ r o x 75'S,
. 3 7 3 , 29'50'\V), ::23-1079 metels, 1 liter of shells;
164OE), 30 ft above sea level, 1 carinolateral (type of Sta. 1007. off South Georgia (Scotia Bank) (59'57'S,
H. ar~tarcticum); Aurora, Sta. 111, off Marie Hyrd 34041rW), 10Ot!-14G4 meters, numerous shells; Sta.
Larid (66'325, 141°39'W), 287 meters, several shells, 1088, off South Shetland Islands (60°49'S, 53'28'W),
2 living specimens; L)kcovery, Sta. 30, off Mac Robert- 586-595 meters, 3 carinae.

Depository: British Museum (Nat. Hist.), London, Supplementary Description: The form of Hathylasrna
Reg. No. 85:17, type (Challenger, Sta. 1 5 0 ) ; corolliforme is variable initially and may change con-
1917.2.9.59-68, type (Terra Nova Expedition) ; Aus- siderably with age. Plates of young individuals are
tralian Museum, Sydney, Nos. P7204, P7206, P7207, usually thin, and flare markedly outward, so that the
P7209, P7210, P7213 (Aurora, Sta. 111) ; Tokyo Uni- aperture is nearly as large as, or larger than, the base
versity of Fisheries, Tokyo (JARE) ; Geol. Survey (pls. XV, XX A ) . Thin plates of this type break ir-
(N.Z.), Lower Hutt, Nos. 7505, 7506, 7612; New Zea- regularly rather than along transverse cleavage planes
land Oceanographic Institute, Wellington (NZARP, (pl. XX B, C ) . However, in other young individuals
Ross Sea, Endeavour, Sta. A-463) ; USNM Cat. No. the plates are relatively thick, have little flare and are
125302 (Eltanin, Sta. 1003) ; 125303 (Sta. 1088) ; generally tall and columnar. When dried, such thick
125304 (Sta. 1054). shells tend to delaminate along obliquely transverse

14'6 ANTARCTIC CIRRIPEDIA
cleavage planes. The plates of large, older individuals to lA<the width of the laterals. As seen from the ex-
tend to be proportionately thin, but some become very terior, the alae are sharply delimited from the parietal
thick (pl. XIX A-C). More important is their having portion of the plate and usually exposed about % the
little or no flare. This is related to the initial growth height of the wall, below which point the parietal por-
pattern, a gradual change in growth pattern during tions of the plates come into contact, but do not inter-
development, and loss through damage and corrosion lock. Internally, the superior alar margin slopes
of the older flared portions of the shell around the obliquely towards the angle formed with the inferior
aperture that had given the flared aspects to individ- margin, and there turns directly downward, parallel
uals when younger. Such plates, when moderately to the line demarcating the lateral boundary of the
thick, delaminate along cleavage planes, beginning sheath, before turning to pass obliquely back toward
parallel to the growth lines and running obliquely the basal angle of the parietes, as the inferior alar
across the plate toward the apex; that is, the plate is margin.
composed of layers (pl. XXII A ) . External alar growth lines closely parallel the in-
Longitudinal thin sections of the plates reveal differ- ferior alar margin. This is because the arthrodial
ences in structure that relate to different growth char- membrane retreats with each growth increment so that
acteristics. In thin forms, lines formed by high organic each line is exposed as the next develops. The growth
content can be seen to loop between major growth lines take an abrupt turn as they approach the superior
ridges, while in thick shells they pass obliquely across alar margin, where a slightly raised welting is pro-
the plate from the major growth lines, apically, toward duced. This welting develops exactly in line with the
the inner side, and thus demarcate the laminations. It lower margin of the sheath and must relate to a com-
is along these lines that the laminations can be sepa- plex fold in the arthrodial membrane at this position.
rated. In thicker individuals cleavage goes from one However, it has no counterpart in the opposing articu-
side to the other, while in thinner shells of large in- lar surface of the adjacent plate.
dividuals, where the calcification pattern loops from The sheath is extremely simple, on the rostrum being
ridge to ridge and no transverse cleavage lines are bounded laterally by the two lines formed by the alar
formed, the plates break more or less irregularly. junctions, on the carinolaterals and laterals by the alar
In general, the transverse pattern is seen in large junction on one side and the superior alar margin on
individuals and the loop pattern in small individuals. the other, and on the carina by the superior alar mar-
However, when a long plate, thin near the apex and gins alone (pl. XVII E-H). Statements that the sheath
relatively thick near the base, is sectioned, the loop has been found lacking in small specimens (Hoek,
pattern is seer1 near the apex and the transverse pat- 1883; Nilsson-Cantell, 1930b) stem from a semantic
tern near the base. The two patterns intergrade com- problem related to imperfections in the original defini-
pletely between these two regions in a single individual. tion. Darwin (1854, p. 38) stated that the sheath in
From this it is clear that the pattern of growth changes the Balanomorpha ". . . sometimes blends insensibly
considerably during life (pls. XXI B, XXII A, K ) . into the lower parts of the compartments and then per-
haps it would not be thought a distinct element . . . ."
Young fragile individuals up to 25 mm or so in
He no doubt should have elaborated this point, for the
height are clothed wilh a persistent epicuticle and 'iperhaps" left cirripedologists with an incomplete defi-
dense spines or hairs along the growth ridges (111.

nition. Whether or not the sheath blends, or is dis-
XVI). The spines are not simply extensions of the epi-
tinct from the lower part, the function and develop-
cuticle, but have their bases extending into the cal-
ment of the area is the same, arid therefore all Ualano-
careous portion of the plate. They developed when
morpha (except Xenobnln~sus) have a sheath. It is
that level represrntetl the basal margin of the plate,
always precisely delimited, by the line or lines formed
and subsequeritly 1)ecame buried by the next growth
by the alae along the articular surfaces and/or by
increment. The epicwticle is for the most part lost in the superior alar margin or margins. Its lower limit
large, heavy-plated, old individuals. may be difficult to detect, but it is always marked, if
The basal margirls of the rostrum and lateral plates ever so faintly, by a line where the arthrodial or oper-
are essentially equal in width, while the carina is equal cular membrane connected the valves to the plates. It
to, or as little as half the width of the rostrum. l'he is the progression of this membrane down the interior
basal margins of the carinolaterals, although showing of the wall and/or ala that is responsible for the basic
no particular teridency towards reduction, are about 14 elements of the sheath. Thickening, overhanging, and

interior extensions toward the base are merely second- although not bilobed, have the cutting- edge . divided
ary elaborations. into two regions; and no marked changes appear to

The opercular valves also vary from thin to thick, occur with growth.
depending on the relative thickness of the wall plates. The cirri tend to increase in number of articles with
When thin, the interior relief is feebly developed, al- size (cf. cirral counts). The anterior ramus of the
though faint ridges or crests for the lateral depressor first pair is longer by about 3 segments; the articles
muscles are evident on the tergum and may be evident of both rami are wider than high and slightly protuber-
on the scutum (pl. XVII A-D). In heavy-shelled forms, ant along the anterior margins. The rami of the second
a distinct, broad pit for the scutoral adductor muscle cirri are subequal, the articles being wider than high
has developed (pl. XVIII D ) . In thin-shelled forms, and becoming more so with age. In larger individuals,
every other growth ridge along the occludent margin the anterior margin of the outer ramus becomes more
of the scutum is deleted, alternately with opposing protuberant and the distal quarter of both rami change
ridges on the adjacent valve, forming notches that from nearly normal (ctenopod: setae in pairs arranged
interlock when the valves are closed. In larger, thicker- like a comb along anterior margin) to antenniform
shelled specimens, while the rudiments of this system (articles articulated so as to be movable through 360°,
can be seen, the occludent margins have broadened setae arranged in whorls at each articulation). The
into flat areas that simply abut. The spur furrow of inner ramus of cirrus I11 is more slender and often
the tergum is depressed below the surface of the valve. longer than the outer ramus; the distal half or % of
The growth ridges traversing the furrow rise abruptly both rami being antenniform (figs. 71C, 72C).
and sharply along the articular margin, producing a The fourth, fifth, and sixth pairs of cirri are nor-
serrated edge in thin-shelled forms, but because of the mal, ctenopod, with the intermediate articles support-
close spacing of the ridges in thick-shelled forms, the ing 5 pairs of setae (fig. 71D, E ) . Penis normal,
margin can be relatively smooth. minutely annulated, without basidorsal point; no cau-
The labrum is thick and can be considered slightly dal appendages or vestiges thereof (fig. 72F). Bran-
bullate. The crest is nearly straight
- with a shallow chiae large and leaf-like.
central depression supporting numerous fine setae and
a few minute irregularly placed teeth or spines (fig.
72P, Q ) . The palps support relatively fine, soft setae Challenger 14
- 17
- 25 - 32 - xx - xx -
along their inner edge; stronger, stiff setae in a line (Hoek, 1883) 14 23 28 37 xx xx
across their outer face, backed by minute ctenae to the
outer margin. The mandible has 4 teeth, the second
Discovery (Nilsson- 9 13 5 24 3 30
Cantell, 1930) 12 15 20 26 30 31
through fourth often having accessory knobs but never
bifid; the inferior angle being relatively simple, sup- Aurora (Bage, 1938) 14
-
19
-
19
-
29+
-
22
-
22?
-
porting a few prominent, or an array of minute, spines. 14 17 25 29+ 22 xx
The form of the mandible changes appreciably with
- -
size (age). In small specimens the teeth are propor-

Endeavour(A-463) 10 22 ?+ 28 30
(small) 11 14 26 24 27 34
tionately large; the inferior angles narrow, projecting
and terminating in several sharp spines. With growth,
the blades become inflated in outline so that the teeth
appear proportionately smaller; inferior angles re- (large) 15 19 27 32f 37 36
- - - - - -
ceding and broadly rounded, supporting a number of
12 21 32 32 36+ 36
relatively small, irregular spines (fig. 72G-K) . The
first maxillae undergo a change comparable to that
seen in the mandibles. In small individuals, the cut-
ting edge below the notch supports about 5-8 large (large) 13 18 28 32 34 37
- - - - - -
spines for about 3/q its length, the lower quarter sup- 12 25 40 36 38+ 37
porting a comb of approximately 6 short, fine spines.
In older individuals, the large spines essentially dis-
appear, being replaced by several times as many finer
ones that differ little from the original fine spines
occupying the lower quarter. The second maxillae,

14s ANTARCTIC CIRRIPEDIA
Renrarks: I n 1883, Hoek described the present spe- in the discovery of early or pre-Pleistocene materials
cies as Balanus corolliformis, taken by the Challenger associated with other marine fossils, certain antarctic
from a depth of 274 meters off Kerguelen Island. The species of which are extinct. Yet Borradaile's material,
species mas not reported again until Nilsson-Cantell as with some, though not necessarily all, comparable
(1930b) described a specimen taken by the Discovery materials subsequently found in or on glaciers, may
from 200-220 meters off the South Shetland Islands. nonetheless be Recent, apparently having came to re-
I n this paper he decided (p. 252, footnote 1 ) that side well above sea level by one or more modes of
Hexelasma hirsutum from the North Atlantic was a uplift (Gow et al., 1965; Weisbord, 1967; Dayton
junior synonym of H. corolliforme, but the contention et al., 1969).
was subsequently investigated and disproved by the Until recently (Zevina, 1968), the occurrence of H.
Southwards (1958). eorolliforme (Hoek, 1883) in antarctic waters has
I n 1916 Borradaile described H. antarcticum, repre- been ignored, except for Nilsson-Cantell's (1930b) re-
sented by a number of disarticulated plates and valves, port, and then the species was erroneously considered
taken during the Terra Nova Expedition from a glacier synonymous with the north Atlantic species.
30 ft above sea level. He was puzzled by the circum- Of Nilsson-Cantell's specimen Utinomi (1965, p. 1 3 )
stance and rather fresh appearance of the specimens, says ". . . it may be an undescribed species or a young
and judged that it could not be decided whether they form of H. antarcticum . . . ." Weisbord (1965) also
were fossil or Recent. Borradaile considered the spe- stated that the Discovery specimen ". . . may perhaps
cies to be closely related to, but distinct from, H. auck- represent a young individual of H. antarcticum Borra-
landicum (Hector, 188S), known from the Miocene of daile." It is in fact a young H. corolliforme. The
New Zealand, and enumerated four diflerences by question that remains concerns H. antarcticum; is it a
which they could be separated. The second and third distinct species or a synonym of H. corolliforme?
differences listed are really for the same character, From published data alone, the question could prob-
namely the internal sculpture of the laterals and carino- ably not be resolved. The trophi and cirri appear simi-
laterals, which leaves us with three. One of these (the lar enough, if allowances are made for changes in size.
longitudinal striations on the rostrum) sezms to be It is the hard parts that seem to differ, the corolliforme-
due to substratum replication, which leaves us t n o type being relatively small and fragile; the antarcti-
that are significant: (1) in H. aucklandicum, the ex- cum-type large and robust, and the two never having
ternal growth lines on the parietal and alar surfaces been reported together. H o ~ e v e r Speden
, (1962) pub-
are finer and more delicate, and ( 2 ) those of the alar lished four text-figures (fig. 14 d-g) of plates from
surface do not parallel the inferior alar margin but the same fossil deposit: a rostrum, carina, carino-
diverge from it before bending abruptly downward lateral, and lateral plate, and from the figures, the first
to the outer alar angle. Rorradaile did not compare two look much like corolliforme, the other two like
his material with Hoek's H. corolliforme from near antarcticum. We have had the opportunity of exam-
Kerguelen Island, nor to H. I~irsutumfrom the north ing this material and this is indeed the case. There-
Atlantic, but if he had he would have found his ant- fore, both forms occur together, at least in fossil de-
arcticum closest to the antarctic form as then known, posits. A thorough examination of the specimens listed
and aucklandicum rather closer to the north Atlantic above. under Materials yielded convincing evidence
form, in regard to these differences. The synonymies that the t ~ forms
o are the same species; that is, what
of the taxa will be returried to shortly. has been called corolliforme is simply a young growth
The problem of whether or not H. antarcticum oc- form of antarcticum, as shown in the longitudinal thin
curred in the Recent was resolved by Rage (1938), sections cut from the same shell. The otherwise ap-
who described living material from 2G7 meters off parently different growth patterns, where the organic
George V Coast taker1 during the Australian Antarctic matrix is seen on one hand looping, and on the other
Expedition, and Utinomi (1')65), who described a liv- traversing the wall, blended from the corolliforme-type
ing specimen from 790 meters off Prince Harald Coast near the apex (the portion having housed the animal
taken during the Japanese Antarctic Research Expedi- when ~ o u n g ) and
, the antarcticum-type near the base
tion. However, neither Rage nor Utinomi compared (the portion laid down later in life).
their material to other known species of Hexelasma. A single, large carinolateral late was referred to
On the other hand, the fossil occurrence of H. antarcti- Hexelasma sp. (rather than to H. antarcticum or If.
cum in Antarctica was demonstrated by Speden ( 1962 ) , corolliforme) by Zevina 11968), because the sheath

occupied 1/2 rather than & : its length. In the present Balanus hirsutus Hoek, 1883, p. 158; Gruvel, 1920,
study we learned that the proportion of the plate occu- p. 55.
pied by the sheath depended on differential growth be- Hexelasma hirsutum: Hoek, 1913, p. 245; Pilsbry
tween the basal and lateral margins of a plate; that is, 1916, p. 330; Southward and Southward, 1958, p. 635;
differences in this proportion reflect environmental Utinomi, 1965, p. 11.
conditions and are not good taxonomic characters. non Hexelasma corolliforme: Nilsson-Cantell, 1930b,
Therefore Zevina's specimen has been referred to p. 252, footnote 1.
Bathylasma corolliforme. Diagnosis: Wall white, conical, low in small and
Marked differences between the cirri and certain of high in large individuals, aperture never flaring; un-
the mouth parts, seen when the extremes in size are eroded surfaces covered with persistent, velvety epicu-
compared, fail completely when intermediate-sized in- ticle; carinolateral plates very narrow, basal margins
dividuals are examined. Thus, all forms previously approximately to ?h width of basal margins of
referred to Hexelasma antarcticum, except that from lateral plates. External alar growth lines removed from
"Sars Bank" (Weisbord, 1965, 1967; Eltanin, Sta. 255 and not closely paralleling inferior alar margin; w-elt-
-see Tetrachaelasma herein), and to H. corolliforme, ing along alar margin very narrow. Scutum with api-
except that from Japan (Kriiger, 1911-see Aapto- cal boundary of adductor muscle pit usually delimited
lasma callistoderma herein), represent the same spe- by a sharp rounded line; tergum with articular mar-
cies. This species ranges from at least the early Pleisto- gin broadly concave. Anterior ramus of cirrus I11
cene to the Recent, in Antarctica. often much elongate, antenniform.
In the discussion of the family Bathylasmatidae it Distribution: Northeast Atlantic: Faeroe Islands
was mentioned that the genus Hexelasma, as previously south to Azores, 944-1829 meters (Southward and
known, contained 3 groups of species and 2 monotypic Southward, 1958). Recent.
taxa which we consider constitute distinct genera:
Material: Sarsia, Sta. 9/1, off continental shelf, south
Bathylasma, Hexelasma, Tetrachaelasma, Tessarelasma,
of Ireland (48'33'N, 10°01'W), 1170-1244 meters,
and Aaptolasma. Species previously included in Hexe-
numerous shells; 9/2 (4S034'N, 10°0O'W), 1042-
lasma, and discussed above as such (corolliforme, hir-
1225 meters, numerous living; P1/62 (4S038.5'N, 9"
sutum, aucklandicum and antarcticum), were removed
42.5'W), 494-567 meters, 1very large.
from Hexelasma and the new genus Bathylasma erected
for them, Bathylasma corolliforme (Hoek, 1883) here Depository: Marine Biological Association, Plymouth.
being designated the type. Bathylasma stands closer to Remarks: Bathylasma hirsutum from the North At-
Tetrachaelasma than to Hexelasma S.S. which, in turn, lantic is not closely related to B. corolliforme from
stands relatively close to Aaptolasma, and with Aapto- the Antarctic. However, it has been confused with this
lasma, Tessarelasma may also hold its closest affinities. species in the past. Nilsson-Cantell (1930b) studied
The similarities between Bathylasma and Tetrachae- a small specimen taken by the Discovery from Brans-
lasma included size, general characteristics of the field Strait, South Shetland Islands, which he at first
plates and valves, and the setation of the cirri. Bathy- identified as B. corolliforme. Subsequently, he had the
lasma hirsutum, B. aucklandicum and Tetrachaelasma opportunity to make a comparison with B. hirsutum
together stand somewhat apart from B. corolliforme and decided, as indicated in a footnote in the same
by virtue of the alar growth lines being hooked near paper, that the two species were synonymous. This
the alar angle, and in this regard they are similar to assertion was thoroughly investigated by the South-
Aaptolasma and Hexelasma. Thus while the genera are wards (1958), and their study demonstrated the two
markedly distinct in some respects, they share a num- to be quite distinct.
ber of characters, and this of course leads to their Bathylasma hirsutum can be separated from B. corol-
inclusion in the same family. Of all known forms of liforme by several characteristics of the hard parts. In
the family, Bathylasma would appear to be the least particular, the articular margin of the tergum, rather
specialized. than being straight, forms a broad curve (pl. XXIII B,
D) ; the growth lines, rather than closely paralleling
Bathylasma hirsutum (Hoek, 1883) the inferior alar margin, stand above it and form a
Plates XXIII, XXIV; Text-fig. 73 divergent hook at the alar angle (pl. XXIV C, D) ; and
Balanus sp. Jeffreys, 1878, p. 414 (fide Southward the carinolateral plates, rather than being $4 to g$
and Southward, 1958). the width of the laterals, are relatively narrow, being

Fig. 73. Bathylasn~nhirsutunz (Hock) : A, labrum with left palp in outline, right palp tirleted;
B, crest of labrr~rri enlarged; C, lelt palp; D and E, right and left mandihles, rcsprctively;
F and G, riglit and left maxillae I, respectively; H, right maxilla 11; ILL, cirri I-IV, rcsprctivcl):
M, intc.rn~ctliatescgmrnts of cirrus VT; N, penis. Sarsin, Sta. P9/2.
approximately to !: the witlth of the laterals A comparison with B n ~ l ~ y l a s mnucklnndicum

n from
(pl. XXIII E l . The trophi and cirri differ in small the Mioce~ieof New Zealarid mill he taker1 up in the
ways. The crest of the labrrrm supports many more section on that species, to follow.
small teeth (fig. 731:) ; the inferior angle of the man- Although B. hirsutum and U. corolliforme are mor-
dible is less produced in small specimens (cf. figs. phologically comparable, B. hirsutum may be consid-
72G, H; 73D, E) ; and the second cirrus is apparently ered more advanced in having the caririolateral plates
always normal (fig. 7 3 5 ) . The inner ramus of the third much reduced in width, the parietal portion forming
cirrus, although usually antenniform, more slender and a thin slip between the carina and laterals. The same
much longer than the outer, may be normal in some trend is seen in Aaptolasma among the bathylasmatids,
individuals. and in certain species of Balnnus and A c a s ~ aamong

the balanids where it is inferred to have led to 4-plated margin before bending abruptly toward the alar angle,
forms. An alternaiive plan in the evolution of 4-plated are all characteristics that point to this relationship.
foims is seen in Pachylasma among the chthamalids, The question then is whether or not B. aucklandicum
where the carinolaterals fuse with the laterals (Darwin, is a distinct species or a junior synonym of B. hirsu-
1854). tum. Since the differences that had separated B. ant-
A new 4-plated bathylasmatid genus reported on arcticum and B. corolliforme have evaporated, and
here, Tetrachaelasma, is otherwise closely related to there seems to be no published information that can
the 6-plated B. hirsutum. In that the carinolaterals are be used to separate aucklandicum and hirsutum other
much reduced in B. hirsutum, it seems most probably than age and relative thickness of the wall plates, one
that 4-platedness in Tetrachaelasma was achieved is inclined to consider forming the synonymy. Having
through a comparable reduction process in a hirsutoid just been confronted with the great range of thickness
ancestor. seen in B. corolliforme, it would be difficult to place
much weight on thickness alone as indicating a signifi-
Bathylasma aucklandicum (Hector, 1888) cant genetic difference when environmental factors
Plate XXV could have played an important role; and the Miocene
Scalpellum aucklandicum Hector, 1888, p. 440. age in itself is not a good criterion when, although
Pollicipes aucklandicum, in part: Benham, 1903, many species have become extinct since, many extant
p. 111; Clarke, 1905, p. 419; Park, 1910, p. 113. species appeared at that time. But fortunately we do
Hexelasma aucklandicum: Withers, 1924, p. 18. not have to test the case by these criteria alone.
Material: Geol. Survey (N.Z.) , 695, locality N38/ In examining the upper portion of very large lateral
524, 1; Miocene, Waitemata Beds, Motutapu Island, and carinolateral plates of B. aucklandicum still pretty
Auckland (approx. 36'45'S, 174'55'E), 1 carino- much imbedded in a sandy matrix, the growth lines
lateral or lateral plate. on the exterior alar surface were observed to rise from
the inferior toward the superior alar margin, and then
Depository: Geol. Survey (N.Z.) .
to curve or hook downward toward the alar angle
Diagnosis: Known only from numerous large (as where they appeared to terminate in sequence along the
much as 18 cm in height) remarkably thin plates (usu- superior margin as a very minute welting. This agrees
ally 2 mm or less in thickness) forming a tall, cylin- with the description and figures of Withers (1913;
drical wall that was probably wider at the aperture 1924), who said the alae were simple, with no distinct
than at the base, but not flaring. The external alar upturned margin at the edge. Although the minute
growth lines are rather delicate and diverge from the welting we detected could be considered an upturned
alar margin before turning abruptly downward to- margin, it is proportionately so small compared to
wards the alar angle (pl. XXV B, C ) . Withers (1913, the overall extent of the plate that the alar margin
1924) states that the alae are otherwise simple, Ye., would be considered effectively simple. However, in
there is no distinct upturned extension at the margin."
clearing away the bounding matrix beyond this point,
However, this portion of the diagnosis must be
it was discovered that this was not the actual superior
emended, for the thin, delicate margin is upturned and
alar margin of the late-the growth lines continued
forms an unusually flat, broad welting l h the width in an upturned direction for a distance equal to nearly
of the entire ala (pl. XXV B ) . This is an important
half the width of the ala, before the actual margin was
characteristic separating this species from B. hirsutum.
The opercular valves are unknown-the tergum reached. This feature is remarkable only in its breadth,
figured but questioned by Withers (1913, pl. 85, for all other species of the Bathylasmatidae have vari-
13a-b) appears to be a plate or valve of a verruco- ously developed weltings along superior alar margins.
morphan. The significance here is that in B. hirsutum, a species
Remarks: While B. aucklandicum has long been com- otherwise very close to B. aucklandicum, only a very
pared with B. antarcticum, or vice versa (Borradaile, narrow welting is developed. There is no reason to
1916; Withers, 19241, the species is in fact very much believe that such a great difference is induced by vari-
more similar to B. hirsutum from the North Atlantic. ations in the environment, but is rather under strict
The general tendency for the plates not to flare at the genetic control, and therefore of great diagnostic value.
aperture, the relatively narrow carinolateral plate, and The finding of this structure in B. aucklandicum, in
the alar growth lines diverging from the inferior alar general so very thin and fragile, was fortuitous. The

welted alar margin is thin and fragile in most species, ward by as much as 90°, so as to align with the articu-
and therefore easily eroded or broken off, so that in lar surfaces of the slightly bowed rostrum. Alar
free plates or fossil materials it is frequently lost. growth lines diverging upward from the inferior alar
margin before turning downward toward the alar
Tetrachaelasma gen. nov. angle, and then turning abruptly upward again, just
Diagnosis: Four solid plates; basis membranous. before reaching the superior alar margin to form a
Tergum similar to that of Bathylasma but articular coarse but narrow welting (pl. XXVIII D). The
margin more or less sinusoidal, rather than straight or carina, the highest but narrowest of the plates, sup-
concave; lateral depressor muscle crests few and reports large alae that internally contribute to nearly
stricted to a small area just below carinal angle. Rami half the total circumference of the sheath. Externally
of both cirrus I1 and I11 antenniform; intermediate the alar growth lines compare closely to those of the
articles of cirrus VI bearing 3 or 4 pairs of major lateral plates.
setae. The scuta occupy entire aperture when closed, the
Distribution: Southeast Pacific-Drake Passage area; terga being hidden by the tall, narrow, wrap-around,
Recent; 1100 to 2300 meters. apical portion of the carina (pl. XXVI C, D). The
Type-species: Tetrachaelasma southwardi gen. et sp. scuta are heavy and triangular in shape, the deep pit
nov. for the adductor muscle being bounded above and
Etymology: With reference to wall of 4 plates. laterally by a distinct line, but opening imperceptibly
below, on to the general interior surface; depressor
Tetrachaelasma southwardi sp. nov. muscle pit deep, V-shaped or cup-shaped, sometimes
Plates XXVI-XXXI, Text-fig. 74 marked by faint crests; occludent margin with every
Hexelasma antarcticum Borradaile, in part, 1916, other growth line deleted, so as to interlock with the
p. 132: Weisbord, 1965, p. 1015 ("Sars Bank") ; 1967, reverse situation on opposing valve; with large articu-
p. 51, pl. 11, figs. 7-8 ("Sars Bank," Eltanin, Sta. lar ridge extending more than half way down the valve
225). (pl. XXVI F, H ) . Terga relatively thin and fragile,
more or less triangular but with articular margin
Material: Eltanin-SOSC Sta. 6, South Pacific (52"10r
thrown into sinusoidal curve; depressor muscle crests
S, 142" 10rW), 2304-2328 meters, 7 specimens; Sta.
few, weakly developed, restricted to the area along the
376, off Falkland Islands (54'03'S, 5S003'W), 1720-
basal margin just below the carinal angle; spur
1739 meters, plates only; Sta. 216, off southern Chile
broadly rounded to slightly pointed, margins virtually
(52"53'S, 75"36'W), 1190-1263 meters, plates only;
continuous with basal and articular margin, external
Sta. 255, off Cape Horn (59'455, 68"50rW), 1207-
furrow very shallow, essentially at the general surface
1591 meters, plates only.
in larger individuals (XXVI E, G ) . Unlike the exter-
Depository: USNM Cat. No. 125305 (holotype; nal surface of the scutum, tergum worn smooth or
SOSC Sta. 6) ; 125306 (paratypes; SOSC Sta. 6 ) ; nearly so, especially near the apex, by having rubbed
125307 (Sta. 376) ; 125308 (Sta. 216) ; 125309 (Sta. on the interior of the carina. Basis membranous.
255). Crest of labrum slightly concave, smooth; interior
Diagnosis: As for genus. surface covered with rows of ridges, spinules and
Description: Solid, thick. white wall plates 4 in num- setules (fig. 74G, H ) . Palps broadly rounded, with
ber: rostrum, right lateral, left lateral and carina numerous setae on outer and posterior margins, and
(pls. XXVI D, XXVIII A ) . Form low and conical in inner surface. Mandible with 4 teeth, teeth simple,
young, to high conical in old (large) individuals. Sur- without accessory knobs; lower margin of fourth tooth
face covered with yellow epicuticle and numerous fine with a few low spine-like elevations; inferior angle
hairs or bristles along growth lines, persistent in young with about 14 irregular spines (fig. 741-L). First
but generally deteriorated and lacking in large indi- maxillae without notch; but lower % of cutting edge
viduals. Rostrum widest plate, only slightly bowed produced beyond upper 1/3, supporting approximately
transversely; internally the sheath occupies about ?h 16 strong spines distributed along length and a group
the height of the plate but is unusually narrow, being of 6 or so short spines, a few pectinated, just above
flanked by very broad articular areas receiving alae of the inferior angle (fig. 74M, N ) . Second maxillae not
lateral plates (pl. XXVIII B) . Laterals nearly as wide bilobed, but setae divided into 2 groups by a median
as rostrum, remarkable in having the alae bent in- nearly naked area (fig. 7 4 0 ) .

SYSTEMATIC ACCOUNT
Fig. 74. Trtrucharlasma southwardi gen. e t sp. nov.: A-D, cirri I-IV, respectively; E, intcrmediate article
of cirrus VI; F, penis and pedicel of cirrus VI; G. labrum, palps delctcd; H, crrst of labrum enlarged; I,
right mandible; J, inrerior angle of right mandible enlarged; K, left mandible; L, inferior angle of left
mandible enl<+rged;M and N, right and left maxillae I, respectively; 0, right maxilla 11. Ellanin-SOSC Sta.
6 (holotype, USNM Cat. No. 125305).
Counts for 2 sets of cirri are given below: Cirrus I with setae arranged in whorls on exterior
I 11 I11 IV V VI surfaces of all but basal segments; interior surfaces
Specimen 1 14 31 37 37 33 32 heavily setose; anterior ramus wider than posterior
- - - - - -
ramus and more heavily armed with setae. Cirrus I1
15 26 52 29 24 35
with both rami antenniform; anterior ramus somewhat
wider, with intermediate articles samewhat lobate along
lesser curvature. Cirrus I11 with both rami antenni-
Specimen 2 16 22 26 28 32 form; anterior ramus, although wider than those of
13 23 62 23 29 32 succeeding and proceeding pairs, tending to resemble
more the setation of cirrus IV than I1 throughout
most of its length; posterior ramus about as wide as

rami of cirrus IV, with all but basal articles tending to the laterals, as described for Pachylasma by Darwin
produce the antenniform effect. Cirri IV through VI (1854). Bathylasma hirsutum is also similar to Tetra-
normal, intermediate articles supporting 3 or 4 pairs chaelasma in the narrow trough-like carina and rela-
of setae along lesser curvature (fig. 74D, E ) . Penis tively thick wall. These, taken with the similarity of
more than twice the length of the pedicel of cirrus appendages, especially the markedly antenniform third
VI, finely annulated, without basidorsal point (fig. cirrus, and the deep bathymetry of both, suggest a very
74F). close phylogenetic relationship.
Etymology: Named for Dr. Alan Southward of the Four-platedness in the Balanomorpha is attained
Plymouth Laboratory, in appreciation of much help through reduction from 6-plated forms (Darwin,
in checking types, providing specimens of Bathylasma 1854). It is apparently an adaptation, in good part,
hirsutum, and many helpful suggestions. to intertidal conditions or to other extraordinary en-
vironmental conditions such as estuaries, living em-
Remarks: Tetrachaelasma southwardi is the deepest
bedded in corals or sponges, inhabiting the deep-sea
known balanomorphan, exceeding the depth attained
or other geographical regions mhere species diversity
by its near relative, Bathylasma hirsutum, by some
and/or competition between barnacle species is rela-
475 meters. It is remarkable in having an extremely
tively low (Newman, 1967). In other words, while
heavy shell. Great thickness is likely an adaptation
4-platedness seems to be well suited to the physical
affording protection, but some workers would no doubt
environment, it does not seem competitive where com-
question this assumption. It is given support, however,
parable 6-plated forms prevail. Eight-plated balano-
by the fact that the scuta too are remarkably heavy
morphans have comparable distributions but are even
and they alone occlude the aperture. The terga on the
more restricted, and therefore also appear to be ex-
other hand are relatively light, actually a little fragile,
cluded from the general littoral by hexamerous types.
and unlike Bathylasma corolliforme, they fit closely
Tetrachaelusma, the deepest known balanomorphan,
into the trough formed by the broad alae, and apical
appears to be another example of this type of competi-
portion of the high carina, and thereby are unexposed
tive exclusion.
when the animal is closed. So protected, thickness in
Only seven complete specimens of Tetrachaelasma
the terga would be superfluous, and this must be the
southwardi have been taken by the Eltanin and then
explanation for their fragile nature which is otherwise
from one station. If it had not been for them, the
so conspicuously out of kezping with the great weight
loose plates taken by the Eltanin at three other sta-
of the rest of the shell. Some loose plates on hand
have been deeply but incompletely drilled by boring tions would have been extremely difficult, if not im-
organisms, presumably gastropods. possible, to interpret or identify (pl. XXIX). It
probably would not have been possible to tell whether
Despite the great weight of the wall plates and the
the loose plates represented a 6- or $-plated species-
broad overlapping surfaces between them, as in Bathy-
that is, whether the carinolateral plates were simply
lasma and Hexelasma they are only weakly held in
not taken in the sample or never developed in the ani-
place, there being no interlocking sutures or calcareous
mal. The great thickness of the plates suggested they
basis to reinforce the wall.
were either Bathylasma or Pachylasma and, without
Tetrachaelasma is further remarkable in having but
opercular parts or appendages to reveal their true
4 plates. Its closest relatives among the Bathylasmat-
nature, it probably would have been assigned question-
idae, in the genus Bathylasma, have 6. Of these, Bathy-
ably to one or the other of these genera.
lasma hirsutum is notable in having the parietal por-
tion of the carinolateral plates very much reduced, in The loose plates from "Sars Bank" (Eltanin, Sta.
some cases to mere slips mhich internally contribute 255), previously identified as Hexelasma antarcticum
little to the sheath, so it would be relatively simple for (Weisbord, 1965, 1967), can now likewise be recog-
the carinolaterals to fail to develop at all in B. hirsu- nized as plates of Tetrachaelasma (pl. XXX). They
t u m , resulting in a 4-plated form virtually identical to are remarkably large-twice as large as the largest
Tetrachaelasma. It is important that this be under- of the complete specimens taken elsewhere. Maximum
stood, both in understanding the relationships between size of the species then is as great as that attained by
these 2 taxa, and in understanding that 4-platedness Bathylasma hirsutum, but considerably less than that
has no doubt developed in this way in Tetrachaelasma, attained by B. corolliforme and B. aucklandicum. Tet-
rather than through fusion of the carinolaterals with rachaelasma, along with related species in the genus

SI-STEMATIC: ACCOUNT 155
Bathylasma, forms a complex of the largest species in Diagnosis: Six solid plates; basis membranous. Spur
the Eathylasmatidae. Furthermore, they inhabit much of tergum narrowing to a blunt rounded point and,
greater depths than any other Balanomorpha. although distinct from basal margin, effectively con-
tinuous with articular margin. Posterior ramus of
Genus Tessareltzsma Withers, 1036 cirrus I11 often antenniform; intermediate articles of
Tessarelasrna Withe~s,1936, p. 591. cirrus V1 bearing 2 or 3 pairs of major setae.
l j i u g r ~ o ~ i s :Four solid plates; basis calc.areous, mar- Type-speri~s: Hexelasma uelutinurn Hoek, 1913,
ginally thickened so as to appear as inflection of the p. 246; by subsequent designation, Withers 1913,
wall. Scutum balanoid, remarkable in having area p. 847; Utinomi 1965, p. 13, emend.; spccimens so
above shallow adductor muscle pit roughened as in designated from Srboga, Sta. 105 only-type-locality
Hesperzbalanus. Soft parts arid tergum unknown. (cf. Aaptolasma leptoderma for explanation) ; Indo-
Lower Miocene. Burma. West Pacific; Recent.
Tessarelasma pilsbryi Withers, 1936

1. Scutum with adductor ridge, and crests for lateral de-
Diagnosis: As for the genus. Type hy monotypy. pressor musc:les . . . . . . . . . . . . . . . . . . . H. arafurae Hoek
Remarks: Withers (1936) referred Tessarelasma to 1. Scutum simple, without adductor ridge or crests.. ... 2
the Chthamalidae because of its similarities to both 2. Ir~tcrniediatearticles of posterior cirri widcr than long;
lesser curvaturc supl~orting2 pairs of setac per seg-
Hexelasma s.1. and Pachylasma. He pointed out that
ment ........................... H . velutirrum llock
the solid wall, lack of radii, compound rostrum, bala- 2. Intermediate articles of posterior cirri longer than wide;
noid scutum and marginally thickened calcareous basis lesscr curvature supporting 3 pairs of setae pcr stag-
are morphological features held in common. There is ment ........................... .H. fosteri sp. nov.
no certainty to which of these two groups, if either,
Hexelasma fosteri sp. nov.
Tessarelasma belongs. In species of Pachylasma there
Text-figs. 75, 76
is a nearly complete trarisition from 8 plates to 4 plates,
through 6-plated forms (Darwin, 1854). Although Diagnosis: Shell white, no persistent. gray, velvety
there are no 8-plated hathylasmaiids, in this family the epicuticle; surface marked by horizontal rows of small
reduction from 6 plates to 4 plates also occurs. Both beads and small bristles paralleling growth lines.
chthamalids and bathylasmatids develop a marginally Crest of labrum supporting row of blunt, round teeth
thickened basis, so that this feature is of little taxo- overlain by groups of fine setules; anterior ramus of
nomic value except that it is not a characteristic of cirrus I11 twice length of posterior, and antenniform.
balanids. While the terga are unknown in Tessare- Intermediate articles of posterior cirri 11L2-2 times as
lasma, the roughening of the scutum above the adduc- long as wide, supporting 2 major and 1 minor pair
tor muscle pit is reminiscent of the solid-walled hes- of setae.
peribalanids, and the plate is othernise more balariid Material: Tui Expedition, Sta. 011-14, north of Auck-
than chthamalid in appearance. The scutum thereby land, New Zealand (30°45'S, 173 51fE), 538-676
indicates balanid rather than chthamalid affinities. The meters, 1 specimen; "Tui Oceanographic Cruise" of
structure of the basis being chthamaloid or hathylas- 1962; with Megalasma striatum Hoek and Trilasmis
matoid on one hand, with the scutum appearing kaempferi (Darwin), the former on spine of echinoid,
balanoid on the other, suggests an intermediate posi- substratum of others unknown (Dr. R. K. Dell, Do-
tion between the chthamalids and balanids. For these minion Museum, Wellington, N. Z.; B. A. Foster,
reasons we tentatively suggest bathylasmatid affinities pers. comm.)
for Tessarelasma. Transverse thin sections of the wall Depository: USNM Cat. No. 125310 (holotype) .
might be revealing, but material was not available for Description: Species known from single specimen
the present study. approximately 5 mm in rostrocarinal diameter and
4 mIn high. Wall essentially identical to that de-
Genus Hexelasma Hoek, 1913 scribed by Hoek (1913, p. 246, 251) for Hexelasma
Hexelasma, in part, Hoek, 1913, p. 24.4; Pilsbry, velutinum and H. arafurae. It differs from the former
1916, p. 329; Kriiger, 1940, p. 30; Withers, 1953, in lacking a persistent velvety cuticle, and from the
p. 99; Utinomi, 1965, p. 13; 1968, p. 30; Zevina latter by presence of horizontal rows of small beads
1968, p. 94. running parallel to the base; a row usually on the

Fig. 75. Hexelasma fosteri sp. nov.: A, entire specimen from above and B, from right side; C and D,
right exterior and left interior scuta, respectively; E and F, right exterior and left interior terga, re-
spectively. Tui Sta. 011 (holotype, USNM Cat. No. 125310).
space between growth lines and another along the ele- ficant that the counts for cirrus II are equal for both
vated growth line itself (fig. 75A, B) . Opercular valves rami and lower than for cirrus I. The posterior ramus
differ from those of H. arafurae, mainly in the scutum of cirrus I11 is elongate and antenniform, comparable
lacking adductor ridge and crests for lateral depressor to cirrus I11 in H. arafurae and contrary to the sit-
muscles (fig. 75C, F). They do not differ at all from uation in several specimens of H. velutinum, described
one of the forms of H. velutinum described by Hoek by Hoek (1913). However, the occurrence of a nor-
(1913, pl. XXX, fig. 5a-d) . mal and an antenniform ramus on cirrus I11 in the
Lahrum thick, with central portion anteriorly con- same species, in both the bathylasmatids and chtha-
vex and minutely squamate; crest only slightly con- malids is known, so that no value can presently be
cave, supporting about 33 rounded teeth, none of placed on this character in the present genus. Articles
which are bifid (fig. 766, H ) . The rounded condition of the posterior cirri are generally 1%-2 times as
cannot be due to wear as each tooth in the central long as wide, the inner curvature of each supporting
region is fronted by a group of setules. Palps un- 2 major and 1 minor pair of setae (fig. 76D). Penis
usually narrow, bearing a conspicuous row of about remarkably short, being about as long as the basal
8 strong setae projecting from the anterior (outer) segment of the pedicel of cirrus VI (fig. 76F). If it
surface. Mandibles resemble those of other species were not that other species of the genus are so de-
in the genus, both in form and in differing somewhat scribed in this regard, the reduced condition might
from one another in the same specimen. The present be considered simply indicative of immaturity.
species has 4 teeth on one side arid 3 on the other, Etymology: Named for Brian A. Foster, University
ar,d differs from previously described species in hav- of Wellington, New Zealand, who kindly sent us the
ing relatively few strong spines jutting forward from specimen of this species.
beneath the inferior angle (fig. 76J, K ) . Nothing re- Remarks: The two previously known species of
markable was noted regarding the first and second Hexelasma, H. velutinum and H. arafurae, were taken
maxillae. by the Siboga Expedition in the Indo-West Pacific
Cirral counts are as follows: and described by Hoek (1913). Subsequent records
(Broch, 1931, p. 53; Hiro, 1939, p. 70) extended the
range of the former from the Banda Sea north to
southern Japan.
The present form, while sharing characteristics with
both previously known species, is, by the structure of
opercular valves, distinct from H. arafurae, but very
In general the counts are low as compared with the similar to H. uelutinum. After a preliminary exam-
other species. The specimen, being small, probably ination we were inclined to attribute cer~ain small
is relatively young and the counts would be expected differences found in the form to geographical varia-
to increase with age. However, it is probably signi- tion, and to consider it simply as representing a south-

ern range extension of H. velutinum, from the Banda ment of spinules on the crest of the labrum, the cirral
Sea south to off the Norfolk Ridge, New Zealand. count for cirrus I1 being lower than for cirrus I, and
However, while the hard parts are very similar, a the articles of the posterior cirri being longer than
critical examination of the feeding structures revealed wide and bearing 3 pairs of setae instead of 2, are
there were greater differences between H. velutinum of particular importance in separating the new species
and the New Zealand form than there were described from H. velutinum.
between H. velutinum and H. arafurae. These, based After this section was written, a specimen identified
on inferences drawn from better known balanomorph- by Broch (1931) as Hexelasnra velutinurn Hoek was
ans, suggest we are dealing with a distinct species. received. It turned out to represent a new species,
Of the differences mentioned in the description, the Aaptolasma leptoderma gen. et sp. nov., described
form and setation of the palps, the pectinate arrange- later in this paper. This led to a more critical
Fig. 76. Hexelasma fosteri sp. nov.: A-D, cirri I-IV, respectively; E, intermediate articles of cirrus VI;
F, pedicel of cirrus VI and penis; G, complete left palp in outline; H, crest of labrum enlarged; I,
labrum, palps removed; J, K, right and left mandibles, respectively; L, left maxilla I ; M, right maxilla
11. Tui Expedition, Sta. 011 (holotype, USNM Cat. No. 125310).

look at H. velutinunz and the conclusions are discussed 2. Carinolateral plates relatively narrow, basal margins
under the new species of Aaptolasma. The outcome % or less width of basal margins of lateral plates.. . 3
2. Carinolateral plates relatively wide; basal margins about
is that H. fosteri sp. nov. now stands as a very dis-
W or more width of basal margins of lateral plates
tinct species. ............................... . A . brintoni sp. nov.
3. Opercular valves white.. . . . . A . callistodrrma (Pilsbry)
Aaptolasma gen. nov. 3. Opercular valves orange ......A. americanum (Pilsbry)
Balanus, in part, Pilsbry, 1911, p. 78. 4. Inner lamina and chitinous layer very thin compared
to outer lamina.. ............A. leptaderma sp. nov.
Hexelasma, in part, Hoek, 1913, p. 245; Pilsbry,
4 . Inner and outer laminae about equal in thickness.
1916, p. 320; Kriiger, 1940, p. 30; Withers. 1953,
chitinous layer relatively thin. .. .A. triderma sp. nov.
p. 99; Utinomi, 1965, p. 13.
Rcmarks: Aaptolasma is a tropical-subtropical genus
Diagnosis: Six plates permeated by a single row of
of five moderately large species (the largest ranging
longitudinal tubes, dividing wall into inner and outer
up to approximately 25 mm high and wide) ; occur-
laminae, as in tubiferous balanids. However, tubes
ring on the shelf and elope at depths from 100 to 800
differ from those seen in all balanids, except certain
meters. The genus is remarkable in having the wall
species of Elminius and Tetraclita, in being filled
permeated by longitudinal tubes filled solidly with a
with a translucent chitinous material throughout their
translucent, yellow, chitinous material. This type of
length. (Aaptolasma differs from Elminius and most
construction is found elsewhere in the Balanomorpha,
species of Tetraclita in having a calcareous rather
in the genera Elminius and Tetraclita, but they occur
than membranous basis, and from Tetraclita in hav-
intertidally, have a wall of but 4 plates, and generally
ing longitudinal tubes filled with chitin at their in-
lack a calcareous basis.
ception and never in more than a single row. Further-
Three of the five species contained in this genus arc
more, Elminius and Tetraclita are 4- rather than 6-
new to science. Aaptolasma callistoderma and A .
plated forms.) Basis calcareous, not permeated by
americanum were previously included in the genus
tubes; thin centrally, often thickened marginally and
Hexelasma. Hexelasma, as originally described, in-
sometimes adhering to the plates so as to appear as
cluded forms with a solid wall, but without a calcare-
inflection of wall. Tergum similar to that in Hex-
ous basis, which clearly separates Aaptolasma from
elasma, but scutum always with strong denticulations
Hexelasma.
for lateral depressor muscles. Cirri like those of
The first 2 pairs of cirri of Aaptolasma are much
Hexelasma in usually having but 2 or 3 pairs of setae
modified as accessory to the trophi, but the third
per article on posterior pairs, but number of speci-
pair of cirri, while showing some modifications in
mens insufficient to determine whether rami of cirrus
this respect, is more similar to the posterior pairs in
I11 are occasionally antenniform.
general setation pattern. This situation is like that
Type-species : Balanus callistoderma Pilsbry, 1911.
seen in Hexelasma S.S. and stands in contrast to that
p. 78.
in Bathylasma and Tetrachaelasma. Yet some species
Etymology: Derived from the Greek, a p t - , invinci- of Hexelasma have the posterior ramus of cirrus I11
ble, and elasm-, a plate, with reference to the ap- antenniform. Bathylasma and Tetrachaelasma tend
parent impregnable nature of the plates. to have both rami antenniform. In this regard then,
Distribution: Western Atlantic and Indo-West Pa- Hexelasma stands in an intermediate position, be-
cific; Recent. A . americanum is known from the tween more generalized Aaptolasma on one hand, and
western Atlantic; the other species from the Indo-West Bathylasma-Tetrachaelasma on the other. Yet this is
Pacific. not the order of similarity with respect to the oper-
cular valves and wall plates. Here Hexelasma is be-
KEY TO SPECIES OF Aaptolasma tween more specialized Aaptolasma and Bathylasma-
Tetrachaelasma.
1. Longitudinal chitin-filled tubes more or less contiguous, Aaptolasma remains closely related to Hexelasmu
forming a continuous or nearly continuous chitinous as presently defined, but is only distantly related to
layer between inner and outer calcareous laminae.. ... 4 Antarctic and North Atlantic Bathylasma. The species
1. Longitudinal chitin-filled tubes more or less arcuate in
remaining in Hexelasma are tropical Indo-West Pa-
cross section, more or less regularly spaced and sep-
arated from one another by pronounced calcareous cific. Aaptolasma, while showing its greatest diversity
bridges ........................................... 2 in the Indo-West Pacific, is found in the western At-

lantic as well, and this distribution is comparable to after the ribs have formed, their being part of the
certain other marine invertebrates such as the Xi- interlocking mechanism seems out of the question.
phosura. The similarity between Hexelasma and The slips of chitin could no doubt act as reinforcing
Aaptolasma in contrast to the other extant genera, rods in the calcareous shell, but since the species of
includes the markedly balanoid opercular parts, teeth the genus all occur in moderately deep water and
on the labrum (Indo-Pacific members), 2 or 3 pairs therefore must not be subject to much mechanical
of setae on intermediate articles of the posterior injury, this too seems unlikely or unimportant. It
cirri and the occurrence of a (?seasonally) reduced could be that the chitinous ribs act as a second-line
penis in several species. defense; as a barrier against chemical erosion and/
While a detailed study on the development of the or the drillings of gastropod mollu~csand other car-
unique wall is not included in this paper, the general bonate borers. The exterior of the wall in bathylas-
process is much as follows: The adult wall consists matids is generally covered with a persistent epicuticle
of an inner and outer lamina separated by longi- which, particularly along the pronounced growth
tudinal tubes, as in Balanus S.S. However, the tubes ridges, extends free of the surface as numerous fine
are solidly filled with a translucent chitinous material. hairs. Once this barrier deteriorates, the shell is very
The primary wall (outer lamina) is basically solid much more susceptible to chemical erosion and in-
and regularly ribbed internally (pls. XXXIII E, vasion of carbonate boring organisms, and presumably
XXXIX E ) . The ribs extend basally, where they are to predations of gastropods. In Aaptolasma, as the
initially formed, as small teeth that lock into the outer lamina deteriorates or is breached, a second chiti-
margin of the calcareous basis. The teeth are simple, nous barrier would be encountered. Although - incom-
without lateral denticulations, and therefore inter- pletely separating the two laminae in most cases, it
laminate figures are not seen in thin sections of the does reduce the contact between them by as much as
mature wall. As the ribs develop, slips of chitin are 50% in most species, and by as much as 90-95% in
deposited in the grooves between them by the hypo- A. triderma and A. leptoderma (pl. XXXIII A-C).
dermis near the basal margin. As the wall moves up- A borer then has a good to excellent chance of en-
ward with growth, an inner layer of calcareous ma- countering the chitinous barrier. This could deter
terial is applied on the interior surface, sealing off the further drilling. While Aaptolasma occurs too deep
chitinous-filled grooves and forming the inner lamina for direct observations, perhaps studies could be car-
of the shell. The junction between the two layers of ried out on certain species of Elminius and Tetraclita
the wall is readily determinable in all species, except occurring intertidally and having a comparably con-
A . americanum (pls. XXXII A, XXXVII A ) . structed wall (pl. XXXIV) . Tetraclita is of particular
The chitin-filled tubes so formed continue to com- interest in this regard because the secondary chitinous
municate with the hypodermis along the basal margin, filling develops primarily on the interior facing sur-
so that the process is continuous during growth. This faces of the large tubes within the wall (pl. XXXIV E ) .
developmental process is similar to the process of tube
Aaptolasma callistoderma (Pilsbry, 1911)
formation in balanids (Newman et al., 1967; Ross
Plates XXXII C, D, XXXV, XLIII C ; Text-fig. 77
and Newman, 1967), except for the deposition of the
chitinous filling and the more superficially applied Balanus callistoderma Pilsbry, 1911, p. 78, fig. 10,
inner lamina, in four of the five species. These dif- pl. 12, fig. 5, ~ 1 15,
. figs. 3-7.
ferences suggest that tube formation has appeared Balanus corolliformis: Kriiger, 1911, p. 55, figs.
independently in the two families. Furthermore, un- 112-114, pl. I, fig. 1, pl. IVYfig. 38.
like the situation in balanids, diametric growth is Hexdasma callistoderma: Hoek, 1913, p. 24,s;
eventually curtailed by calcareous fusion of the wall Pilsbry, 1916, p. 332, fig. 99; Utinomi, 1965, p. 12.
with the basis, as in certain chthamalids (Newman, Diagnosis: Basal margin of carinolateral plate about
1961a). 1/3 to width basal margin of lateral plate. Longi-
The development of the teeth at the basal margin tudinal chitin-filled tubes in wall not continuous with
of the wall is clearly an adaptation related to mechan- one another, arcuate in cross-section, separated by
ically interlocking with a calcareous basis, as in more or less regularly spaced broad calcareous
balanids and certain chthamalids (Newman et al., bridges. Lower margin of alae rarely hooked at free
1967), but the function of the chitinous slips is by angle, exterior surface faintly marked by growth
no means as obvious. Since the slips begin to form lines. Opercular valve white, tergal spur broadly

Growth lines of the paries stand as low, thin eleva-

tions spaced a millimeter or so apart, with minute
growth ridges between, and do not correlate with the
growth lines on the exterior of the alae (pl. XXXV
A-D). Alar growth lines are all of one type, run
parallel to the inferior alar margin for a short distance
and then diverge widely from the inferior alar mar-
gin before bending at an abrupt angle downwards
towards the alar angle. There is an abrupt change
again in the alar growth lines at the superior alar
margin forming a thin but relatively rough welting
of downward directed growth ridges, which may also
be seen from the interior along the margin of the
sheath. In this species, the welting clearly forms a
functional element of the wall. The superior alar angle
abuts against the margin of the sheath essentially as a
point; that is, there is very little contact between the
Fig. 77. Aaptolasn~a callistoderma (Pilsbry) : A, inter- two edges. The area of contact is enlarged consider-
mediate segment of cirrus VI: H, labrum, palps removed;
C, crest of labrum enlarged. Albatross, Sta. 3741.
ably by the welting, which develops just above the
alar angle, as the shell grows. While the alar angle in
this species meets the basilateral margin of the sheath
rounded, removed about y2 its width from articular in the usual manner, the margin of the sheath, raiher
margin. Crest of labrum with irregular elevations than simply abutting against the alar angle, is grooved
but untoothed. Japan, between 100 and 140 meters; so as to overlap the ala slightly. It is in the groove
Recent. that the reinforcing effect of the welting occurs, result-
Material: Albatross, Sta. 5063, off Ose-zaki, Japan ing in a much more intricately articulated wall than
(approx. 35ON, 138.j0E), 141 meters, 1 specimen; exists in other members of the family. (fig.- 70).
Sta. 3741 (approx. 35.5ON, 138.5OE), 115-124 meters, Growth lines on the sheath are conspicuous and
3 specimens. widely spaced at the carinal end, but become fine and
Depository: USNM Cat. No. 33690 (holotype, Alba- relatively inconspicuous at the rostra1 end. A patch
tross, Sta. 5068; paratypes, Albatross, Sta. 3741). has been worn smooth on each side of the carinal
sheath, apparently by being rubbed by the terga.
Supplementary Description: The cirri are quite com-
The longitudinal parietal tubes are filled with a
parable to those of Aaptolasma brintoni, except rami
yellow, translucent, chitinous material, and separate
of the first pair are subequal in length and the seta-
the paries into inner and outer laminae (pl. XXXII
tion of the intermediate articles of the posterior pairs
C, D ) . The outer or primary lamina is about 1/3 as
is more complex, there being a row of 4 or 5 short
thick as the inner lamina, and the calcareous portions,
stiff setae below the upper major pair of setae (cf.
extending at intervals between the tubes to the inner
figs. 77A, 78P). The number of articles per ramus in-
lamina, represent internal ribs which are formed
creases markedly with size, as can be seen in the counts
basally as marginal teeth that lock the wall in the
for 2 specimens given below :
margin of the basis. Dark areas seen in transverse
I I1 I11 N V VI thin sections represent areas of high organic content,
and by their orientation it can be observed that the
Specimen 1 11 xx xx 21 29 32
- - - - - inner lamina is formed of 2 parts; an inner one ex-
(Carinal height: 10 mm) 9 xx 18 21? 34 34
tending across the ribs and forming the inner wall of
12 xx 11 27 30 30
- - - - - - the tubes, the outer apparently consisting simply of
10 18 18 24 31 32 concentric layers applied to the interior after the first
Specimen 2 18
-
18
-
24
-
38
-
40
-
41
-
layer developed, rather than from the base as the first
(Carinal height: 25 mm) 16 23 30? 35? 43 43 layer was developing. The chitinous material forms
xx 17 23 40 41 44 an almost continuous lamina in the alae.
- - -
17? 22 G? 37 43 43 One of the characteristics of the Bathylasmatidae is

SI STEMATIC ACCOITNT 161
that members lack radii. Radii are defined as any Material: Albu~ross, Sta. 2663, Blake Plateau (29'
elaboration of the margin of a parietes, where it ovcr- 39'N, 79"49'W), 770 meters, 1 specimen; Sta. 2662
laps an ala so as to fill or partially fill the space that (29"24'N, 7Y043'W) , 734 meters, 4 plates.
develops between that margin and the alas-parietal Depository: USNM Cat. No. 14559 (holotype) ;
junction of the adjoining plate. An elaboration of the 48093 (paratypes).
margin can be recognized as something other than
the parietal wall by a distinctly different appearance Supplementary L)rscriptior~: High, cylindrocorlical
as compared to the paries, marked primarily by a shell and opercular valves pale orange in color; cov-
precise change in the direction of the growth lines. ered by persistent epicuticle but without trace of hris-
In the present species, there are slight marginal de- tles or hairs along growth lines. Parietal growth lines
velopments that might be construed as radii, especially widely spaced; not correlated ~ i t hfiner lines on alae,
if single rather than articulated plates were being ex- as in A. callistoderrna. Alas growth lines closely paral-
amined. The present material (Albatross, Sta. 3741 ) leling inferior alar margin, running upward toward
consists of what mere four individuals growing one on superior margin, then bending d o m n ~ a r din a rounded
top of another, the two below heirlg represented by a curve toward alar angle before turning abruptly up-
few plates supporting the others. The other two then ward again at an angle to form a welting along the
are growing on irregular and finite surfaces. Where superior alar margin. Welting visible on interior sur-
particularly restricted, definite alterations and distor- face of sheath, forming broad abutting surface against
tions to their symmetrical growth pattern have resulted. articular ridge on adjacent plate overlapping alar an-
Certain plates of these tw o specimens have overlapping gle. Growth lines in carinal sheath strongly developed,
parietal margins slightly modified and the right carino- but fine and inconspicuous in rostra1 sheath. Sheath
lateral of the uppermost specimen has what might of carinolateral plate occupying alar portion only, as is
readily be considered a radius. However, the degree generally the case in the genus.
to which these are developed is related directly to the Wall permeated by U-shaped tubes filled w i ~ ha yel-
degree of distortion of the normal growth pattern, low, translucent chitinous material, separating calcare-
which in turn has been induced by irregularities in or ous portions into outer and inner laminae-but tke
limitations of the substratum. These then cannot he actual division hetween the 2 laminae not clearly de-
construed as radii in the usual sense, but rather simply fined as in other species of the genus (pls. XXXIT
adjus~mentsmade in comperlsatiorl for dificulties in A, 6, XXXVII A, B). In transverse thin-section, the
growth; that is, they are environme~ltally induced calcareous porlions of the all appear to be tlivisihle
anomalies rather than true elenierits of the wall. Per- into 3 principal regions: (1) an outer layer, bounded
haps more has been said of this than mizht seem neces- exteriorly by the epicuticle and interiorly 11y a dark
sar), I ~ u tthis is because all contingencies are not cov- undulating line outlining what would appear to have
ered in the definition of raclii and room for misinter- been the primary wall and ribs of the shell; ( 2 ) an
pretaiion of the case in hard would o t h e r ~ i s ebe intermediate layer delimited by the undulating line
possible. separating it from the outer layer, and, towards the
interior surface, by the chitin-filled tubes and dark
Anptolasma americnnum (Pilsbry, 1916) protrusions extending between them; and ( 3 ) an inner
Plates XXXII A, B, XXXVI, XXXVII layer, bounded by the aforementioned inner margin
lIe?;ela.smn arnericanum Pilsbry, 1916, p. 330, text- of the intermediate layer and the interior surface of
fig. 98a-d, pl. 69, figs. 1-3; Utinomi, 1965, p. 12. the wall.
Diagnosis: Basal margin of carinolateral about 0.36 From these patterns the follom~i~lg growth sequence
width basal margin of lateral plate. Longitudinal can be deduced: The primary wall is solid and pro-
chitin-filled tubes not continuous, separated by longi- videcl with teeth that lock to some extent with the mar-
tudinal calcareous bridges; strongly arcuate in cross- gin of the basis. These teeth form ribs on the interior
section. Lower margin of alae slightly hooked at free of the wall, as the animal grows. Secondary calcifica-
angle, external surface faintly marked with growth tion first enlarges the ribs and then not quite com-
lines. Opercular valves pale orange; tergal spur pletely fills the spaces between them. This is sug-
broadly truncate and slightly removed from articular gested because it would be otherwise difficult to under-
margin. Labrum undescribed. Western Atlantic; stand how the concentric layers extending from the
Recent. ribs were formed at the same time the filler-material

was being applied; the former must have preceded the again at a sharp angle to form a ridged welting that
latter. Concomitant with this development, the incom- is only weakly indicated on interior alar surface
pletely filled between-spaces received the chitinous (pl. XXXIX A, B). Interior surface of wall marked
slips, and this suggests the process to this point took by longitudinal, low and regularly spaced ribs which
place very close to the basal margin. It was then fol- terminate basally as teeth that lock in margin of the
lowed by the development of the inner layer, first in basis (pl. XXXVIII F-I). Basis remarkably thick, as
a manner that brought the undulating interior surface compared to other members of the genus, surface
inward to a relatively uniform interior surface, and marked by radiating grooves representing the course
then by a series of concentric layers that line the en- of the basal teeth of the wall during growth. There
tire interior of the shell. The last step was probably is no marginal thickening of the basis in the examples
not only concomitant with but responsible for the fu- on hand.
sion of the wall plates with the basis, resulting in cessa- Wall, as seen in transverse thin-section, bears a strik-
tion of growth. ing resemblance to that of A. callistoderma, before con-
This remarkably developed wall is clearly a special- centric layers thicken the inner lamina (cf. pls. XXXII
ization over the simpler system seen in the other spe- C, XXXIII E ) . On the other hand, the laminae being
cies and, along- with the colored shell and opercular about equal in thickness, the tubes tending to be rela-
valves, sets A. americanum from the western Atlantic tively straight, and the general thinness of the wall,
well apart morphologically from the Indo-West Pacific recalls A. triderma. In this, and also by the lack of
members of the genus. intralaminate figures in the outer lamina, and dark
regions extending inward from the tubes in the inner
Aaptolasma brintoni sp. nov.
lamina, it differs from A. callistoderma and in general
Plates XXXIII E, F, XXXVIII, XXXIX A-E;
bears a closer resemblance to A. triderma.
Text-fig. 78
Opercular parts white; exterior of the scuta covered
Diagnosis: Basal margin of carinolateral about 0.58 with persistent yellowish-brown epicuticle, articular
width basal margin of lateral plate. Longitudinal margin straight, the articular furrow extremely shal-
chitin-filled tubes not continuous, only slightly arcuate low, the articular ridge relatively weakly developed.
in cross section, separated by more or less regularly These features give the plate a truly balanoid appear-
spaced, broad calcareous bridges. Lower margin of ance (pl. XXXVIII B, D ) . The scutoral adductor
alae curved but without basally directed hook at free muscle pit is deep, and an adductor ridge with a slight
angle, exterior surface weakly marked by growth lines. roughening above runs nearly parallel to the articular
Opercular valves white; tergal spur essentially continu- ridge; lateral depressor muscle pit has several crests,
ous with articular margin, truncated, and forming as it does in other members of the genus in which
-
abrupt angle (approx. 130') with basal margin. scuta are known; the occludent margin is toothed,
Crest of labrum with about 50 teeth. South China through the deletion of alternate growth lines.
- "
Sea; Recent. Of the tergum the articular margin is likewise nearly
Material: Naga Expedition, Sta. 60-212, off Da Nang, straight and continuous with the spur (pl. XXXVIII
Vietnam (15040fN, 109O22.9'E) , 110-198 meters, 1 A, C ) . The articular furrow is shallow; crests for the
complete specimen on brachiopod. lateral depressor muscles are numerous; the basal mar-
Depository: USNM Cat. No. 125311 (holotype) . gin meets the spur at a remarkably sharp angle; and
Description: Known from one complete specimen and the spur is unusually truncate. The exterior of the ter-
several calcareous bases attached to a large articulate gum is remarkably sculptured; the area from the apex
brachiopod (pl. XXXVIII) . Form conical, with spaces to the basal margin, overlying the depressor muscle
between apical ends of paries filled by alae whose su- crests, slightly elevated above the general surface; the
perior margins run nearly parallel to base. Scuta, area intervening between this and a welted ridge run-
when closed, filling aperture. Wall plates yellowish ning down the inner side of the spur furrow, low and
brown; covered with brownish epicuticle and short relatively smooth. Both sides of the spur furrow
fine bristles along beaded growth lines, but this is gen- bounded by a welting, the furrow itself flat-bottomed
erally worn away in upper areas. Growth lines on and lying below the general surface of the valve.
alae not correlating closely with those of paries, run- The labrum is broadly concave, but without the
ning upward parallel to inferior alar angle before slightest indication of a notch; the crest supports more
curving outward towards alar angle and then upward than 50 small irregular but sharp teeth, backed by

SYSTEMATIC ACCOUNT
Fig. 78. Aaptolasma brintoni gen. et sp. nov.: A-D, cirri I-IV, respectively; E, penis and pedicel of
cirrus VI; F, labrum, palps deleted; G, crest of labrum enlarged; H, right palp; I, left mandible; J, in-
ferior angle of left mandible enlarged; K, right mandible; L, inferior angle of right mandible enlarged;
M, N, right maxilla I ; 0,right maxilla 11; P, intermediate segments of cirrus VI. Naga Expedition, Sta.
60-212 (holotype, USNM Cat. No. 125311).
small bristles (fig. 78F, G ) . Palps are spatulate, mar- spines. The second maxillae are obovate, marginal
gins but not outer surfaces setose. Mandible with 4' setae divided into 2 groups, but the appendage not
teeth; second and third with small accessory knobs, bilobed.
fourth with accessory spines; the inferior angle slightly Anterior ramus of cirrus I longer by 2 segments
produced, terminating in a single spine the lower mar- than posterior ramus, wider and more heavily setose
gin of which has l or 2 small serrations. First maxil- and with setae arranged in semiwhorls at each articu-
lae with lower 2/3 of the cutting edge elevated, support- lation. Anterior ramus of cirrus I1 longer by 1 seg-
ing 12 or so heavy spines and a cluster of 4 or 5 ment, wider and with anterior margins of segments
shorter ones just above the inferior angle; the upper slightly protuberant. Rami of cirrus I11 subequal, very
1/3 of cutting edge with 2 large and 6 or so smaller similar, more than twice the length of cirrus 11, with

anterior margin slightly protuberant but bearing 3 so developed as to form a nearly continuous margin
pairs of setae per article and in general resembling around the aperture. Rostrocarinal diameter 19.5 mm,
more the posterior than anterior pairs. Cirri IV- height approximately 12 mm, aperture approximately
VI normal, anterior margins bearing 2 to 3 pairs of y5 diameter of base. Shell off-white to light, yellowish
setae per article (fig. 78D, P ) . Counts for the single brown; surface of paries covered with brownish cuticle,
specimen are given below. Penis short, but twice the marked by heavy transverse growth lines supporting
length of the pedicel of cirrus VI, without basidorsal numerous chitinous hairs or bristles. Alae likewise
point (fig. 78E). marked and bristled; growth lines correlating with
those of paries, paralleling inferior alar margin, first
upward from base before turning downward at, and
then upward to the alar angle, forming broad welting
along the superior alar margin. Welting not visible
on interior of sheath but broadly abutting margin of
alar angle; the angle not overlapped internally by
articular ridge of the sheath.
Remarks: Aaptolasma brintoni, with its high alae Carinolateral plate wide, about % width of lateral;
bounding the aperture, is more similar to A . triderma superior alar margin running subparallel to base;
than to A . callistoderma, A . americanum and A . lepto- sheath developed from apex out to alar angle, with
derma. The nature of the wall as seen in cross section peculiarly raised portions over parietal apex. Measure-
also suggests this relationship. Unfortunately, the ments for wall plates given below (in mm).
opercular valves of A . t r d e r m a are unknown, so that
Right Right Left Left
the comparison cannot be carried further. However, Ros- Lat- Carino- Carino- Lat-
the opercular valves of A. brintoni, especially the ter- trum eral lateral Carina lateral
gum, are very distinct from those of other species of Height
- - - - - eral -
11.4 - 9.5 12.4 9.4 11.0
the genus, and considering the apparent similarities Basal width 14.2 - 5.9 10.8 4.0 7.8
between A. brintoni and A. triderma in form and de- Thickness (at 1.3 - 0.8 1.4 0.8 0.9
tailed structure of the wall, one might expect that the sheath)
opercular valves would also be similar. Basis thin, calcareous, marginally thickened and
Etymology: Named for Dr. E. Brinton, Scripps In- fused to wall so that growth of shell in the individual
stitution of Oceanography; Associate Scientific Direc- on hand had been curtailed. The relatively thin wall is
tor, Naga Expedition. separated into two laminae by a thin chitinous layer
which itself is divided transversely at irregular inter-
Aaptolasma triderma sp. nov. vals by calcareous bridges (pls. XXXIII D, XXXIX
Plates XXXIII C, D, XXXIX F, G G ) . The bridges represent ribs, but are small and could
Diagnosis: Basal margin of carinolaterals about yz not have formed much of an interlocking arrangement
width basal margin of lateral plates. Longitudinal with the basis. The chitinous layer forms an appar-
chitin-filled tubes laterally elongate, thin, nearly ently continuous sheet in the wide but thin alae.
straight in cross section, separating wall into inner and Etymology: In regard to the 3 layers of the wall; 2
outer laminae connected infrequently by small, incon- calcareous, separated by a chitinous layer.
spicuous, calcareous bridges. Inferior margin of alae
Remarks: Aaptolasma t r d e r m a , with its relatively
with abrupt basally directed hook at alar angle, ex-
wide carinolateral plates, low cylindroconic profile
terior surface strongly marked by g r o ~ t hlines. Oper-
and wide, high alae, stands closer to A. brintoni than
cular and soft parts unknown.
to A . callistoderma and A . americanum, in which the
Material: Jyn 111 Expedition, Sta. 51, off Tateyama, carinolaterals are narrow to very narrow at the base,
Japan (34'54.3'N, 13g043.4'E), 549 meters, 1 speci- and the summits of the alae are markedly oblique
men on rock. rather than nearly parallel to the base. Aaptolasma
Depository: USNM Cat. No. 125312 (holotype) . triderma stands very much apart from all preceding
Description: Known from a single set of articulated species by the development of the chitinous layer, be-
wall plates attached to a rock. without opercular or soft tween the inner and outer laminae, into an almost con-
parts (pl. XXXIX F, G ) . Form cylindroconical, alae tinuous sheet, and the extension of this sheet virtually

throughout the alae. The thinness of the calcareous Malerial: Danish Expedition 1922, Amboina, Sta. 58,
portions of the wall is no doubt related to the extensive Kepulauan Kai (Kei Islands) (5O2YS, l3Z037'E), 290
elaboration of the chitinous third layer. A wall of this meters, on brachiopod, one specimen.
type is found in A. leptodernaa, taken up next. Depository: Zoological Museum, Copenhagen (Sta.
Aaptolasma leptoderma sp. nov. 58, holotype).
Plates XXXIII A, B, XL; Text-figs. 79, 80 Supplemenlary Description: The cirri are comparable
H e ~ r ~ l a s mvelutinurn:
a Rroch, 1931. p. 53, in part to other members of the genus; third pair more or less
(Sta. 59 and 251; not Sta. 105). intermediate in form between second and fourth; Icsser
Hrxrlasn~a velz~linurn: Hroch, 103I , 1). 5.3, in part curvature of inte~mediatesegments of posterior pairs
(Sta. 58, ? Sta. 3 arid 491. hearing 2 major pairs of setae; all rami riormnl (one
Diagno~is: Longitudinal chitin-filled tubes apparently antenniform) (fig. 8OA-F, H) . Basal posterior mar-
contiguous, forming a continuous chitinous layer be- gins of the posterior pairs are serrated. Cirral counts
tween inner and outer calcareous laminae; basal mar- (given below) similar to those in Aaptolasrna and
gin of caririolateral plates about width of basal Heselasma, especially H. velutinum Hoek from Siboga
margins of lateral plates; lower margin of alae turn- Station 251.
ing abruptly downward to form hook before turning
upward to join superior alar margin; exterior of alae
distinctly marked by fine growth lines; tergal spur
obliquely truncate (pointed), removed by about half
its width from articular margin; crest of lahrum con-
cave, M ith numerous small sharp teeth.
Fig. 79. Aapiola~malepiodrrn~agen. ct sp. nov.: A, labrum, palps deleted; R, rrcst of labrum enlarged;
C, right palp; D-F, righl mandible, maxilla I and maxilla 11, respectively. An~Goina, Sta. 58 (holotype,
Zool. Mus., Copenhagen).

Fig. 80. Aaptolasma Ieptoderma gen. et sp. nov.: A-C, cirri 1-111, respectively; D-F, portions of cirrus
IV; G, pedicel of cirrus VI and penis; H, intermediate articles of cirrus VI. Amboina, Sta. 58 (holotype,
Zool. Mus. Copenhagen).
The unnotched concave crest of the labrum supports mature, but it is normal for this genus, as for Hexe-
approximately 42 teeth (fig. 79A). The mouth parts lasma.
are comparable to those known for other species of Of the opercular parts Broch (1931, p. 5 3 ) said,
the genus with no particular distinctive features. The they ". . . differ a little from Hoek's description [for
penis is remarkably short, being about as long as the H. velutinum], the longitudinal furrows of the scuta
pedicel of the sixth cirrus (fig. 80G). It is strongly being very indistinct, the furrow along the tergal mar-
annulated throughout its length and the tip supports gin even failing in one of the valves." This statement
a whorl of setae, but as in other members of the family was based on Hoek's description which in turn was
it lacks a basidorsal point. This apparently reduced apparently based on the specimen figured on plate
condition might lead one to believe the form was im- XXVI, fig. 2c from Station 105. However, the state-

ment does not hold for Hoek's specimen figured on of the inner calcareous lamina. A narrow white band
plate XXVI, fig. 5c from Station 251, where it can be appears around the basal margin where the calcareous
observed that the longitudinal furrows are lacking. basis adheres and the chitinous layer and inner lamina
Only the right opercular valves are present in Broch's had not yet developed (pl. XL I, J ) . The sheath is
specimen, and the external surface of the scutum illus- elevated above the general shell surface but is not
trated here is entirely without furrows (pl. XL A ) . overhanging, and the interior surface of the sheath
The scutum is covered with a persistent yellow epi- is remarkably smooth and shiny. The area below the
cuticle and has numerous fine bristles along the growth sheath is smooth, without ribs, and marked by faint
ridges. Internally, the articular ridge extends down- whitish longitudinal lines. The exterior surfaces of
ward about 1h the length of the articular margin; the the alae are marked by fine growth ridges and these
adductor muscle pit is distinct but shallow and resides turn abruptly downward before reaching the superior
on a centrally thickened portion of the valve; lateral alar margin, to form a hook-like portion; they then
depressor muscle crests are present but weakly de- turn abruptly upward to form a thin welting along the
veloped, and rostra1 depressor muscle crests are barely superior alar margin (pl. XL F, G) .
visible (pl. XL C ) . The tergum has a persistent yel- The plates have the following measurements (in
low epicuticle and a marked but shallow spur furrow; mm) :
the spur is obliquely truncate and removed from the Right Right Left Left
Ros- Lat- Carino- Carino- Lat-
articular margin by about half its width (pl. XL B, D ) . strum eral lateral Carina lateral eral
- - - - - -
The wall is conical, appearing yellow in color when Height 9.2 9.7 10.3 10.8 8.3 -
viewed externally, due to persistent epicuticle, and Basal width 10.1 6.1 2.9 8.1 3.2 -
chitinous layer within the shell. The epicuticle is pro- Thickness (at 0.9 1.2 1.0 1.1 1.1 1.1
duced into bristles along the growth lines, but the shell sheath)
is by no means velvety. The yellow color is more pro- The basis is calcareous, but it is thin and much re-
nounced when viewed from within due to the thinness mained attached to the substratum when the barnacle
A Ba/anus (Ausfrobolonus) vesfifus Darwin Bolonus (Balanus) loevis Bruguiere

A Ba/anus (Ausfrabalonus) floscu/us Dorwin Ba/anus (8afhyba/onus/ penfacrini Hoek
v Balanus (Ausfrobalanus) imperolor Dorwin + Ba/anus (Chirona) everrnanni P I lsbry
o tBo/onus (Austrobalonus) sp. Pi lsbry
+ tBalanus spp. Hennig
Chart 11. Distribution of Balanidae.

was removed, only a narrow rim remaining attached 132O0.2'E, 204 meters) in having a velvety rather
around the basal margin of the mall plates (pl. XL J ) . than simply hirsute epicuticle; in being cylindrical
rather than conical in form; in having a scutum
Etymology: With reference to the thinness of both
marked externally by two longitudinal depressions;
the shell and the chitinous layer within.
and in having the tergal spur blend with rather than
Remarks: The specimen designated here as the type distinctly set off from the basiscutal margin. There are
of Aaptolasma leptoderma mas received from Dr. Tor- apparently no marked differences in the mouth parts,
be11 Wolff, through the courtesy of Dr. Victor A. Zullo. and the cirral counts are quite similar to those in other
It had been collected by Dr. T. Mortensen during the members of the genus. While a cylindric form can be
Danish Expedition to the ICepulauan Kai (Kei Islands) acquired by many conical barnacles when crowded,
in 1922 (Amboina, Sta. 5 8 ) , and identified by Dr. H. opercular valves and the nature of the epicuticle do
Broch (1931, p. 53) as Hexelasma velutinum Hoek, the not ordinarily vary to the degree seer1 betmeen Hoek's
species subsequently designated as the type for the two groups and we believe he inadvertently confounded
genus Hexelasma s.1. (Withers, 1913; Utinomi, 19651. two species.
Since we have been involved in a revision of this genus, With the exception of the calcareous basis (which
it was desirable to study at least one described member one must assume Hoek would have seen had there
of Hexelasmn s.s. and this specimen was obtained to been one) and the chitinous layer in the mall (mhich
satisfy this need. It was therefore with some disap- could not have been seen by Hoek in that he did not
pointment that, upon examination of the specimen. it grind thin sections), Hoek's Station 59 and 251 ma-
could not be attributed to H. velutinum Hoek, but terials appear virtually identical to Broch7s Station 52
represented a new species of Aaptolasma. material (S029/S, 132037'E7 290 meters), and it is
Broch (1931) had already mentioned certain differ- not improbable that they represent the same species.
ences seen in his material from Mortensen's Station 5U, Their inclusion in the synonymy is objective and proof
as compared ~ i t hHoek's general description of H. one n a y or the other awaits further evidence.
uelutinum, specifically the evternal features of the This problem does not meaken the generic concept
scutum, as discussed above. Upon examination it mas
surrounding these two species groups. By the nature
observed that the exterior of the specimen had a hir-
of their appendages they stand closer to one another
sute rather than velvety cuticle and a calcareous rather than to Rathylasma, but the specialization in the wall
than membranous basis, the latter characteristic being of Aaptolasnaa sets it aside from all ba~hylasmatids.
quite contrary to the generic dilgriosis of the genus
By structure of the wall, A. leptoderma is most simi-
Hexelasma S.S. Thin sections of the wall revealed a
lar to A. triderma, but the inner lamina is markedly
chitinous layer separating the calcareous wall into
thinner than the outer, rather than being about equal,
inner and outer lamina in a manner entirely compa-
and the chitinous layer is apparently coritinuous rather
rable to that seen in Aaptolasrna.
than being broken intermittently by calcareous bridges.
While detection of the irlt~icatestructure of the shell
Furthermore, A. leploderma is high and conical, with
wall was beyond the scope of the work carried out hy
the aperture markedly smaller in diameter than the
earlier workers in this group, determination of the na-
base. The opercular parts of A. triderma are un-
ture of the basis rests on a routine observation. Hoek
known; while those of A. leptoderma bear little resem-
specifically stated that Elexelasma has a membranous
blance to those of A. callistoderma and less to A. brin-
basis, and it must be assumed that this observation
toni, they are quite similar to those of A. americar~um.
was correct. That it was correct seems likely since
However, the terga are white rather than orange; the
the new species of IIe.zrrlasrna from New Zealand de-
spur is more removed from the articular margin: the
scribed here lacks both a calcareous basis and the in-
articular margin is nearly straight rather than concave;
tricate structure of the wall. We therefore concluded
and the spur furrow is below rather than flush nit11
that while the two genera are admittedly closely re-
the surface of the valve.
lated, Aaptolasma is distinct from Hexelasma.
Hoek (1913) based his description of Hexelasma
Family BALANIDAE
Leach, 1817
velutinum on s~ecimensfrom three stations. The ma-
terial from one of these (Sta. 105, 6"08/N, 121°19'E, Balanidae: Pilsbry 1916, p. 48 (ex. Tetraclitinae
275 meters) differed from the other two (Sta. 59, Gruvel, 1903, p. 160 (= Tetraclitidae nom. trans].
10°22.7'S, 123"16.5'E, 390 meters; Sta. 251, 5O28.4'S, Ross, 1968, and species previously described under

Hexelasma, included in Raihylasmatidae fam. nov. lacking in small spccimens; interior of wall with more
herein) . or less regular ribs terminating as marginal denticles
No living, benthic members of the Ralariiclae are locking with calcareous basis; scutum with lateral
known south of the Antarctic Convergence. Species depressor muscle crests; tergum with short oblic~uely
of Coronula arid Xenohalanus occur on cetaceans, truncated spur.
but owe their distribution to the migratory activities Material: Eltczrtin, Sta. 1431: off Duriedin, New Zea-
of their hosts and therefore do not constitute part of land (45°:57'S, 170°58'E), 51 meters, numerous
the resident antarctic fauna. Members of the genus spccimens or1 trochid gastropods, with B. decorus.
Balur~z~s apparently inhabited antarctic coasts in the
L)eposilory: USNM Cat. No. 125313.
Pleistocene ( IIerinig, 191I \ , hut subgeneric or specific
aflinities have not been determinctl. The following Remarks: This New Zealand species is known from
material is inclutled in this report either because it Campbell Island to North Aucklarid peninsula. Foster
was taken by the Ellanin, or hecause study was neces- (1967) says that it occurs in the low intertidal under
sary in dealing with systematic problems surrounding stones and on mollusk shells washed ashore, supgest-
higher taxa to which t ~ u eantarctic forms belong. irig it is predominantly a subtidal form. The Ellar~ir~
material, from 51 meters of water, bears out this sup-
Genus Balanus DaCosta, 1778 position. Foster (1967) has developed a very useful
Subgenus Austrobalanus Pilsbry, 1916 key for the New Zealand Balanornorpha, and in the
same paper provides in addition to notes on their
Austrobalanus Pilslsry, 1916, p. 218; Nilsson-Can-
ecology and biogeography, illustrations of the oper-
tell, 1021, p. 330; Davadie, 1963, p. 78; Foster, 1967,
cular valves.
p. 80.
Diagnosis: Wall of 6 compartments, solid, basal Balanus (Austrobalanus) jlosculus sordidus
margins roughened with irregular points and ridges Darwin, 1854
(nearly r e ~ u l a r in B. vestitus) ; basis calcareous, Plate XLVI A, Text-fig. 82
somelimes extremely thin; radii narrow or wanting, Balanus flosculus var. sordidus Darwin, 1854, p.
sutural septa irregular. Scutum with adductor ridge 290, pl. 8, fig. 5b; Fletcher, 1938, p. 115, fig. 15, pl.
long and strong; usually with crcsts for lateral de- 10, fig. I ( ? ) .
pressor muscle (Pilsbry, 1916).
Diagnosis: "Shell glohulo-conical, dirty white, with
Type-species: Balanus imperator Darwin, 1G54: Pils-
numerous sharp, narrow, longitudinal folds or ridges;
bry, 1916, p. 218.
with the internal basal cdges of the parietes rough
Remarks: The austrobalanids are presently confined with irregular points and ridges; radii narrow or ab-
to Australia, New Zealarid, and South America. sent; basis excessively thin, in appearance absent.
Withers (1953) reported the species below from Scutum with crests for the lateral depressor muscle;
1-ower Oligocene deposits of New Zealand (Chatham tergum very narrow, with the spur pointed', (Darwin,
Island), and Fletcher (1932) attributed materials of 1854, p. 290).
late Cenozoic age from Kerguelen Islands, to the Re-
Origir~ulLlescriplion: "This form is very common
cent B. ( A . ) flosculu,s sorddus Darwin from Tierra
on the tidal shores of the Strait of Magellan, and of
del Fuego. Some of the unidentified balanids from
the southernmost parts of Tierra del Fuego, near
the late Cenozoic of Graham Land (Hennig, 1911)
Cape Horn: it lives attached to rocks, mytili, and logs
may prove to belong to this austral group.
of wood, and is associated with Chthamalus scahrosus.
Balanus (Austrobalanus) vestitus Darwin, 1E54 It almost certainly is the most antarctic form of the
Plate XLI, Text-fig. El genus Balanus. If I were guided by external appear-
ance alone, I should certainly separate this form
Balanus vestitus Darwin, 1854, p. 23.5, pl. 8. fig. 3; specifically from B. flosculus, but, as will be seen in
Hutton, 1279; Gruvel, 1905, pp. 248, 249, figs. 277, the followirig description, the differences consist only
278; Broch, 1922, p. 322, fig. 61; Withers, 1923, p. in var. sordidus being much fuller and rather differ-
36; 1953, p. 77; Moore, 1944, p. 333; Foster, 1967, ently coloured, in the longitudinal folds being sharper
p. 23, fig. 5. and more prominent, and in the whole shell being
Diagnosis: Shell generally white, covered by jellow rather more globular, and on an average rather larger;
epidermis; alae tinted pink; radii weakly developcd, but in the true 12. flosculus there is considerable vari-

Fig. 81. Balanus (Ar~.strobalanus)vestitus Darwin: A, lahrum, palps dclcted; B, right palp; C E, right
mandible, maxilla I and maxilla 11, respctively; F-I, cirri I-IV, respectively; J , penis, arrow indicating
basidorsal point. Eltanin, Sta. 1431.
ation in all these respects, as there likewise is in var. examination, to find any suficient diagnostic char-
sordidus; thus some of the cylindrical varieties of acters.
the latter have less prominent ridges than even var. "The shell in var. sordidus is generally globulo-
@osculus. In general appearance I have seen some conical, dirty white, frequently with a green tinge,
nearly, but not exactly, intermediate forms; there- from the growth of confervoid matter. Orifice small.
fore, I do not feel positive that these forms may not The exterior surface is covered with numerous promi-
be specifically distinct, but have failed, after careful nent, narrow, sharp ribs or folds, the basal margin

Balanus hoekianus Pilsbry, 1911, p. 77, pl. 13, figs.

3-7, pl. 15, figs. 1, 2, text-fig. 8 a-b (for full syn-
onymy see Tarasov and Zevina 1957, p. 230).
Diagnosis: Scutum not striate longitudinally ; ter-
gum with spur furrow appearing in young stages ex-
ceeding &lo mm high, becoming progressively better
developed with growth; radii narrow, but lacking in
small specimens. Lower angle of mandible bluntly
Fig. 82. Balanus (Austrobalanus) flosculus sordidus Darwin: pointed, supporting a few knob-like teeth; becoming
A, Recent specimen from Tierra del Fuego (redrawn from expanded in older specimecs, with teeth proportion-
Darwin, 1854, pl. 8, fig. 5B) ; B, Late Cenozoic specimen from ately smaller and sharper.
Kerguelen Island (redrawn from Fletcher, 1938, fig. 1 5 ) .
Material: Albatross, Sta. 34'87, Bering Sea (52"10'N,
173'45'W), 148 meters, 6 specimens identified as B.
being serrated with projecting points where the folds
evermanni by Pilsbry, 1907a. Albatross, Sta. 4778,
terminate. When the radii are not developed, the
Bering Sea (52" 12'N, 179"52'E), 78 meters, 1 speci-
sutures are very often obscure. Internally, the shell
men designated type for B. hoekianus by Pilsbry,
is faintly tinted of a port-wine purple. In all points
1911 (USNM Cat. No. 38666).
of structure this form is identical with the true B.
posculus. In some few specimens the whole exterior Depository: USNM Cat. No. 52132 (Albatross, Sta.
surface was disintegrated and smooth; and this is 3487).
generally the case with the upper parts of the shell. Remarks: Under the discussion introducing the
Some other specimens, which had grown crowded to- Bathylasmatidae it was mentioned that Metabalanus
gether on wood, had become cylindrical, and conse- occupied an equivocal position among the generalized
quently the orifice was as large in diameter as the or simple balanids. It was therefore necessary for
shell, namely, half an inch: in some of these cylindrical
varieties the sheath was entirely soldered to the walls.
The largest specimens which I have seen were .6 of an
inch in diameter; and above one inch in height"
(Darwin, 1854, p. 292).
Subgenus Chirona Gray, 1835

Chirona Gray, 1835, p. 37; Pilsbry, 1916, p. 203;
Metabalanus Pilsbry, 1916, p. 200, fide Tarasov and
Zevina, 1957, p. 230.
Diagnosis: Parietes and basis solid; radii wanting,
or weakly to strongly developed and then sometimes
with crenulated edges; opercular parts balanoid, spur
of tergum well developed; third cirrus resembling
very much more the second than the fourth; labrum
with deep notch, crest provided with marginal teeth;
penis ( ? ) with basidorsal point; cirri without teeth
but with spinules extending onto anterior margins.
Type-species: Balanus hameri (Ascanius, 1767) :
Pilsbry, 1916, p. 203.
Balanus (Chirona) evermanni Pilsbry, 1907

Text-fig. 83
Fig. 83. Balanus (Chirona) evermunni Pilsbry: A,
Balanus evermanni Pilsbry, 1907a, p. 203, pl. 7, mandihle, Albatross, Sta. 3487 (USNM Cat. No.
figs. 7-14, pl. 8, figs. 14, pl. 10, fig. 1, pl. 11, fig. 1, 52132) ; B, mandible; C, labrum, palps deleted.
text-fig. 4a-c. Albatross, Sta. 4778 ( U S N M Cat. No. 38666).

us to examine Merabalanz~sand attempt to determine illustrated by Tarasov and Zevina (1957, p. 231, fig.
its position. As Hoek (1913, p. 245) remarked, Pils- 90) apparently would not have had radii when 8 mm
bry's (1911) description of the mouth parts, espe- high. Therefore, could Metabalanus be a young stage
cially the labrum, was incomplete, but he felt the of Chirona and if so, is it C. evermanni?
similarities great enough to place Pilsbry's species in The tergum of Metabalanus has no furrow, but in
his new genus, Hexelasma. Pilsbry (1916, p. 200) C. evermanni a deep, nearly closed external furrow
disagreed with this assisnment and returned the spe- runs from the apex to the end of the spur. While the
cies to Balanus, saying (p. 203), "The relationship degree of open or closedness of the spur furrow may
I formerly thought I saw with Hexelasma was in the vary, it is usually either present or absent in a given
superficial character of the absence of radii. Every- species. Thus by this criterion, the two forms would
thing else about this barnacle shows it to be a he distinct.
Balanus." He was nonetheless influenced by Hoek's With regard to the cirri, the first 3 pairs in Meta-
views on the matter for he erected a new subgenus, balanus are relatively uniform in length and are
Metabalanus, to accommodate it. noticeably shorter than the posterior pairs. Pilsbry
The principal difference separating Metabalanus (1916, p. 205) remarked, "The cirri do not differ
from Chirona was the apparent lack of radii, but much from Chirona except the rami of the anterior
Pilshry remarked (p. ZOO), "Whether the absence of pairs are decidedly unequal and therc are no minute
radii is sufficient to distinguish Metabalanus from spinules on the segments of any cirri." While the
Chirona is somewhat doubtful." The basis for this actual numbers of articles per ramus of normal cirri
remark is not explained, but in the discussion of may increase with age (size) in thoracican cirripeds,
Balanus (Chirona) evermanni (p. 214) he mentions, the relative proportions usually do not, and therefore
"The youngest individual seen, from [Albatross] these two forms would be interpreted as different in
station 3487 . . . has no radii." He then goes on to this respect.
describe, on the basis of observations on larger speci- The labrum of Metabalanus was neither figured nor
mens, how the radii appear and increase in breadth described. We were fortunate in being able to obtain
with growth. Tarasov and Zevina (1957, p. 230) en- Pilsbry's original preparation (USNM Cat. No.
countered this difficulty in their material, came to 38666) and prepared the accompanying illustration
consider M. hoekianus synonymous with C. evermanni, (fig. 83C). It can be observed in comparison with
and merged Metabalanus with Chirona. Pilsbry's various figures of the labrum in Chirona
From what has been said, it is clear these forms that the two types differ markedly in form. The
are balanids and need not be considered further labrum in Metabalanus is more like that in Bathy-
in the formation of the family Bathylasmatidae. How- balanus, the crest being nearly straight across, with
ever, it does behoove us to make a few comments on the shoulders at the median incision turning directly
the synonymy of Metahalanus with Chirona, proposed downward, where in Chirona the shoulders are gen-
by Tarasov and Zevina. On the basis of published erally broader and the margins of the incision more
information, there is insufficient information to fully gently sloping, tending to overlap basally in micro-
evaluate their decision. We compared M. hoekianus scopical preparations. IIowever, the labrum of C.
with the description of C. evermanni and, without evermanni illustrated by Tarasov and Zevina (1957,
additional knowledge of the ontogenetic changes that p. 232, fig. 91v) appears intermediate between the two
may occur in the latter species, were left with the types so that the character seems of little value here.
impression that the two still might be distinct species. There is a marked difference between the mandi-
The analysis was as follows: hles of the two forms, that of Metabalanus having a
The single specimen described as Metabalanus peculiarly molariform inferior angle, that of C. ever-
hoekianus by Pilsbry was relatively small, standing manni being more of the cutting type. But again
but 8 mm high, and lacked radii. Pilsbry (1916, p. the differences could possibly have resulted from
211) illustrates two 100 mm specimens of Chirona allometry in growth.
evermanni at one half natural size. By measuring At this point, no decision on the validity of the
4 mm down from the apex of the rostra, carinolaterals synonymy could be made. In order to reach a solu-
or laterals on these illustrations, it can be seen that tion, the most direct approach would be to see speci-
radii were developed when the individuals were but mens of B. evermanni small enough to lack radii and
8 mm high. On the other hand, the large specimen therefore comparable in size to B. hoekianus. A re-

quest was made to the U. S. National Museum and went on to say. "In this respect it approaches those
exactly the material required was obtained. Morr- deep sea species that are characterized by the absence
over, it was from the same lot of material Pilsbry had of radii as uell, and which for this reason I think
found the small specimen lacking radii that had raised it hest considered as forming a new genus, for which
doubts in his mind, as mentioned above. Study of the name Hexelasma is proposed in this report."
this material revealed that the radii are actually ac- Henelasma in this sense included species described
quired as the individual exceeds 10 mm or so in earlier in the genus Balanus (Challenger Expedition
length; the spur furrow also begirls to develop at this Keport, 1883), and he was inclined to continue the
stage, and the inferior angle of the mandible changes genus in the Balanidae. In this context he felt that
from that tlrscribed for Metabalanus (fig. 83A, B) IZexelasma was related to Ralanus through Batlly-
to that seen in B. ( C . ) evermanrzi. Thus, the syn- balanus. Following this lead, Pilsbry (1916) placed
onymy of the Iormer wit11 the latler, proposed by Hezelasrna in the Chthamalidae, raised Bathybalanus
Tarasov arltl Zevina, is confirmed. to generic rank and implied that it also belonged in
the Chthamalidae. Subsequent authors have simply
Sul~genusBathybalanus Hoek, 1913
followed (see Table 3 ) .
Balanus pentacrini Hoek, 1913, pp. 100,230 (Sectio : When initially confronted with what to do with
Bathy-balar~us [sic] ) , pl. XXIV, figs. 4-10. Bathybalanus, the question seemed to be simply
Bathyhalanus pentacrini: Pilsbry, 1916, p. 328; whether to leave it in the Chthamalidae or include it
Kriiger, 1940, p. 451; Tarasov and Zevina, 1957, in the new family Bathylasmatidae. As far as the
p. 251 (?=Solidobalanus s.s., cf. Henry and Mc- literature was concerned, the only thing that seemed
Laughlin, 1967, p. 4 3 ) . to preclude its being considered a bathylasmatid was
Type-species: Balanus (Bathybalanus) pentacrini the development of radii. Upon receiving material
Hoek, 1913. for dissection, the whole problem evaporated. First,
Diagnosis: Plates solid, rostrum with radii; radii Hoek had minimized the balanid nature of the labrum
well developed, with marginal denticles locking with (fig. E4A). Not only is a notch developed in a man-
adjacent plate; basis calcareous, solid, locking by ner seen in some balanids, but the teeth and chitinized
marginal teeth with regular ribs in the wall; third shoulders are also balanoid. The third cirrus is en-
cirrus rescmbling the second more than the fourth; tirely balanoid too, resembling the anterior rather
labrum with shallow notch or incision; penis with than the posterior pairs (cf. fig. 84F-H). Finally,
basidorsal point. the penis was found to support the balanoid basidorsal
point (Fig. 841). Thus Bathybalanus belongs to
Balanus (Bathybdanus) pentacrini Hoek, 1913
neither the Chthamalidae nor Bathylasmatidae. It is
Plates XLII, XLIII B; Text-fig. 84
rather clearly a balanid, as Hoek originally proposed.
S y n o n y n ~ yand Diagnosis: As for subgenus; tjpe by
Corifrorited with the question of the affinities of
monotypy (Pilsbry, 1916, p. 328).
Batlrybalarrus with other balanids, one is taken by the
Material: Siboga, Sta. 253, South of Palau Taam fact that it appears inseparable from the subgenus
(j048.2'S, 132" 13'E), 304 meters, on Pentacrinus. Solidobalanus s.s.: the wall, particularly the radii,
Depository: Zoological Museum, Amsterdam. and ribs interlocking with the calcareous basis, the
Remarks: In the introduction to the Bathylasmati- form of the scutum and deep depressor muscle pit (pls.
dae, generalized or primitive balanomorphans were XLTI, XLIII B ) , and the mouth parts and unarmed
discussed. Some of these clearly belonged to this new cirri (fig. 84) are characteristics that when taken
family while others were already well established together point in this direction. If it were not that
members of the Chthamalidae or Balanidae. Two Henry and Mc1,aughlin (1967) had greatly expanded
taxa however, hletabalanus arid Bathybalanus, occu- the concept of the subgenus by placing Hesperibalanus
pied more or less equivocal positions. Material repre- in synonymy with Solidobalanus, we would not hesi-
senting both taxa was obtained so that the problem tate to provisionally place Batl~ybalanusin synonymy
could be approached objcctively. The first has just with Solidobalanus as defined (Hoek, 1913 ; Pilsbry,
been covered; Bnlhybalanz~s is taken up here. 1916). Since it is beyond the scope of this paper
While Hoek (1913) considered Bathy1)alanus a sec- to resolve problems surrounding the union of Solzdo-
tion of Ralanus, he remarked that it was unusual in hulanus and H e ~ ~ e r i b n l a n uits , seems best to continue
the structurc of the labrum in having no notch. He to recognize Bathyhalanus as a subgenus, realizing

ANTARCTIC CIRIUPEDIA
Fig. 84. N n / n r u ~ . s (Buthqbn/cznr~s) pentucrini Hork: A, lal)ru~n,palps drlrted; R, palp; C-E, rigllt man-
diblv, maxilla I and rnaxilla 11, respectively; F-11, cirri 1, 111, and IV, resprctivrly; I, penis wit11 basi-
dorsal point. Siboga, Sta. 253 (Zool. hlus., Amstrrdam).
that as presently uritlerstootl, the designation indicates FPfclanus (Balanus) luevis luevis Rruguiere, 17:10
nothing fundamental in our knowledge of the balanids. Plates XIII, XLIV; Text-figs. 85, 86
Subgenus Ralanus da Costa, 1778 Balanus laevis: Nilsson-Cantell, 1957, p. 18 (non
Balar~us:Pilsbry 1010. p. 77, synonymy. subgen. Megabalanus) ; Davadie, 1903, p. 36 (B. loe-
Diagnosis: Wall but not radii porous; basis calcare- vis [sic] ) ; Richards and Craig, 1963, p. 112.
ous, solid,or porous: cirrus 11I armed with teeth or Diagnosis: Basis porous; radii narrow but filling
spinules (Pilsbry, 1916) . space between plates; scutum with 1 to 3 external
Type-species: Balnnus hnlanus ( LinnP, 1758) : Pils- longitudinal furrows, internal articular ridge termi-
bry, 1916, p. 49. nating below in a free point.

illaterial: Eltanin, Sta. 219, off Cape Horn (55' including the Strait of Magellan. Kolosviry (1950)
4 7 5 , 66"17'W), 115 meters; Sta. 343 (54"05'S, has described B. laevis /ossilis from the Lower Mio-
5S052'W), 119 meters; Sta. 969 (54'565, 65'06'W), cene of Hungary. The record for B. laevis from Ha-
229-265 meters. waii (Nilsson-Cantell, 1921) is erroneous as pointed
Depository: USNM Cat. No. 125314 (Sta. 219) ; out by Nilsson-Cantell (1957, p. 17, footnote 1 ) .
125315 (Sta. 343) ; 125316 (Sta. 969). Darwin's record from California, perpetuated by
Nilsson-Cantell (1957, p. 19) and Davadie (1963,
Remarks: Balanus laevis is known from the Lower p. 38) is also erroneous. Nilsson-Cantell (loc. cit.)
Miocene of Europe and North Africa (Davadie, 1963) finds it impossible to uphold Darwin's living varieties,
and Pleistocene of South America (Darwin, 1854; but our material is insufficient to add further to the
Richards and Craig, 1963), but is presently found matter. The Eltanin material is wholly typical: wall
living only around the southern part of South Amer- covered by a persistent yellow epidermis rather than
ica (S. Brazil to Tierra del Fuego and Falkland naked; scuta with one wide rather than one or two
Islands to Peru). Three varieties described by Darwin narrow external furrows; the characteristics that
(1854) were considered subspecies by Pilsbry (1916, distinguish it from B. laevis nitidus.
p. 121). One, B. laevis coquimbensis, having an
elongate compartmentalized basis and growing in Subgenus Megabalanus Hoek, 1913
clumps, occurs in Pleistocene deposits from the Strait Megabalanus: Pilsbry, 1916, p. 51, synonymy.
of Magellan to Coquimbo, northern Chile. The other Qiagnosis: Wall, basis and radii permeated by pores.
two are extant: B. laevis nitidus ranging from Peru Type-species: Balanus tintinnabulum (Linnk, 1758) :
to Concepci6n, Chile; B. laevis laevis, further south Boek, 1913, p. 158.
Fig. 85. Balanus (Balanus) laevis laevis BruguiBre: A, labrum, palps deleted; B, palp; C, man-
dible; D, inferior angle of maxilla I enlarged; E, maxilla I ; F, maxilla 11. Eltanin, Sta. 969.

Fig. 86. Balanus !Balanus) laetis laevis Bruguikre: A-C, cirri I, 111, and IV, respectively; D, penis wit11
basidorsal point. Eltanin, Sta. 969.
Balanus (Megabalanus) decorus Darwin, 1G54 Depository: USNM Cat. No. 125317.
Plate XLV; Text-figs. 87, 88 Remarks: Balanus decorus is apparently closely re-
Halanus Hecorus Darwin, 1854, p. 212, pl. 2, fig. lated to B. cnmphelli Filhol, 1885 (Gruvel, 1905;
6a, b ; Hutton, 1879, p. 328; Cliilton, 1911, p. 317; Brocli, 1922). Both are endemic to New Zealand,
Jennings, 1918, p. 60; Young, 1929, p. 158; Withers, but until recently 8. campbelli was known only from
1924, p. 25; Linzey, 1942a, p. I ; 1942b, p. 279; Campbell Island. Thus it might have been suspected
Moore, 1944, p. 333; Foster, 1967, p. 81, fig. 3. of being a southern race of B. decorus, arid Pilsbrv
Diagnosis: Tergum without beak, scuta strongly (1916, p. 54, footnole 1 ) remarked that its systematic
sculptured externally, basal margin of tergum between position was uncertain. While Broch (1922) heltl
spur and basiscutal angle concave; inner lamina of that the structural differences in the opercular valves
parietes normal; parietes and radii rose pink, the and parietes were so great that the two species seem
latter darker (New Zealand, Miocene-Recent) . distinctly separate, his material was of B. campbelli
Material: Eltnnin, Sta. 1431, off Dunedin, New Zea- from Campbell Island, where B. decorus mas not
land (45'37'S, 170°58'E), 51 meters, numerous speci- then known to occur, so that the geographical impli-
mens on trochid gastropods, with B. vestitus. cations could not be excluded.

Recently both forms have been reported in New presumed to be of Pleistocene age, are particularly in-
Zealand; B. campbelli is now known to occur on the teresting. From the illustrations it appears there are
Otago Peninsula (Foster, 1967). While this precludes several species represented. One (pl. XLVI B) bears
the problem of geographical races, it opens the door a superficial resemblance to Bathylasma, but the
to problems of ecophenotypy, at least as far as the presence of radii preclude this assignment. The degree
characteristics of the shells are concerned. However, of development of the radii makes it likely that this
Foster (op. cit.) has discovered that the opercular specimen is a illegabalanus, or Balanus, rather than
membrane in B. campbelli is yellow with a red lining, an Austrobalanus. Another specimen (pl. XLVII C)
becoming blue at the rostra1 angle, while in B. de- is remarkable in appearing to have more than six
corus it is pale pink. According to present knowledge, plates, in which case it could not be Balanus s.1. as
a single species does not vary this markedly, and in presently defined. Two other specimens (pl. XLVII
consideration of the characteristics of the shell: B. A, B) are represented only by bases, on scallop
campbelli and decorus should be considered separate shells. The radiating pattern of ribs on these indicates
species. there were well developed teeth on the basal margins
Unfortunately, the colors of the opercular mem- of the wall plates of the type seen in porous walled
branes are lost in materials fixed in ordinary pre- Balanus. A third form (pl. XLVI C) appears to be
servatives so that nothing can be said of them here. yet another distinct type of Balanus. Clearly this
Otherwise, the Eltanin material is wholly typical B. interesting collection should be restudied.
decorus.
Genus Coronula Lamarck, 1802
Balanus spp. indet. Diagnosis: Orifice of body chamber larger than
Plates XLVI B, C, XLVII A-C
basal openings; sheath as long as inner wall; radi-
Balanus sp.: Hennig, 1911, p. 10, pl. 2, figs. 3-7. ating ribs on either side of sutures unbranched or
Remarks: The barnacle remains reported as Balanus asymmetrically branched; opposed sides of terminal
sp. by Hennig (1911), from Cockburn Island and flanges crenulate; radii less than half thickness of
&BCDEF_O 25 rnm
Fig. 87. Balanus (Megabalanus) decorus Darwin: A, labrum, palps deleted; B, palp; C, right man-
dible; D, mandibular hypodermis of right mandible; E, F, maxilla I and maxilla 11, respectively.
Eltanin, Sta. 1431.

I
Fig. 88. Balanus (Megabalanus) decorus Darwin: A-C, cirri I, I11 and IV, respectively; D,
penis with basidorsal point. Eltanin, Sta. 1431.
compartmental plates, leaving cavity between radii any variety of C. diaderna (pl. 111) ; other specimens
and adjacent alae. being globulo-conical, but rather less globular than
Type-species : Coronula diaderna (Linnk, 1767) , the ordinary form of C. diaderna. The orifice is near-
p. 110, by subsequent designation. ly hexagonal, and the whole internal cavity, when the
opercular membrane is removed, can be seen from
Coronula reginae Darwin, 1854
one point of view, owing to the contraction of the
Text-fig. 89
lower part of the cavity and small size of the basal
Coronula reginae Darwin, 1854, p. 419, pl. 15, fig. membrane. The surface of the shell is smooth, but
5, pl. 16, fig. 4; Nilsson-Cantell, 1939, p. 238. has, even up to the summit, a peculiar frosted ap-
Diagnosis: "Shell globulo-conical or depressed, with pearance, different from that of the foregoing species,
longitudinal, much flattened ribs, having their edges caused by the longitudinal striae being rather more
crenated, and their surfaces striated and granulated; distinct, and being crossed by beaded, very delicate
orifice hexagonal: radii thin. not exceeding one fifth transverse lines of growth. The ends of the transverse
of the thickness of a compartment: terga absent" loops, forming the exterior surfaces of the compart-
(Darwin, 1854', p. 419). ments, are much flattened, even more so that in C.
Original Description: "Shell conical, straight-sided; balaenaris. The lines of junction between the loops,
some specimens being nearly as much depressed as are finely serrated, as in C. diaderna; and internally
C. balaenaris, and considerably more depressed than they are solidly filled up, instead of being formed into

a set of tubes by longitudinal septa, as in C. baluen- and probably in old specimens there would have been
aris. The under side of the shell, with its folded u~alls, four pairs.
presents an appearance intermediate between the "The present species differs from both the foregoing
variable appearance of this part in the above two only in its rather more conical and straight-sided
species. outline, smooth, frosted surface, and in the narrow-
"The sutural edges of the radii offer by far the ncss of the sutural edges of the radii, and consequent
most remarkable character, in their thinness from large size of the chambers between the radii, and
top to bottom; for they hardly exceed one fifth of alae. It resembles C. diadema, as far as the shell is
the thickne~sof the compartment, measured from the concerned, in the external ribs or transverse loops
external surface to the base of the sheath; hence a having their lines of junction serrated and in being
very large cavity is left between the radii and alae: solidly filled up-in the shape of the orifice and of the
in the thickness of the radii the three species already internal cavity of thc shell-in the shape and struc-
described form a series. C. reginae at one end and ture of the alae-and in the basal edge of the sheath
C. balaer~aris at the other. The sinuous plates, form- being free. It comes nearer to C. diadema than to C.
ing the lower part of the radius, are coarser and balaenaris in the structure of the radii. It differs
stand rather further apart than in C. diadema. The from C. diadema, and comes near to C. balaenaris, in
alae are thick, and have the same outline, being nar- the external ribs being flattened, instead of being con-
row at their basal margins and broad at top, as in C. vex, and in the lines of growth being very delicate.
diadema, with their sutural edges similarly construct- Rut as it resembles C. diadema in the several fore-
ed: the basal edge of the sheath likewise projects going characters of its shell, in the opercular valves,
freely. in all parts of the mouth (excepting the labrum), and
"Opercubm. This resembles most closely that of in the cirri, it is very much more nearly related to
C. diadema. There are no rudiments of terga. The that species than to C. balaenaris" (Darwin, 1854,
scuta cannot be distinguished from those of C. dia- pp. 4 2 M 2 1 ) .
dema.
Csronula diadema (LinnZ, 1767)
"Mouth. The labrum has a row of inwardly curved
Plate I11
little teeth along the whole crest, and these I did not
notice in C. diadema: there is only a trace of the Coronula diadema Darwin, 1854, p. 417, pl. 15,
prominence on the outside at the bottom of the cen- figs. 3-3b, pl. 16, figs. 1, 2, 7 ; Nilsson-Cantell, 1939,
tral notch. The hails on the basal exterior margin of p. 237.
the palpi are moderately long. The mandibles have Diagnosis: "Shell crown-shaped, with longitudinal
five teeth. In the Cirri, the first pair resembles in its convex ribs, having their edges crenated; orifice hex-
peculiar structure those of C. diadema. In the sixth agonal: radii moderately thick, very broad: terga
pair, however, the segments support only three pairs absent or rudimentary" (Darwin, 1854, p. 417).
of main spines: but the specimen was not very large, Supplementary Description: "The crown-like shape
of the shell is well expressed by its name of Lliadema,
but the crown tends to pass into a cylinder. The
radii are extremely broad. The orifice is large, and
neatly hexagonal: when the operculum is removed the
whole inside of the cup-formed shell can be seen at
once, for the flat membranous basis is much smaller
than the orifice. The under side of the shell is deeply
concave. The outside of each compartment is formed
by broad, rounded, and somewhat prominent, rarcly
divided, ribs (i.e., the transver~eends of the folded
walls) ; these ribs are closely united together by finely
serrated lines of junction. Their surfaces outside are
delicately striated longitudinally, anti plainly cros~ed
Fig. 89. Coronuln reginae ]Darwin: On l~umpbacli by irregular, transver~eridges, especially in the lower
. fig. 5 ) .
whales (redrawn from Darwin, 1854, ~ 1 15, part of the sh1.11. The largest sprcimen which I have

180 ANTARCTIC CIRRIPED1.h
seen Mas two and a half inches in diameter and two outside thr bottom of the medial notch, there is a
in I-wight. small hard prominence. The palpi are broad; on their
"Scuta. These are placed close together at the ros- basal exterior margins there is a short row of spines,
tral end of the orifice, and are imbedded in the b r o a n - which do not equal in length the width of the palpi,
ish, tough, longitudinally plicated, horny substance, and therefore are not so long as in C. balaenaris. The
which extends far beyond both ends of the valves. In mandibles have five main teeth, of which the second
outline they are mitre-shaped, or rounded and sub- arid third show only an obscure rudiment of being
triangular, a little curved, and more or less elongated, tlouble; between these two teeth, and between the
being most so in young specjmens; they are, however, third and fourth tooth, there is a small intermediate
1e.s elongated and rather more massive than in C. tooth: the inferior angle is narrow, rounded, and
halaer~nris. Terga,-these seem entirely absent in most ~pinose. The extremity of the apodeme of the maxil-
specimens; but in one I found a ludiment, namely, lae is expanded.
a short thin plate of shell, barely visible to the rialied "Cirri. I have only to remark, illat the pedicel of
eye, extending parallel and near to the tergal margin firs[ cirrus is extremely broad, and that the rami are
o l the scutum. The lips of the aperture of the sack are set on in an unusually crooked manner: the basal
prominent, and highly so towards the caririal end. segment of the shorter and broader ramus of this
"Strzrcture oJ Shell and Radii. Owing to the sllell cirrus has its dorsal surface produced into a plate
not spreading much at the base, new folds in the fringed with very fine hairs" (Darwin, 1854, pp. 418-
walls are much seltlorner formrd, and therefore the 419).
external longitudinal ribs (i.e. the terminal trans- Genus Xenobnlanus Steenstrup, 1851
versely elongated loops), are much seldomer divided,
Diagnosis: Lengthened body not contained i11 shell
than in C. balaenaris or regir~ae;even rather large
wall; shell star-shaped, small, composed of 6 corn-
specimens sometimes having only the origirial eighteen
partmental plates emheddecl in skin of host; oper-
folds. Whcn new folds are formed, only one is formed
cular valves ahsent.
on one, viz., the rostral, side of each suture, instead of
Type-species: Xcnobalanus globicipitis Steenstrup,
on 130th sides, as in C . balaenaris The inner erids ol
original
the folded walls, surrounding the basal membrane,
are narlow, instead of being almost square, as in C. Xenobnlanus globicipitis Steenstry, 1351
balaenaris. The lower edge 01 the sheath, which pro- Plate XLVIII
jects freely, descends almost to the level of the basal Yenobalanus g l o l ~ i c l p i t i ~ :I)arwin, 1854, p. 440,
edges of the walls. The outer ends of the folded walls, pl. 17, figs. 4a-c.
forming the transverse loops, are interr~allyfilled up "General Appearance. The shell is in an almost
solidly by calcareous matter, insteail of by septa rudimentary condition, arid appears like a small white
forming tubes, as in C. balaenar5s. Thr ratlii are a irregular star, imbedded up to its top in the skin of
little thicker in the lower and outer than in the upper the porpoise. Out of this thin, star-shaped shell, a
and inner part of each compartment; in the middle, cylindrical flexible, prduncle-formed botly springs,
they do not reach the sheath by about half the thick- rzhich forms the main part of the animal; it is nar-
nes5 of tl.~ecompartmcnt, and consequer~tly they are row where coming out of the central cavity of the
sepaiated from the plates on which the alae rest, by star, but soon acquires its full diameter; at the upper
laige cham1)ers which extend up to n c a ~ l ythe apices end it has a reflexed hood, and hence is broader, and
of the t oinpartmerits: the extent, however, to which this has the appear~xriceof forming a capitulum, like
the upl'e' cnds of these chamhe~rhave been solidly that of a pedunculated Cir~ipede. This pseudo-capi-
filled up, varies a little. The sinuour plates forming tulum is formed by a membranous reflexed collar or
the main portion of the compour~d ratlii are rather hood, which is kery narrow at the lower end of the
thinrier and closer together than in C. Oaluer~arzu. orifice, close under the mouth, and hecomes wider and
The alae are thick, being thickest in the midtlle part, wider towards the upper and carinal or posterior end
and there equal the radii in thickness; their lower of the orifice; hen(,? the lower reflexed edge of the
margins are 1 ery short compared with their upper hood is only slightly oblic~ueor even nearly transverse.
margins, hence they are almost wedge-formed. The orifice leading into the sack is large, arid nearly
"Mozrth. The teeth and fine hairs on tlie labrum in the same straight line with the peduncle; it is a
are sometimes obscure, and sometimes plain: close little l~ollowed out in the middle at tlie upper end,

and on each side of this medial hollow, there is a membrane. The transverse ridges are surmounted by
small rounded projection or horn, not perforated, but knobs arising from the longitudinal septa; and the
hollow, as may be seen by turning up the hood and knobs themselves are capped by other little heads,
looking at its under side. These two little horns curi- which are not represented in the drawing.
ously bring to mind the ear-like appendages in Con- "The internal cavity of the shell is small: it ap-
choderma aurita (Otion), but these latter are per- proaches a hexagon in shape, with the rostral side
forated, open into the sack, and point outwards. The very short, and the lateral sides curved inwards. It
peduncle-formed body answers to the main part of is lined by a rather thick sheath, which descends
the shell in Tubicinella, and the hood, as it would very near to the basal membrane; the sheath is di-
appear, to the lips of the sack-aperture, which project vided into very distinct, successive convex zones of
between its scuta and terga; of these valves there is growth. The external membrane of the pseudo-pe-
not here a trace. The whole surface is smooth, and duncle is attached with remarkable strength to these
is formed by rather thin membrane, of an orange rib-like zones of the sheath. The alae are represented
colour; but from the colour of the underlying corium, by mere angular shoulders, received into very slight
the whole appears of a dark chocolate red, the re- notches, and placed at the inner ends or entrances of
flexed hood being rather lighter coloured. the double walls, or rays. With respect to the radii,
"The largest specimen which I have seen was very they also are in an extremely rudimentary condition;
nearly two inches in length: in this specimen the but a thin layer of shell, apparently continuous and
star-shaped shell measured, from extreme point to homologous with the sheath, extends from the sheath
point, nearly a quarter of an inch in diameter, but along both sides of each ray, and on the rostral side
the internal cavity only about one eighth of an inch. (whence the radius ought to arise), about half-way
"Structure of Shell. The almost rudimentary shell from the end of the ray, gives rise to a projection or
consists of a small, thin, six rayed disc, formed of ridge which runs from the top to the base of the shal-
six compartments, each of which, instead of being low shell. From this longitudinal ridge, septa, parallel
outwardly convex, is deeply bowed inwards. The to the basis, extend to nearly the extremity of the ray
narrow sutures separating the six compartments, run or double wall. These represent the radii; but they
along the middle of the six rays, each ray being never grow, so as to increase the diameter of the shell.
composed of the bowed ends of the walls of the ad- These radii evidently correspond to the additional or
joining compartments. The rays are a little curved pseudo-radii in Coronula, which in that genus lie be-
towards the carinal end of the shell. It is remarkable tween the parallel, as here, and folded parietes. Of the
that the rostrum is smaller and less deeply folded true radius, having the same thickness as the paries, I
inwards than the other compartments, and the lateral here saw only traces of an internal, very slight, longi-
compartments are a little smaller than the carino- tudinal ridge running up the shell, close to the outer
lateral compartments, which is exactly the reverse extremity of each ray or double wall.
of what is usually the case. Only about four zones "The membranous Basis is united all round to a
of growth have been preserved in any specimen, and rather wide flap of membrane which is inflected from
consequently the shell is very nearly of the same the outer surface of the shell. The cement-glands
diameter at the top and bottom; for the upper end appear to be mere enlargements of the cement-trunks,
of the shell is rapidly removed, as in Tubicinella, by which latter extend in two nearly straight and paral-
the scaling off of the upper rims of the sheath, and lel lines. From each gland two cement-ducts proceed,
by the disintegration of the walls. The zones of one of which runs parallel to the cement-trunk.
growth are commonly not piled exactly over each "Mouth. Labrum unusually prominent, as meas-
other, but rather obliquely. Each zone projects, form- ured from its basal margin to the crest, which is but
ing a prominent, sharp, toothed ridge round the shell. slightly notched, hairy, and without teeth. Palpi
In Coronula and its allies, the outer lamina of the broad, heart-shaped, clothed on their inner sides by
wall is formed by the union, a little above the basal a thick brush of spines, which here, as on the other
margin, of ledges running along the sides of the longi- gnathites and cirri, are almost all doubly serrated.
tudinal septa. In Xenobalanus similar ledges are less On the outer margin of the palpi there are a few
perfectly joined, and apertures seem always to be longer spines. Mandibles villose, with five teeth, of
left in transverse rows under the transverse toothed which the fifth is very small and of irregular shape:
ridges. The apertures, of course, are covered by the inferior angle is broad and pectinated. There are

no intermediate teeth between the second, third and of bristles: there is no knife-edged projection at its
fourth teeth, as in the three foregoing genera. Had I dorsal base.
not known that the lower main mandibular teeth were "Imbedment. The shell is imbedded up to its sum-
always laterally double in the Balaninae, and had I mit, but the shell is very shallow. The imbedment
not observed how obscure this structure was in Coro- seems due either to the compression of the epidermis
nula and Tubicinella, I should have overlooked the of the porpoise, or to its formation beneath the shell
merest vestiges of double teeth in the present genus; having been checked; the outline of the true skin under
indeed, in some specimens the teeth seemed to be ab- the dark-coloured epidermis is not in the least affected.
solutely single. The maxillae are villose; their edge The epidermis fills up the bay-like spaces formed by
exhibits a trace of being notched under the two great the inwardly folded walls, and firmly adheres to them.
upper spines. The outer maxillae are bilobed, but "Afinities. The genus presents very varied affinities
not very plainly: between these organs there is no to Tubicinella, Coronula, and Platylepas. To the latter
little prominent mentum, as in the three previous it is more especially allied in the compartments, being
genera. singly folded inwards, though the sides of the folds
"Cirri. The cirri are short, particularly the three are not here closely pressed together, as in Platy-
anterior pairs. The segments in all are singular, from lepas: in both these genera the fold is less deep in
being so much compressed, so short, and of such great the rostrum, of which fact, in Platylepas, the final
breadth; they are protuberant in front. In the second cause is evident, but here there seems no cause, ex-
and third cirri, the broad lateral faces of the seg- cepting the simple one of affinity. Xenobalanus is
ments, with the exception of the posterior face of the further allied to Platylepas, in the lesser size of the
posterior ramus, are almost bare of spines. In the inner fold of each branchia, compared with the outer,
three posterior pairs of cirri, the segments are pro- and in the structure of the cement glands, and to a
tuberant in front, and support three pairs of short certain extent in that of the sheath. To Coronula
thick spines, with an intermediate tuft; the dorsal the special alliance is shown by the remarkable char-
tufts are unusually small: their pedicels are remark- acter of the pseudo radii lying between the parallel
able from the upper segment, and the upper part of and adjoining walls, and in the general character of
the lower segment, being produced into a rounded the cirri: in Coronula we have the terga sometimes
protuberance, dotted with spines. In the third pair quite aborted, and the scuta of small size, thus ex-
there is only a trace of this structure; and in the hibiting a tendency to the entire absence of opercular
second pair the anterior margin of the pedicel is valves, so remarkable in Xenobalanus. To Tubicinellu,
straight, and clothed with three tufts of bristles. The the alliance is still more plainly shown in the external
pedicel of the first cirrus is very broad, and clasps shape of the whole animal, wider at top than at bot-
the mouth. tom,-in the opercular membrane descending almost
"The cirri arid mouth are dark chocolate red, like to the very base-in the relative positions of the differ-
the outside of the animal and the upper part of the ent parts and organs-in the upper end of the shell
sack. The thorax is redder and paler. The four continually scaling off-in the prominence of the
posterior articulations of the thorax are straight and transverse external ridges-in the sheath being- di-
transverse; the next segment, or that corresponding vided into successive zones of growth, and being pro-
with the second pair of cirri, is slightly inflected, in longed nearly to the basal membrane-and in the
the usual way, towards the prosoma. The prosoma edges of the alae being smooth. Internally, the re-
is pale coloured, extraordinarily elongated, and blunt- semblance is also plainly shown, in the strength of
ly pointed; it extends down about one third of the the internal tunic of the sack-in the branchiae
length of the whole animal. The orifice leading into springing from two approximate fleshy crests-in the
the acoustic sack forms a freely depending little point freely. depending acoustic orifices-in the form of the
-
beneath the basal articulation of the first cirrus. The thorax and prosoma-and consequently of the alimen-
stomach in the uppermost part is deeply and closely tary canal" (Darwin, 1854, pp. 440-445).
plaited lori;itudinally, but has no caeca; it runs down
to the lower point of the prosoma, and is then doubled Order RHIZOCEPHALA Miiller, 1862
back on itself, so that it is very long. Diagnosis: Parasitic crustaceans without appendages
"Generative System. The probosciformed penis is or digestive tract in adult stage; recognized as Cir-
short and thick, and covered with very minute tufts ripedia by characteristic nauplius larvae or cyprid

stage, or both. Parasitic on crustaceans, primarily Original Description: "External cuticle thick and
Decapoda. callous, with grooves and shallow pits. The surface
is covered with small excrescences, which have a
Suborder KENTROGONIDA Delage, 1884 length of approximately I) p and form a dense cov-
Diagnosis: Rhizocephala with kentrogon stage es- ering of the outer layer. Internal cuticle with retina-
tablishing endoparasitic phase; development generally cula of large size, containing numerous spindles,
including naupliar stages. which vary in length from 15 p to 55 p.
"The specimen has a length of 98 mm., its height
Family PELTOG-~STRIDAE
Lilljeborg, 1861 attains 53 mm., and the thickness amounts to 31 mm.
Genus Briarosaccus Boschma, 1930 As compared with all other Rhizocephala hitherto
Diagnosis: "Body slightly elongate, curved. Stalk known it really is a gigantic animal. The thick ex-
about in the central part of the dorsal surface. Mantle ternal cuticle possesses a number of grooves which
cpmin:, at one extremity. Mesentery from the mantle spread from the stalk toward the ventral region of
opening to the posterior part of the dorsal surface. the parasite. Between these grooves and especially at
thin, except in the region of the stalk. Collecteric the ventral region the surface of the mantle shows
at the left and right side of the mesentery, numerous small depressions, giving the surface a
highly lobular. Testes paired [ ?male cell receptacles], dotted appearance. The stalk is surrounded by a
situated in the dorsal part of the visceral mass, parallel strong shield, consisting of chitin of a harder kind
to long a-cis" (Boschma, 1930, p. 1). and of a darker color than the remainder of the
external cuticle. The mantle opening is found at one
Briarosaccus callosus Boschma, 1930 side of the animal, which consequently has to be re-
Text-fig. 90 garded as the anterior pole. It is surrounded by a
Briarosaccus callosus Boschma, 1930, p. 2, figs. 1-8; thick sphincter, as a result of which the parts around
Boschma, 1962, p. 58, text-figs. 1-24, pl. 1 (= Pelto- the mantle opening slightly protrude. forming thereby
gaster sp. Weltner, 189213, p. 3 ) . a kind of wall" (Boschma, 1930, pp. 2-3).
Fig. 90. Briarosaccus callosus Boschma: from an anomuran crab (Lithodes agassizii
Smith) ; A, individual from right side and B, from left side ( f r o m Boschma, 1962, fig.
1, A, B ) .

APPENDIX 1. Localities and Bathymetry of Previously Known Antarctic Cirripedia
North of
Species South of Antarctic Convergence Antarctic Convergence Source Remarks
Ascothoracica:
Ascothorax bulbosus South Georgia (approx. 55"S, 37"W) Hecgaard, 1951 :186 On Amphiura belgica and
Heegaard, 1951 100-175 meters A . microplax
Acrothoracica:
j-Brachyzapfes elliptica var. Alexander Island, east coast; 73"S, 68"W, Taylor, 1965:39 In belemnite rostra
gigantea Taylor, 1965 fossil material; Upper Aptian,
Cretaceous
Rhizocephala:
Briarosaccus callosus South Georgia (approx. 55"S, 37"W) ; Boschma, 1962 :58 On Paralonzis granulosa and
Boschma 1930 13-728 meters Lithodes antarcticus
Falkland Islands; 11-207 Boschma, 1962:58 On Paralomis granrrlosa and
meters Lithodes untarctic~~s
Off SE coast of United Boschma, 1962 :58 On L. agassizii;
States; 494 meters off Fernandina,
Florida ( type-locality)
54"2'5OUN, 16G045'W Boschma, 1962 :58 On L. aequispinus
(Bering Sea) ; 742 meters
Strait of Magellan Boschma, 1962 :58 On P. granulosa
Thoracica:
Lepadomorpha :
Arcoscalpellum angulare 6lo25'30"S, 53"46'W (Clarence Island) ; Nilsson-Cantell, 193011:239 Single record
(Nilsson-Cantell, 1930) 342 meters
A. antarcticum (Hoek, 1883) 65"4ZtS, 7go49'E; 3065 meters Hoek, 1883:95 Single record
A. aurorae (Bage, 1938) 66"32'S, 14lo39'E ; 287 mrters Rage, 1938:6 Single record
( =Litoscalpellum)
'4. berndti (Gruvel, 1907) Gauss, winter station (approx. 66"S, Gruvel, 1907a:161; Single record
90°E) ; 350 meters 1909:207
A. bouveti (Nilsson-Cantell, Bouvet Island (approx. 56"S, 3"E) ; Nilsson-Cantell, 1939:233 Single record
1939) 40-45 meters
A. bouvieri (Gruvel, 1906) Flagon Point; McMurdo Sound, Ross Gruvel, 1906:272; 1907:3 Type-locality; on hydroids
Sea; 18-37 meters
Approx. 78"201S, 171°00'W (Ross Sea) ; Nilsson-Cantell, 1926 :2
550 meters
65"34'S, 109"12'E; 515-550 meters Zevina, 1964 :252
67"45'S, 147"lO'E ; 900-920 meters Zevina, 1964 :252
(Knox Coast to Ross Sea)
65"34'S, 109"lZ'E ; 515-550 meters Zevina, 1968 :92

APPENDIX 1. (Continued)
North of
-- -
A . bouvieri (continued) 67"45'S, 147"lO'E; 9CC920 meters Zevina, 1968 :92

67"41'S, 45"45'E; 7C157 meters Zevina, 1968 392
A. breuecarinatlim (Hoek, 1883) 46"46'S, 45'31'E; 2475 meters Hoek, 1883:82
46"16'S, 48"21'E; 2880 meters Hoek, 1883:82
59"25'S, 97"23'E ; 4540 meters Zevina, 1%4:253

65"34'S, 109"12'E ; 515-550 meters Zevina, 19643253
A . compactum (Borradailr, Off Granite Harbor, McMurdo Sound, Borradaile, 1916:130 Type-locality
1916) Ross Sea; 92 meters
64"201S, 56"38'W (S.E. of Seynlour Nilsson-Cantell, 1921:I98 On ~ o l ~ c h a etube
te
Island) 150 meters
64"36'S, 57"42'W (S.W. of Snow Hill Nilsson-Cantell, 1921 :I98
Island) ; 125 meters
75"56'S, 178"35.5'W (Ross Sea) ; 567 Nilsson-Cantell, 1939:229 On hydroids
meters
A. convexum (Nilsson-Cantell, 54"11'S, 3h018'W (South Georgia) ; Nilsson-Cantell, 1921 :I94 Type-locality
1921) (=Litoscalpellum) 252-310 meters
53"51'S, 36"21'30"W (South Georgia) ; Nilsscn-Cantell, 1930b:244
200-236 meters
Cumberland West Bay, South Georgia; Nilsson-Cantell, 1930b :244
110 meters
Stromness Harbor to Larsen Point, South Nilsson-Cantell, 1930h:244
Georgia; 122-136 meters

A. discoveryi (Gruvel, 1906) Discovery, winter quarters; 9 meters Gruvel, 1906:271; 1907a:Z
(=Litoscalpellum)
7"56'S, 164"12'E (E. of Borradaile, 1916:128 Terra Nova, Sta. 340
Solomon Islands) ; and 356
293 meters
Off Granite Harbor, McMurdo Sound, Borradaile, 1916:128 On Amrnothea glacialis
Ross Sea; 92 meters
74"25'S, 179"3'E (Ross Sea) ; 289 meters Nilsson-Cantell, 1928:ll Terra Nova, Sta. 294
Commonwealth Bay (approx. 67"S, Bage, 1938:s On pycnogonid
142"E) ; 46 meters
6lo25'30"S, 53"46'W (Clarence Island) ; Nilsson-Cantell, 1939:230 On Colossendeis sp.
342 meters
From 54"04'S, 36"27'W to 53"58'S, Nilsson-Cautell, 1939:230
36"26'W; 155-178 meters
Vol. 14
A. gawsi (Gruvel, 1907) Gauss, winter station (approx. 66"S, Gruvel, 1907a:159, Single record
90°E) ; 380-385 meters 1909:204
North of
A . improvisum (Hoek, MS) 53"55'S, 108"35'E (Kerguelen Islands hlurray, 1893 :SO9
region) ; 23054755 meters
A. liberun~(Nilsson-Cantell, 64"48'30MS, 63"3lr30"W (Palmer Archi- Nilsson-Cantell, 1930b:233 Single record; on gorgonian
1930) pelago) ; 259 meters
A . n~agnaecarinae (Nilsson- 64"48'30"S, 63"31130"W (Palmer Archi- Nilsson-Cantell. 1930b :236
Cantell, 1930) pelago) ; 259 meters

64"56'S, 65"35'W (Palmer Archipelago) ; Nilsson-Cantell, 1930h :236
315 meters
A . njmphonis Borradaile, 1916 W. shcre Mc3Iurdo Sound, Ross Sea; Uorradaile. 1916: 129 On Nymphon sp.;
( = A . discoveryi) 146 meters type-locality
A . recrtrvirostrurn (Hoek, 52"04'S, 7lo22'E; 274 meters Hoek, 1883: 77 On rocks; single record
1833)
A . triangulare (Hoek, 1883) 37"17'S, 53"52'W; Hoek, 1883:130
1080 meters
Near Ballenj Islands; 6 i o S , 168"E Zebina 1964 :253
A. ventricosr~m(Hoek, 1907) 1Oo35.6'S, 124"11.7'E; Hoek, 1907h :98
2050 meters
5Oo11'S, 50°50'W; 2675 meters Nilsson-Cantell, 1921 :205
South Georgia (approx. 55'5, 37'W) Nilsson-Cantell, 1921 :205
A. weltneri (Gruvel, 1907) Gauss, winter station (approx. Known only from Antarctic Gruvel, 1907a: 159 On hjdroids and bryozoans;
66"S, 90"E) ; 350, 380, 385 meters 1909 :205 10 separate collections
Scalpellum korotkevitshae 67"41'S, 4So45'E; 70-157 meters Zevina, 1968 :89

Zcvina, 1968
i=L;toscalpellum)
S. leoni Zevina, 1968 69"211S. 14"06'E; 67&830 meters Zevina, 1968 :90
( =G) mnoscalpellum)
S. vanhoffeni Gruvel, 1907 Gauss, winter staticn (approx. Known only from Antarctic Gruvel, 1907b:138; On brjozoans; 28 separate
66"S, 90"E) ; 350, 380, 385 meters 1909:202 collections (1909)
~Euscalpellumnntarcticum 63"55'S, 57"301W; 63"59'S, 57"25'W; Withers, 1951 :157
Withers, 1951 (Graham Land) ; fossil (U. Cretaceous)
tZeugrnnto1ep;as georgiensis Annekov Island, South Georgia; Aptian Withers, 1947: 18
Withers, 1947 (Cretaceous)
tCretiscalpellum apttensis Alexander Island, E. coast; 73"S, 68"W; Taylor, 1965:37; Withers,
var. antarcticum Taylor, Upper Aptian (Cretaceous) 1935 :147
1965
South Shetland Islands (approx. Gruvel, 1911a:292 On Balaenoptera borealis
Vol. 14
Conchoderma auritztm
(LinnC, 1767) 62"S, 58°F') ; on whales and Megaptera longimana I
ca
-1
-. .-
North of
.-
C. auritum (continued) South Georgia (approx. 5S0S, 37"W) ; Nilsson-Cantell, 1921 :240 On Coronula diadems on
on whales Megaptera boops
South Georgia (approx. 5S0S, 37"W) ; Nilsson-Cantell, 1930b :249 On Coronula reginae on
on whales blue whale
South Georgia; on whales Nilsson-Cantell, 1930b :249 On Coronula diadema on
humpback whale
South Shetland Islands (approx. Nilsson-Cantell, 1930b:249 On teeth of sperm whale
62"S, 58"W) ; on whales
South Shetland Islands (approx. Nilsson-Cantell, 1930b:249 On Coronula diadema on
62"S, 58"W) ; on whales humpback whale
South Shetland Islands (approx. Cosmopolitan; on Coronula; Nilsson-Cantell, 1930b:250 On baleen plates of
62"S, 58"W) ; on whales on whales (teeth and ba- fin whale
leen plates), fish, eels, ships,
buoys and floating objects
57"20'S, 32"W; on whales Nilsson-Cantell, 1939:235 On Coronula diadema on
humpback whale
South Georgia (approx. 55"S, 37"W) ; Nilsson-Cantell, 1939 :235 On baleen plates of
on whales fin whale
South Georgia (approx. 55'S, 37'W) ; Nilsson-Cantell. 1939:235 On Coronula reginae on
blue whale
on whales
South Georgia (approx. 5S0S, 37'W) ; Nilsson-Cantell, 1939:235 On teeth of sperm whale
on whales
Lepas australis Darwin, 1851 Antarctic waters, circumpolar; on Tasmania (Van Diemen's Darwin, 1851:89 On floating objects; hull of

floating objects Land) New Zealand; HMS Beagle; Laminaria, etc.
Patagonian coast; on float-
ing objects
Tasmania Guiler, 1952 :20 On driftwood
Region of Kerguelen Islands ( ? ) Honolulu, Hawaii ; on Weltner, 1900 :306
floating objects
Off Cape of Good Nope, Gruvel, 1907a: 161; On Macrocystis sp.
S. Africa; on floating 1909:224
objects
Off St. Helena; on floating Gruvel, 1907a: 161; On ship bottom
objects 1909:224
Between Stewart Island Nilsson-Cantell, 1926:7 On algae
and Campbell Island, New
Zealand
Vol. 14
42"36'30"S, 18O19'30"W; Nilsson-Cantell, 1930b:247 On Macrocystis sp.

on floating objects
North of
A
-. --
L. australis (continued) 48"50tS, 53"56'W ; Nilsson-Cantell, 1930b:247 On /Macrocystis sp.

on floating objects
65'155, 64"42'5OUW (Drake Passage) Nilsson-Cantell, 1930b:247 On Macrocystis sp.
53"40'S, 6lo10'W; Nilsson-Cantell, 1930b:247
on floating objects
48"501S, 53"56'W; Nilsson-Cantell, 1939:235 On Macrocystis sp.

on floating objects
52"19'S, 52"11'W; Nilsson-Cantell, 1939:235 On c macro cyst is sp.
on floating objects
46"S, 6O0O5'W ( ? 150 Nilsson-Cantell, 1939:235 On Larninaria sp.
meters)
48"S, 99"E; on floating Nilsson-Cantell, 1939:235 On algae
objects
Islas Juan Fernindez; Val- Nilsson-Cantell, 1957:6 On algae
paraiso, Chile, south to
Cape Horn; on floating
objects
L. australis zveltneri Nilsson- Islas Juan Fernindez; Tal- Nilsson-Cantell, 1929:487 On .Macrocqstis sp.
Cantell, 1929 cahuano, Chile; on floating
objects
Lepas hillii (Leach, 1818) Macquarie Island, south of New Zealand Cosmopolitan on floating Rage, 1938:7
objects

Verrucomorpha :
Verruca (Altiverruca) 48"37'S, 55"17'W; Hoek, 1883: 134 Type-locality
gibbosa Hoek, 1883 1892 meters
7"51'N, 2lo39'W; Nilsson-Cantell, 1928:30
3056 meters
29"55'S, 178"14'W; (Ker- Nilsson-Cantell, 1928:25 Type-locality for
madec Islands) ; 952 meters V . sulcata
29"45'S, 178"111W (Ker- Nilsson-Cantell, 1928:25
madec Islands) ; 1153 meters
70°S, 80°48'W (approx.) ; 555 meters Nilsson-Cantell, 1928:25 Type-locality for
V . mitra
4lo07'N, 65"26'30"W ; Nilsson-Cantell, 1928:25 Type-locality for
3129 meters V. bicornuta
6"18.8'N, 49"32.5'E (E. Nilsson-Cantell, 1929a :470
Vol. 14
V . (ti.) gibbosa somaliensis

Nilsson-Cantell, 1929 Africa) ; 1079 meters
I-
APPENDIX 1. (Continued) \O
o
North of
V. (A.) gibbosa somaliensis 2'58.8'N, 47"06.1'E (E. Nilsson-Cantell, 1929a:470 On spines of Cidaris sp.
(continued) Africa) ; 1289 meters
Balanomorpha :
Hexelasma antarcticum 75"02'S, 163"49'E (Terra Nova Bay) ; Antarctic only Borradaile, 1916 :132 Fossil, in glacier ;
(Borradaile, 1916) $9 meters type-locality
(=Bathy/asma)
66"32'S, 141"39'E ; 307 meters Bage, 1938:8 Living material
59"45'-59"4fj'S, 6S050'W ; Weisbord, 1965:1015 ; Shells only
1208-1592 meters 1967:51
59"56.8'S, 34O41.1'--34"32.7'W; Weisbord, 1965:1015; Shells only
1006-1153 meters 1967:51
67"48.5'S, 33"41'E (off Riiser-Larsen Utinomi, 1965:3 1 partially complete living
Pen.) ; 790 meters specimen and shells
67O51.55, 33" 13.5'E (off Riiser-Larsen Utinomi, 1965 :3 Shells only
Pen.) ; 630-680 meters
63"29.7'S, 3Zo02.7'E (off Riiser-Larsen Utinomi, 1965 :3 Shells only
Pen.) ; 590 meters
68O09.75, 34"34'E (cff Riiser-Larsen Utinomi, 1965 :3 Shells only
Pen.) ; 830 meters
68"10'S, 35"04'E (Gunnerus Bank) ; Utinomi, 1965:3 Shells only

620 meters
78"04'S, 167"501E; 100-200 ft above Speden, 1962 :746

sea level ; Pleistocene to Recent
78"28'S, 165"50fE; 3-10 ft above Speden, 19623746
sea level
H. corolliforme (Hoek, 1883) 5Z004'S, 71°22'E (off Kerguelen Hoek, 1883 :I55 Type-locality ; on pebbles
Islands) ; 280 meters and spines of Cidaris sp.
63"17'20"S, 59O48'15"W (South Nilsson-Cantell, 1930t):252
Shetland Islands) ; 200 meters
Off South Shetland Islands; Zevina, 1968:94
28-200 meters
Off Scott Island; 500-900 meters Zevina, 1965:94
Off Kerguelen Islands; 141 meters Zevina, 1968:94
63"32'S, 98"18'E; 420 meters Zevina, 1968:94
tBalanus (Austrobalanus) Kerguelen Islands ; frssil (750 ft South America; Recent Fletcher, 1938 :115
Vol. 14
flosculus var. sordidus above sea level; ? Pleistccene)

Darwin, 1854
North of
tBalanus sp. Cockburn Island, Graham Land, Hennig, 1911:10

64"20'S, 56"50'W ; fossil, on scallop
( ? Miocene)
Cornnula diadema South Georgia (approx. 55"S, 37"W) ; Nilsson-Cantell, 1921 :371 On ,Megaptera boops
(LinnB, 1767) on whales
South Georgia (approx. 55"S, 37"W) ; Probably cosmopolitan Nilsson-Cantell, 1930b:256 On humpback whale
on whales
South Shetland Islands (approx. Nilsson-Cantell, 1930b:256 On humpback whale
62"S, 58"W) ; on whales
57"201S, 32"W (South Georgia) ; Nilsson-Cantell, 1939:237 On humpback whale
on whales
C. reginae Darwin, 1854 Approx. 78'20fS, 17lo00'W, Ross Sea Cosmopolitan Nilsson-Cantell, 1926 :15 On blue whale
South Georgia (approx. 55"S, 37"W) ; Probab1)- cosmopolitan Nilsson-Cantell, 1930h:256 On blue whale
on whales
South Georgia (approx. 55"S, 37"W) ; Nilsson-Cantell, 1939:238 On blue whale
on whales
Xenobalanus globicipitis South Shetland Islands (approx. Calman, 1920:165 On Balaenoptera physalus
Steenstrup, 1851 62"S, 58"W) ; on whales
14"45'N, 18"34'W Nilsson-Cantell, 1930h:258 On whale

Deception Island, South Shetland Cosmopolitan; on cetaceans IYilsson-Cantell, 1930373258 On fin whale
Islands; on whale
+ Extinct or fossil forms.

Vol. 14
APPENDIX 2. Summary ol Stations Yielding Specimens Covered in Thii Siudy

-- -
Sta. Lat. Long. Datc Depth ( m e ~ e r s ) Gear
- - - - -- -- - -
L'L~~~~~L-USAKP
18 April 3, 1962 Blalie
43 June 12, 1962 Menzics
63 June 21, 1962 Mcnzies
112 July 20, 1962 Mcnzics
117 .luly 25, 1962 Engine room
strainer
July 28, 1962 IKMW T
Sept. 16, 1962 Mcnzics
Sept. 23, 1962 40' O ~ t e r
Sept. 23, 1962 l'eterscn grab
Sept. 27, 1962 4,0' Otter
Oct. 10, 1962 Kock dredge
Oct. 20, 1962 10' Blake
Oct. 26, 1962 Rock dredge
Dec. 2, 1962 40' Otter
Dee. 3, 1962 40' Otter
Uec. 3, 1962 Menzies
Dee. 4', 1962 Rock dredge
Dee. 20, 1962 Men~ies
Uec. 20, 1962 IICMWT
Dec. 20-21, 1962 Rock dredge
Dec. 21, 1962 5' Blake
Dec. 31, 1962 5' Ulalce
Jan. 1, 1963 5' Blake
Jan. 5, 1963 5' Blake
Jan. 7, 1963 5' Blake
Jan. 9, 1963 5' Blake
Feb. 15, 1963 Menzies
March 14, 1963 Menzirs
Rlarch 14, 1963 5' Blalte
Aug. 23, 1963 10' Blake
Oct. 25, 1963 5' Blake
Oct. 28-29, 1963 hlerlzies
Jan. 26, 1964 5' Blake
Jan. 28, 1964 5' Blalie
Feh. 10-11, 1964 5' Blake
Feh. 14, 1964 10' Blake
Marc11 13, 1964 5' Blake
March 13, 1964 10' Blake
March 14, 1964 10' Blake
Marc11 15, 1964 10' Blake
April 2, 1964 10' Blalce
April 8, 1964 5' Blake
April 15, I964 5' Blake
April 15-16, 1964 5' Blrike
April 17, 1964 5' Blakc
April 17, 1964 5' Ixlakc
June 17, 1964 5' Blaltc
June 23-24, 1964 5' lllalic
June 25, 1964 3 m IKhlWT
June 28-29, 1964 3 m IKMWT
June 30, 1964 3 m JKMWT
Ang. 4, 1969 IKMWT
Aug. 5, 1964 TKMWT
Aug. 6-7, 1964 TKMWT
Ang. 8, 1%4 IKMWT

APPENDICES 193
Std. Ldt.
--
1201 56" 14's Aug. 8, 1961. IKRlWT

1%03 55"56'S Aug. 10, 1964 IKMWT
1206 58" 14's Aug. 11, 1964 IKMWT
1220 6O052'S Aug. 15, 1964 IKRIWT
1224 6Z0O6'S Aug. 16, 1964 IKMWT
1235 59"52'S Aug. 20, 196% IKMWT
1270 57"37'S Sept. 1, 1964 IKMWT
1428 5lo06'S Feb. 10 20, 1965 3 m IKRIWT
1429 49" 15's Feb. 21, 196.3 40' Ottrr
1430 49"19'S l'cb. 22, 1965 40' Ottcr
14431 45"37'5 Feb. 23, 1965 40' Ottcr
SOSC 6 52"IO'S Rlarch 21, 1965 lloek drcdgc
A1batroc~-IJSBCF
2042 39"33'N July 30, 1883 Deep sea trawl
2196 39"35'N Aug. 6, 1884 Large bcam trawl
2221 39"05'30"N Sept. 6, 1884 Large hcarn trawl
2222 39"03'15"N Sept. 6, 1884 Large beam trawl
2662 29"24'30"N May 4, 1886 Large beam trawl
2663 29"39'N May 4, 1886 Large beam trawl
2711 38"59'N Sept. 6, 1886 Large bcam trawl
2731 36"4S1N Oct. 25, 1886 Large beam trawl
2741 37"44'N Sept. 17, 1887 1,argc bcam trawl
2839 33"08'N May 8, 1888 1,argc bcam trawI
2980 33 "49'54"N Feb. 12, 1889 1,argc bcam trawl
3487 5Z010'N July 13, 1893 1,argc beam trawl
3697 Off Manazuru- May 5, 1900 8' Tanner l ~ c a mtrawl
Misaki, Hon-
shu, Japan
3741 35 "N May 17, 1900 8' Tanner heam trawl
4353 15 mi off Point March 14, 1904 8' Tanner Lean1 trawl
Lorna, San Die-
go, California
4382 Off Isla Corona- March 18, 1904. 10' Blake trawl
do Nortc, San
Dicgo, Cali-
fornia
4397 33"10'15"N April 1, 1904 8' Tanner heam trawl
4<778 52"12'N June 5, 1906 12 ' Tanner beam trawl
5068 35"03'N Oct. 15, 1906 12 ' Tanner beam trawl
Umitaka-Mar u-JARE
4 67"48.5'S Feh. 7, 1957
6 67"51.5'S Feb. 7, 1957
S6ya-JARE
4 68"29.7'S Jan. 31, 1958
6 68"lO.O'S Jan. 27, 1958
7 68O09.75 Jan. 29, 1958
New Zcaland Geological Survey

695 (approx.1 1887(?)
36"45'S
GS7505 (dpprox.) Dee. 26, 1958 3-10' above sea level
78"28'S
GS7506 (approx.) June 6, 1958 610' above sea level
78"28'S
GS7512 (approx.) Dee. 22, 1958 10&200' above sea
78"4'S level

19'4 ANTARCT~C CIRRIPEDIA
- -
Sta. Lat. Long. Date Depth (meters) Gear

-- - -
Sarsia-MBAUK
P9jl 48"33'N 1O0O1'W May 3, 1957 1170-1244 4' rock dredgc
P9/2 48"34'N 1O000'W May 3, 1957 1042 1225 4' rock dredgc
P1/62 48"39.5'N 9"42.5'W 494-567
Enstz~ard-DUMBL
7617 33"58.7'N 75 "42.0'W June 21, 1967 2280
7552 33 "39.5'N 75"41.0'W June 15, 1967 3010
Lndenuor~r-NZARP
A-4-56 74-30's 119"40'W Jan. 15, 1959 238-301 Crab
A-463 72"20'S 174"50'E J a n . 21, 1959 465-4458 Grab
A-465 72"5S1S 175"301E Jan. 22, 1959 399
A-625 75"OO'S 163"58'42"E Feb. 5, 1961 460
Chain Cruise 50--WHO1
81 34"41'N 66"28'W July 2, 1965 5042 Epibenthic trawl
E1>ihcnthic trawl
Naga Expedition-SIO
60-212 15"40'N 109"22.9'E Feb. 27, 1960 110-198 10' benthic Blake
'I'ui-AU
011 30°45'S 173"51tE July 6, 1962 538-676
Jpn-111 Expedition-SIO
51 34"54.3'N 139"43.4'E July 17, 1961 549 Pebble dredge
Challenger
150 52"4'S 71°22'E Feb. 2, 1874 274
Terra Nova
- 75"s 164"E 30' above sea levcl
1)iscouery-BANZARE
301 60" 48's 71 "24'E Dec. 27, 1929 456
Siboga Expedition
253 S048.2'S 13Zo13'E
An~boina-Danish Expctlitiotr 10 Kci Islands
58 5"29'S 132"37'E May 12, 1922

APPENDICES
APPENDIX 3. Eltanin Cirripedia Listed A(:cording to Station
Sta. 18 Arco~cal~ellum vitreum (Hoek, 1883) Sta. 969 Scalpellun~gibberum Aurivillius, 1892
A r c o ~ c a l ~ e l l umichelottianum
m (Seguenza, 1876) Balanus laeuis BruguiBre, 1789
Arcoscalpellum sp., cf. A. sinuatum (Pilsbry, Sta. 981 Scalpellum gibberum Aurivillius, 1892
1907) Sta. 991 Gymnoscalpellum tarasovi, sp. nov.
Arcoscalpellum sp. Sta. 993 Cryptophialus tomlinsoni, sp. nov.
Sta. 43 Neoscalpellum eltaninae, sp. nov. Australscalpellum schizmatoplacinum, sp. nov.
Sta. 63 Arcoscalpellum hirsutum (Hoek, 1883) Sta. 995 Gymnoscalpellum tarasovi, sp. nov.
Sta. 112 Arcoscalpellum acicularum, sp. nov. Sta. 997 Gymnoscalpellum tarasovi, sp. nov.
Sta. 117 Lepas jascicularis Ellis and Solander, 1786 Sta. 1003 Australscalpellum schizmatoplacinum, sp. nov.
Conchoderma virgatum (Spengler, 1790) Arcoscalpellum multicostatum, sp. nov.
Sta. 122 Lepas anatifera LinnB, 1758 Bathylasma corolliforme (Hoek, 1883)
Sta. 216 Tetrachaelasma southwardi, sp. nov. Sta. 1054 Bathylasma corolliforme (Hoek, 1883)
Sta. 217 Scalpellum gibberum Aurivillius, 1892 Sta. 1067 Verruca gibbosa Hoek, 1883
Sta. 219 Balanus laevis Bruguikre, 1789 Bathylasnza corolliforme (Hoek, 1883)
Sta. 222 Scalpellum gibberum Aurivillius, 1892 Sta. 1084 Scalpellum vanhoffeni Gruvel, 1907
Sta. 255 Tetrachaelasma southwardi, sp. nov. Arcoscalpellum weltneri (Gruvel, 1907)
Sta. 268 Arcoscalpellum formosum (Hoek, 1507) Litoscalpellum fissicarinatum, sp. nov.
Sta. 289 Arcoscalpellum africanum ( Hoek, 1883) Sta. 1088 Bathylasma corolliforme (Hoek, 1883)
Sta. 337 Scalpellum gibberum Aurivillius, 1892 Sta. 1089 Cryptophialus tomlinsoni, sp. nov.
Sta. 339 Scalpellum gibberum Aurivillius, 1892 Sta. 1150 Arcoscalpellum latusculum, sp. nov.
Sta. 340 Arcoscalpellum parallelogramma (Hoek, 1883) Neoscalpellum schizoplacinum, sp. nov.
Sta. 343 Balanus laevis Bruguikre, 1789 Sta. 1161 Lepas australis Darwin, 1851
Sta. 372 Arcoscalpellum imbricotectum, sp. nov. Sta. 1162 Lepas australis Darwin, 1851
Sta. 375 Lepas australis Darwin, 1851 Sta. 1167 Lepas australis Darwin, 1851
Sta. 376 Tetrachaelasma southwardi, sp. nov. Sta. 1170 Lepas australis Darwin, 1851
Sta. 377 Scalpellum gibberum Aurivillius, 1892 Sta. 1185 Lepas australis Darwin, 1851
Sta. 410 Australscalpellum schizmatoplacinum, sp. nov. Sta. 1188 Lepas australis Darwin, 1851
Arcoscalpellum angulare (Nilsson-Cantell, 1930) Sta. 1196 Lepas australis Darwin, 1851
Arcoscalpellum liberum (Nilsson-Cantell, 1930) Sta. 1200 Lepas australis Darwin, 1851
Litoscalpellum simplex, sp. nov. Sta. 1201 Lepas australis Darwin, 1851
Sta. 413 Gymnoscalpellum tarasovi, sp. nov. Sta. 1203 Lepas australis Darwin, 1851
Sta. 426 Arcoscalpellum liberum (Nilsson-Cantell, 1930) Sta. 1206 Lepas australis Darwin, 1851
Sta. 430 Gymnoscalpellum tarasovi, sp. nov. Sta. 1220 Lepas australis Darwin, 1851
Sta. 439 A r c o ~ c a l ~ e l l u angulare
m (Nilsson-Cantell, 1930) Sta. 1224 Lepas australis Darwin, 1851
Ar~oscal~ellum liberum (Nilsson-Cantell, 1930) Sta. 1235 Lepas australis Darwin, 1851
Litoscalpellum discoveryi (Gruvel, 1907) Sta. 1270 Lepas australis Darwin, 1851
Sta. 480 Arcoscalpellum formosum (Hoek, 1907) Sta. 1428 Lepas australis Darwin, 1851
Sta. 557 Brochia bulata, sp. nov. Sta. 1429 Smilium acutum (Hoek, 1883)
Arcoscalpellum parallelogramma (Hoek, 1883) Arcoscalpellum buccinum, sp. nov.
Sta. 558 Arcoscalpellum parallelogramma (Hoek, 1883) Sta. 1430 Arcoscalpellum buccinum, sp. nov.
Sta. 671 Arcoscalpellum multicostatum, sp. nov. Sta. 1431 Balanus vestitus Darwin, 1854
Sta. 791 Arcoscalpellum vitreum (Hoek, 1883) Balanus decorus Darwin, 1854
Sta. 803 Arcoscalpellum darwinii (Hoek, 1883) SOSC Tetrachaelasma southwardi, sp. nov.
Sta. 945 Arcoscalpellum darwinii (Hoek, 1883) Sta. 6
Sta. 948 Litoscalpellum walleni, sp. nov.

ANTAHCTIC CIRRIPEDIA
APPENDlX 4.. Ellanin Cirripedia Listed According to Depth
Species Station
Conchorlerma vzrgatum (Spenglcr, 1790)

Lcprzs jascicularis Ellis and Solander, 1786
Surface Lepas anatifera Linn6, 1758
Surface Lepas australis Darwin, 1851
Surface Lepas ar~stralisDarwin, 1851
Surfacc Lepas australis Darwin, 1851
Surface Lepas australis Dai win, 1851
4049 Scalpellum gibberr~mAurivillius, 1892
79-80 Scalpellum gibherum Aurivillius, 1892
92 Scalpellum gibberum Aurivillius, 1892
106--110 Scalpellum gibberun~Aurivillius, 1892
115 Balanus laeuis BruguiZre, 1789
119 Bala~zuslaeuis Brugui&re, 1789
128-165 Arco~calpellumangzrlarr (Nilsson-Cantell, 1930)
Litoscalpellum discoveryi (Gruvcl, 1907)
Arcoscalpellum liberum (Nilsson-Cantell, 1930)
Balanus vestitus Darwin, 1854
Balanzts drcorr~sDarwin, 1854
Arcoscalpellr~m buccinum, 31). nov.
Australscalpellurn schiznanloplocinrcm, sp. nov.
Arcoscelpellum multsco.\lutu~n, sp. nov.
Bathylasma corolliforrrie (Hoek, 1883)
Australsca/pellum schizn~atoplatinurn, sp. nov.
Arcoscalpcll~~m a1zg7~10r~ 1930)
(Nils5011CLLIICCII,
Arcoscalpellum laberum (N~lsson-Cantell,1930)
Litoscalpellum simplex, sp. nov.
Arcoscalpellum multicostaturn, sp. nov.
Scalpellum gibberum Aurivillius, 1892
Ralanus laeuis BruguiZre, 1789
Scalpellum vanhoffrni Gruvrl, 1907
Arcoscalpellum weltneri (Crovel, 1907)
Litoscalpellum fissicarinaturn, sp. nov.
Cryptophialus tomlinsoni, sp. nov.
Australscalpellum schizmatoplacinunz, sp. nov.
Smilium acutum (Hock, 1883)
Arcoscalpellum buccinum, sp. nov.
Scalpellum gibberum A~~rii~illius, 1892
Arcoscalpellum pamllelogramma (Hock, 1883)
Bathylasma corolliformr (Hock, 1883)
Cryptophialus tomlinsoni, sp. nov.
Arcoscalpellum parallelogran~naa (Hock, 1883)
Gymnoscalpellum tarasovi, sp. nov.

APPENDICES
AI'I'ENDIX 4. (Continued)
Depth (meters) Station
Arcosculprllurn libr,rum (Nilsson-Cantrll, 1930)

Rathylasma corolliforme ( Hoek, 1883)
Broci~ia bulnta, sp. nov.
Arcosculprllun~ parallelograrrzma (Hoek, 1883)
Tetrachuelasma southwardi, sp. nov.
Verrucu gihbosa Hoek, 1883
Ucrthylusma corolliforrne (Hock, 1883)
Gymn~o.sca1~1ellurntaraso~ii,sp. nov.
Trtracharla.smu southwardi, sp. nov.
T~trcrcharlnsn~a southwardi, sp. nov.
Arcoscalprllnm Izirscctun~ (Hork, 1883)
Scnlpellum gibhrrrtnz Aurivillius, 1892
Arcoscul~~rllurn imbricotr~cturn,sp. nov.
Tetrachaelasrna sor~thwardi,sp. nov.
Gymnoscalpellum tarasoui, sp. nov.
Arcoscalpellum forn~osum (Hoeli, 1907)
A rcoscalpel/um africanun~ (Hock, 1883)
Arroscalprllum formosurr~ ( IIock, 1907)
Arcoscalpellurn. vitreum (Hoek, 1883)
Arcoscalpellum michelotticrnun~(Scgucnza, 1876)
Arco.scalpellr~m sp., cf. A. sinl~cctum (Pilsbry, 1907)
Arcoscalpellum aciczdarum, sp. nov.
Arcoscalpellum darwinii (Hoek, 1883)
Arcoscalpellum darwinii (Hoek, 1883)
Litoscalpellum walleni, sp. nov.
Arcoscalpellum uitreum (Hoek, 1883)
Arcoscalpellr~m latusculum, sp. nov.
Neoscalpellun~schizoplacinum, sp. nov.
AVc~oscalpellun~rltnninac, sp. nov.
- - -

APPENDIX 5. Bathymetric Distribution for Sampling Devices Used during Eltanin Expeditions Covered in This Report
Under each type of gear the three columns list, from left to right, number of attempts at depth given, total hours on
bottom, and number of species recovered. Parentheses enclose data from south of the Antarctic Convergence, extracted from
the unparenthesized totals for each depth interval. Species counts given in totals, with depth or within a depth interval, are
accumulative and therefore do not indicate number of species present (cf. Table 2).
Depth
(meters) 5' Blake 10' Blake Menzies Rock Dredge Peterseu 41)' Otter Totals
0499 17 5.41 12 20 8.30 6 5 3.08 0 14 12.78 8 94 42.02 23

( 8 1.95 10) ( 6 2.41 5) ( 2 2.00 0) ( 4 1.51 0) ( 32 15.02 15)
18 14.06 2 7 3.81 1 6 3.30 3 2 2.66 1 53 28.99 7
( 12 5.96 1) ( 3 2.08 1) ( 2 1.16 0) ( 1 2.36 0) ( 25 14.66 2)
8 5.15 5 2 2.50 1 7 5.96 1 1 1.25 0 30 19.92 8
( 7 4.65 4) ( 1 0.50 1 ) ( 3 2.41 0) ( 0 0 01) ( 19 12.62 6)
8 6.11 1 3 3.00 0 1 1.06) 2 0 0 0 16 11.19 4
( 6 4.45 0) ( 0 0 0) ( 0 0 0) ( 0 0 0) ( 8 4.45 0)
7 6.25 0 2 3.00 1 5 7.13 0 0 0 0 22 20.91 1
( 6 5.75 0) ( 0 0 0) ( 2 1.68 0) ( 0 0 0) ( 13 10.04 0)
14 14.83 1 3 1.78 1 12 11.03 1 0 0 0 37 33.30 4
( 14 14.83 1) ( 1 1.001) (11 9.53 1 ) ( 0 0 0) ( 31 28.02 4)
15 15.30 0 3 2.93 0 24 24.48 0 0 0 0 43 44.57 0
( 15 15.30 0) ( 0 0 0) (17 15.01 0) 0 0 0) ( 37 32.17 0)
11 12.45 1 4 4.16 0 31 41.05 0 0 0 0 54 64.02 1
( 10 10.61 1 ) ( 1 2.000) (26 32.98 0) ( 0 0 0) ( 43 50.42 1 )
21 19.55 5 7 8.60 0 16 19.75 2 0 0 0 52 52.48 7
( 14 13.08 5) ( 1 1.13 0) (12 13.15 2) ( 0 0 0) ( 32 31.19 7)
30 33.80 0 5 2.00 0 14 10.45 0 0 0 0 55 48.601 0
( 22 25.48 0) ( 41.58 0) (10 7.86 0) ( 0 0 0) ( 39 36.67 0)
5 6.00 0 1 0.31 0 5 6.00 1 0 0 0 13 13.26 1
( 5 6.00 0) ( 1 0.31 0) ( 4 4.00 0) ( 0 0 0) ( 12 11.26 0)
2 2.00 0 0 0 0 6 9.9.3 0 0 0 0 8 11.93 0
( 2 2.00 0) ( 0 0 0) ( 2 2.10 0 ) ( 0 0 0) ( 4 4.10 0)
0 0 0 0 0 0 0 0 0 0 0 0
( 0 0 0) ( 0 0 0) ( 0 0 0) ( 0 0 0)
1 1.58 0 0 0 0 0 0 0 0 0 0
( 1 1.58 0) ( 0 0 0) ( 0 0 0) ( 0 0 0)
0 0 0 0 0 0 I 1.00 a 0 0 0
( 0 0 0) ( 0 0 0) ( 1 1.00 0, ( 0 0 0)
0 0 0 0 0 0 1 1.001 0 0 0 0
( 0 0 0) ( 0 0 0) ( 1 1.00 0) 0 0 0)
Totals 157 142.49 27 57 40.39 I0 134 145.16 10 17 16.69 9
(122 111.69 22) (18 11.01 8) (93 93.88 3) ( 5 3.87 0)

REFERENCES
Annandale, N. 1925. Report on a collection of Cirripedia (barnacles)
1905. Malaysian barnacles in the Indian Rluseum, with a from South African waters. Fish. Mar. Biol. Sur-
list of the Indian Pedunculata. Mem. Asiatic Soc. vey, Union of South Africa, Rept. 4, no. 6, pp. 1-5.
Bengal, vol. 1, no. 5, pp. 73-84, pl. viii. Batham, E. J., and J. T. Tomlinson
1906. Natural history notes from the R.I.M.S. "Investiga-
1965. On Cryptophialus melampygos Berndt, a small bor-
tor", Capt. I. H. Heming, R.N., commanding. Series
ing barnacle of the order Acrothoracica abundant
111, no. 12, Preliminary report on the Indian stalked
in some New Zealand molluscs. Trans. Roy. Soc.
barnacles. Ann. Mag. Nat. Hist., Ser. 7, vol. 17,
New Zealand (Zoology), 1-01. 7, no. 9, pp. 141-154,
pp. 389-400.
23 figs., 1 pl.
1908. Crustacea (Entomostraca). Illus. Zool. R.I.M.S.
"Investigator", pt. 11, pls. 111-V. Belloc, G.
1909a. Description of a barnacle of the genus Scalpellum 1959. Catalogue des types de cirrhipPdes du MusCe
from Malayasia. Rec. Indian Mus., vol. 3, no. 3, Oc6anographique de Monaco. Bull. 1'Inst. OcCanogr.
pp. 267-270. Monaco, vol. 56, no. 1157, pp. 1-7.
1909b. An account of the Indian Cirripedia Pedunculata.
Mem. Indian Mus., vol. 2, no. 2, pp. 61-137, 2 pls. Benharn, W. B.
1910a. Description of a new species of Scalpellum from 1903. On some remains of a gigantic fossil cirripede from
the Andaman Sea. Rec. Indian Mus., vol. 5, no. the Tertiary rocks of New Zealand. Geol. Mag.,
2, pp. 115-116. vol. 10, no. 3, pp. 110-119, pls. IX and X.
1910b. The Indian barnacles of the subgenus Smilium, with
remarks on the classification of the genus Scalpellurn. Berndt, W.
Rec. Indian Mus., vol. 5, pp. 145-155, 2 figs. 1907. Uber das System der Acrothoracica. Arch. Naturg.
1910c. Notes on Cimpedia Pedunculata in the collection Jahrg., vcl. 73, no. 1, pp. 287-289.
of the University of Copenhagen. K~benhavn Nat.
Eorradaile, L. A.
Medd., pp. 211-218, pl. iii.
1911. Some barnacles of the genus Scalpellurn from Irish 1916. Crustacea. Part 111. Cirripedia. British Antarctic
seas. Ann. Mag. Nat. Hist., Ser. 8, vol. 7, pp. ("Terra No\aW) Expedition, 1910. Nat. Hist. Rept.,
588-590. Zool., vol. 3, no. 4, pp. 127-126, figs. 1-7.
1913. The Indian barnacles of the subgenus Scalpellum.
Boschma, H.
Rec. Indian Mus., vol. 9, no. 4, pp. 227-236.
1916. Barnacles from deep-sea telegraph cables in the 1930. Brrarosaccus callosus, a new genus and new species
Malay Archipelago. J. Straits Branch Roy. Asiatic of a rhizocephalan parasite of Lithodes agassizii
Soc., no. 74, pp. 281-302, pl. IV-VI. Smith. Proc. U.S. Natl. Mus., vol. 76, art. 7, pp. 1-8,
8 figs.
Aurivillius, C. W. S. 1962. Rhizocephala. Discovery Rept., vol. 33, pp. 55-92,
1892. Neue Cirripedien aus dem Atlantischen, Indischen text-figs. 1-24, pl. I.
und Stillen Ocean. K. Vet.-Akad. Forhandl. Stock-
holm, no. 3, pp. 123-134. Broch, H.
1894. Studien iiber Cirripeden. K. Svenska Vet.-Akad. 1912. Die P l a t t e n e n t ~ i c k e l u n bei
~ Scalpellum strlmii M.
Hand]., vol. 26, no. 7, pp. 5-107, pls. 1-9. Sars. K1. Norske Vidensk. Selsk. Skr., no. 4, Trond-
1898. Cirrhipi.des nouveaux provenant des campagnes hjem, pp. 1-14, 7 figs.
scientifiques de S.A.S. le Prince de Monaco. Bull. 1922. Papers from Dr. Th. Mortensen's Pacific Expedi-
Soc. Zool. France, vol. 23, pp. 189-198. tion 1914-1916. X. Studies on Pacific cirri~eds.
Vidensk. Medd. Dansk Naturh. Foren., vol. 73, pp.
Bage, F. 215-358, 77 figs.
1938. Cirripedia. Australasian Antarctic Expedition 1911- 1924. Cirri~edia Thoracica von Norwegen und dem Nor-
1914, Sci. Rept., Ser. C, Zoology and Botany, vol. wegischen Nordmeere. Eine systematische nnd
2, pt. 7, pp. 5-14, 1 fig., pls. V-VIII. biologisch-tiergeographische studie. Vidensk. Skr.
I. Mat.-Naturv. Kl., no. 17, pp. 5-121, figs. 1-35,
Barnard, K. H. pls. 1-11].
1924. Contributions to the Crustacean fauna of South 1927. Studies on Moroccan cirripeds (Atlantic coast). Bull.
Africa. 7. Cirripedia. Ann. South African Mus., Soc. Sci. Nat. Maroc, vol. 7, pp. 11-38, figs. 1-10, pls.
vol. XX, pt. 1, pp. 1-103, pl. I. I-VI.
99

1931. Papers from Dr. Th. Mortensen's Pacific Expedition 1851b. A monograph on the subclass Cirripedia, with fig-
1914-1916. LVI. Indomalayan Cirripedia. Vidensk. ures of all species. The Lepadidae; or, peduncu-
Medd. Dansk Natur. Foren., vol. 91, 146 pp., 41 figs. lated cirripedes. Kay Soc., London, pp. v-xi; 1-400,
1917. Cirripedes from Indochinese shallow-waters. Av- pls. I-X.
handl. Norslce Vidensk.-Akad., Oslo, 1, Mat.-Naturv. 1854. A monograph on the subclass Cirripedia with fig-
Kl., no. 7, pp. 3-32, figs. 1-8. ures of all the species. The Balanidae, the Ver-
1953. Cirripedia Thoracica. The Danish Ingolf-Expedi- rucidae, etc. Ray Soc., London, pp. vii-viii; 1-684,
tion, Copenhagen, vol. 3, no. 14, pp. 1-17, 12 figs. pls. I-XXX.
1959. Cirripedia. Thoracica, Family: Lepadidae. Con.
Internatl. L'Exp. Mer, Zooplankton Sheet 83, figs. Davadie, C.
1-6. 1963. Etude des balanes d'Europe et d'Afrique. SystB-
1961. Benthonic problems in antarctic and arctic waters. matiqne et structure des balanes fossiles d'Europe
Norske Vidensk.-Akad., Oslo, no. 38, pp. 5-32, 6 et d'Afrique. Ed. Centre Natl. Rech. Sci., pp. 3-146,
figs., 1 map. 55 pls.
Calman, W. T. Dayton, P. K., G. A. Robilliard, and A. L. DeVries

1918. On ballnacles of the genus Scalpellum from deep- 1969. Anchor ice formation in McMurdo Sound, Ant-
sea telegraph cables. Ann. Mag. Nat. Hist., Ser. 9, arctica and its biological effects. Science, vol. 163,
vol. 1, pp. 96-124, figs. 1-7. pp. 273-274.
1920. A whale-barnacle of the genus Xenobalanus from
antarctic seas. Ann. Mag. Nat. Hist., Ser. 9, vol. Dedlessandri, G.
6, pp. 165-166. 1895. Contribuzione a110 studio dei cirripedi fossili d'Ita-
lia. Bol. Soc. Geol. Ital. Roma, vol. XIII, pp. 234-
Caras, R. A.
314, pls. iii-v.
1962. Antarctica; Land of frozen time. Chilton Books, 1897. La pietra da Cantoni di Rosignano e di Vignale
Philadelphia, 209 pp. (Basso Monferrato) . Studi s~ratigraficie paleonto-
Chapman, F. logici. Mem. Soc. Ital. Sci. Nat. Milano, vol. VI,
1914. Australasian fossils; a student's manual of pale- pp. 1-98> pls. i and ii.
ontology. Melbourne and London, 341 pp., 150 figs., 19%. Studi monografici sui cirripedi fossili d'Italia.
1 map. Palaeontogr. Italica, vol. XII, pp. 207-324, pls. xiii-
xviii.
Chilton, C.
1911. Scientific results of the New Zealand Government de Graaf, F.
Trawling Expedition 1907. Crustacea. Rec. Canter- 1952. Some notes on the genus Lepas LinnB, 1767. Beau-
bury Mus., vol. 1, no. 3, pp. 285-312, pl. LVIII. fortia, no. 14, pp. 1-6, figs. 1-2.
Clarke, E. Dell, R. K.
1905. The fossils of the Waitemata and Papakura series. 1952. Marine biology, In The Antarctic today a mid-
Trans. New Zealand Inst., Wellington, vol. 38, pp. century survey. Frank A. Simpson, ed., New
413421, pl. xxxii. Zealand Antarctic Soc., Wellington, 389 pp., 1 fig.,
Clarke, R. 45 pls.
1966. The stalked barnacle Conchoderma, ectoparasitic on
Djakonov, A. M.
whales. Norsk Hvalfangst-Tidende, no. 8, pp. 153-
168, 7 figs., 5 tables. 1914. Ascothorax ophioctenis n.g. und n. sp.-ein nener
Endoparasit aus der Gruppe der Ascothoracidae.
Codez, J. Vorl. Mitt.-Trav. Soc. Nat. C. R. SBances Petrograd,
1957. Etudes de cirripkdes acrothoraciques fossiles: Di- vol. 485, no. 1, pp. 158-164; 1 fig.
plome d'Btudes supQieures, Poitiers (not seen).
Ekman, S.
Codez, J., and R. de Saint-Seine
1953. Zoogeography of the sea. Sidgwick and Jackson,
1957. RBvision des cirriphdes acrothoraciques fossiles. Ltd., London, 417 pp.
Bull. Soc. Geol. France, Ser. 6, vol. 7, pp. 699-719,
pls. 37-39, tables 1-3. Fletcher, H. 0.
Cornwall, I. E. 1938. Marine Tertiary fossils and a dcscription of a Re-
1962. The identification of barnacles with further figures cent Mytilus from Kerguelen Island. B.A.N.Z. Ant-
and notes. Canadian J. Zool., vol. 401, pp. 621-629, arctic Res. Exped. 1929-1931, Ser. A, vol. 2, Geol.,
pl. 1-11. pt. 6, pp. 101-116, fig. 15, pls. 1 0 ~ 1 2 .
Darwin, C. Foster, B. A.
1851a. A monograph of the fossil Lepadidae, or, pednn- 1967. A guide to the littoral balanomorph barnacles of
culated cirripedes of Great Britain. Palaeontogr. Soc., New Zealand. Tuatara, vol. 15, no. 2, pp. 75-86,
Lo~ndon,88 pp., tables 1-5. figs. 1-13.

KEFEKENCES 201
Cow, A. J., W. F. Wccks, G. Ilendrickson, a ~ r dK. Rowland 1556. Suyrplcmcnt to a list of the Crustacea of Tasmania.
1965. Nrw light on tlie motlc of uplift of the fish and Iicc. ()urm Victoria Rlus., Tasmania, New Ser.,
fossiliferous moraines of the McMurdo ice shelf. rlo. 5. pp. 1 8 .
J . Clariol., vol. 5, no. 42, pp. 813-828.
Hapgood, C. H.
1962. Ancient knowledge of Anicrica and Antarctica.
182.5. A synopsis of tlrc genera of cirripedes arrangctl Itlraca, vol. 8, no. 26, pp. 479 488, figs. 1-5.
in natural familirs, with a description of some new
species. Ann. Philos., New Ser., vol. 10, no. 2, p p IIcctor, J.
97-1 07. 1888. Specimens of a large fo5iil stalkctl cir~ipcdc. Trans.
Roy. Soc. New Zealand In\t., Wellington, vol. 2D,
Gruvrl, A. p. 440.
1900a. Sur urre cs1)i.c.e nouvelle du genre Sccrlpellum provc-
nant d(: la collection tlu MusCum tl'tiistoirc Na-
turellr de Paris. Bull. Mus. d'liist. Nat., I'aris, 1951. Antarctic parasitic copepotls and an ascothoracitl
vol. 6, pp. 188-189. cirriped from brittle-slars. Vidensk. Rlcdtl. J ) B I I S ~ .
190011. Sur c~uelquesesp6c.e~nouvcllcs du genre S c a l p e l l ~ ~ m Natur. Foren. Copenhagen, vol. 113, pp. 171-1W,
provenant de la campagne du "Talisman." Rull. 7 figs., 2 pls.
RPus. d'llist. Nat., Paris, vol. 6, pp. 189-194.
1902a. Cirrhipides. Expkditions scientifiques du "Travail- llenrrig, A.
leur" et du "Talisman," Paris, pp. 1-178, 18 figs., 1911. LC corrglomCrat pleis~ockne B Pecten de I'ile Cock-
7 pls. burn. Schwcdischen Sudpolar Expedition 190-1903,
19021,. Revision dcs cirrhipkles appartcnant B la collccticin vol. 111, Geologie und Palaetrntologie, pp. 1-72, 4
du MusCum d'F[istoirc Naturelle. Arch. Mus. d'llist. figs., 5 pls.
Nat., Ser. 4., vol. 4, pp. 215-312, pls. XI-XIV.
19013. Revision tles cirrhipkdes appartenant B la collection Hmry, U. P., and P. A. McLaughlin
tlu MusCum tl'llistoire Naturelle. Nouv. Arch. Mus. 1967. A revision of the subgenus Solirlobal(znz~s Hoek
d7Hist. Nat., Scr. 4, vol. 5, pp. 95-1701. (Cirripedia Thoracica) including a description of
1905. Mcmographie des cirrhipkdes ou thEcostracGs. IlIas- a new species with complemental males. Crusta-
son et Cic., Cditcurs, Paris (reprinted 1965 by A. ceana, vol. 12, pt. l , pp. 4'3-58, figs. 1-3, pl. 1.
A ~ h e rand Co., Amsterdam, Netherlands, will1 -472
Hiro, F
pp., 427 figs.)
1906. CirrliipPdes du 1)iscovcry. Cull. Mus. d'Hist. Nat., 1932. Report on the Japanese species of tlic genus Ca-
Paris, vol. 12, pp. 270-273. lantica (Cirripedia) . Annot. Zool. Japan, vol. 13,
1907a. Crustacca. VI. Cirrhipkdes. Natl. Antarrtir Expe- no. 5, pp. 67-482, 5 figs., pl. 30.
dition. Nat. Hist., vol. 111. Zoology and Rctany, 1933. Report on the Cirripedia collected by the surveying
Brit. Rlus. (Nat. Hist.), London, pp. 1-4, 1 pl. ships of the Imperial Fisheries Experimental Station
1907b. Nutc prklin~inaire sur les cirrhipides p$donculi~s on tlie continental shelf bordering Japan. Rec.
rccurillis par 17cxpGdition antarctiriue allcmantl': Oceanogr. Works, Japan, vol. 5, no. 1, pp. 11-84,
tiu "Causs." Rull. Soc. Zool. France, vol. 32, pp. 22 figs., 3 pls.
157-163. 19%. 'l'he fauna of Aklceshi Bay. 11. Cirripedia. J. Fac.
19W. Die Cirripcdien der Deutschen Sudpolar-Expedition Sci., Hokkaido Impcrial Univ., Ser. 6, Zoology,
1901-1903. Dcutsche Siidpolar-Expedition 1901- vol. 4, no. 4, pp. 213-229, pl. X.
19013, vol. 11, Zoologie, no. 3, pp. 195-229, pls. 23-26, 1937. Studies on cirripedian fauna of Japan. 11. Cir-
19lla. ExpCdition antarctique fran~aise du "Pourquoi- ripcds found in the vicinity of the Seto Marine
Pas?", dirigde par M. le Dr. J. B. Charcot (1908- Biological Laboratory. hlem. Coll. Sci., Kyoto Im-
1910). Liste des cirrhipkdes. Bull. Mus. d'llist. pcrial Univ., Ser. B, vol. 12, no. 3, art. 17, pp.
Nat. Paris, vol. 17, p. 292. 385-478, 43 figs.
1911b. Sur dcux espkces nouvelles de cirrhipGdes apparte- 1939. Studies on the cirripedian fauna of Japan. IV.
nant i la collection du Muskum. Bull. Mus. d'Hist. Cirripeds of Formosa (Taiwan), with some gco-
Nat. Paris, no. 5, pp. 29CL291. graphical and ecological remarks on the littoral
1912. Note prkliminaire sur les cirrhipkdes rccueillis pen- forms. Mem. Coll. Sci., Kyoto Imperial IJniv., Ser.
dant les campagnes de S.A.S. le Prince de Monaco. B, vol. 15, no. 2, art. 8, pp. 245-284, 16 figs.
Bull. 1'Inst. Oceanogr., Paris, no. 241, pp. 1-7.
1920. Cirrhipkdes provenant des campagnes scientifiques IIoek, P. P. C.
de S.A.S. le Prince de Monaco, 1885-1931. Results 1883. Report on the Cirripedia collccted by H.M.S. Chal-
Campagnes Sci. Albert 1"' no. 53, pp. 3-88, pls. 1-7. lenger during the years 1873-1876. Report on the
scientific results of the voyage of H.M.S. Challenger
Guiler, Eric R. during the years 1873-1876, Zoology, vol. 8, pt. 25,
1952. A list of the Crustacea of Tasmania. Rcc. Queen pp. 1-169, 3 figs., pls. 1-13.
Victoria Mus., Tasmania, vol. 3, no. 3, pp. 15-44. 1884. lbid., Part 28, Anatomical Part, pp. 1 4 7 , pls. I-VI.

1904. An interesting case of reversion. Proc. Sect. Sci., 1942b. The body appendages of Balanus decorus. Trans.
K. Akad. Wetensch, Amsterdam, vol. 7, pt. 1, pp. Proc. Roy. Soc. New Zealand, vol. 72, pp. 1-5, 1 pl.
90-94, figs. 1-5.
1907a. Cirripedia. Rtsultats voyage du S.Y. Belgica, 1897-
1899 (Expedition Antarctique Belge). Rapp. Sci. 1929. A report on some cirripeds collected by the S.S.
Zool., Anvers, pp. 3-9, 4 figs. "Albatross" in the Eastern Pacific during 1891 and
1907b. The Cirripedia of the Siboga-Expedition. A. Cirri- 190%. Bull. Mus. Comp. Zool., Harvard, vol. 69,
pedia Peduuculata. Siboga-Expeditie, vol. 31, pp. no. 15, pp. 527-538, pls. 1-3.
1-127, pls. 1-10.
Menzies, Robert
1913. The Cirripedia of the Siboga-Expedition. B. Cirri-
pedia Sessilia. Siboga-Expeditie, vol. 31, pp. 129-275, 1963. General results of biological investigations on the
PIS. 11-27. deep-sea fauna made on the U.S.N.S. Eltanin
1914. Cirripedia from the "Michael Sars" North Atlantic (U.S.A.R.P.) during Cruise 3 between Panama and
Deep-Sea Expedition 1910. The "Michael Sars" Valparaiso, Chile in 1962. Int. Revue Ges. Hydro-
North Atlantic Deep-Sea Exped., vol. 3, pp. 3-5. biol., vol. 48, no. 2, pp. 185-2001, figs. 1-14.
Hutton, F. W. Moore, L. B.
1879. List of the New Zealand Cirripedia in the Otago 19U. Some intertidal sessile barnacles of New Zealand.
Museum. Trans. Proc. Roy. Soc. New Zealand Inst. Trans. Proc. Roy. Soc. New Zealand, vol. 73, no. 4,
pp. 315-334, pls. 4 U 7 .
vol. 11, art. XXXIV, pp. 328-30.
Murray, J.
Jeffreys, J. C.
1895. A summary of the scientific results obtained at the
1878. On the Mollusca procured during the "Lightning"
sounding, dredging, and trawling stations of 1I.M.S.
and "Porcupine" expeditions, 186&18701. Part I.
Challenger. Rept. H. M . S. Challenger (1872-18761,
Proc. Zool. Sac., pp. 393416.
Summary, vol. 1, pt. 1, p. 509 (only).
Jennings, L. S.
Newman, W. A.
1915. Pedunculate Cirripedia of New Zealand and neigh- 1961a. On the nature of the basis in certain species of the
bouring islands. Trans. Proc. New Zealand Inst., Hembeli section of Chthamalus (Cirripedia, Thora-
Wellington, vol. 47, art. XXX, pp. 285-293, figs. 1-3. cica). Crustaceana, vol. 2, pt. 2, pp. 142-150, 2 figs.
1918. Revision of the Cirripedia of New Zealand. Trans. 1961b. On certain littoral species of Octolasmis (Cirripedia,
Proc. New Zealand Iust., Wellington, vol. 50, art. Thoracica) symbiotic with decapod Crustacea from
111, pp. 56--63.
Australia, Hawaii, and Japan. Veliger, vol. 4, no. 2,
pp. 9%107, pls. 21-23.
Kolosvary, G.
1967. A new genus of Chthamalidae (Cirripedia, Balano-
1950. Descrip~ions of three new fossil Tertiary barnacles m o r ~ h a ) from the Red Sea and Indian Ocean. J.
from Hungary. FGldtani Kozlony, vol. 8, pts. 7-9,
Zool., London, vol. 153, pp. 42-35 6 figs.
pp. 1-6, figs. 1-4. - Cirripedia. I n Reef and shore fauna of Hawaii,
L. G. Eldredge ed., B. P. Bishop Mus. Spec. Publ.,
Kriiger, P.
Honolulu, Hawaii (in press).
1911. Zur Cirripedenfauna Ostasiens. Zool. Ang., Leipzig,
vol. 38, pp. 4 5 9 4 4 . Newman, W. A., V. A. Zullo, and S. A. Wainwright
1940. Cirripedia. I n Bronns Klassen und Ordnungen des 1967. A critique on recent concepts of growth in Balano-
Tierreichs, Leipzig, bd. 5, abt. I , buch 3, teil 3, pp. morpha (Cirripedia, Thoracica). Crustaceana, vol. 12,
1-560, figs. 1-391. pt. 2, pp. 167-178, 1 fig., 3 pls.
Lacaze-Duthiers, H. Nicol, D.
1880. Histoire de la Laura geraridae, Type nouveau de 1966. Descriptions, ecology, and geographic distribution of
crustacC parasite. Arch. Zool. Exper. Gen., Paris, some antarctic pelecypods. Bull. Am. Paleontol.,
vol. 8, pp. 337-381, figs. 1-7. vol. 51, no. 231, pp. 5-102.
Leach, W. E. Nilsson-Cantell, C. A.
1825. A tabular view of the genera composing the Class 1921. Cirripeden-studien. Zur Kenntnis der Biologie, Ana-
Cirripedes, with descriptions of the species of Otion, tomie und Systematik dieser Grnppe. Zool. Bidrag,
Cineras, and Clyptra. Zool. J., vol. 2, pt. 6, art. Uppsala, vol. 7, pp. 75-394, figs. 1-89, pls. 1-3.
XXIII, pp. 208-215. 1926. Antarktische und subantarktische Cirripedieu. Gesam-
melt van S. Vallin 1923-1924. Arkiv Zool., Stockholm,
Linzey, J. T. vol. 18A, no. 27, pp. 1-16, figs. 1-5.
1942a. The balanomorph barnacles of the Kermadec Islands. 1927. Some barnacles in the British Museum (Nat. Hist.).
Trans. Proc. Roy. Soc. New Zealand, vol. 71, pp. Proc. Zool. Soc., London, pp. 743-7W, figs. 1-19,
279-281. pl. I.

REFERENCES 203
1928. Studics on cirripeds in the British hluheun~ (Nat. Pilsbry, H. A,, and A. A. Olsson
Hist.). Ann. Mag. Nat. Hist., Scr. 10, vol. 2. no. 7, 1951. Trrtiary and (:rt:taccous C:irripetlia from north%-est-
pp. 1- 39, figs. 1-16. ern South America. l'roc. Acatl. Nat. Sci., Philatlcl-
1929a. Cirripedien tles genus Verrnca tler 1)cutsehen Tief- phia, vol. 1013, pp. 197-210.
see-Expedition auf ti en^ 1)ampler "Valdivia" 1898-
1899. Zool. Jahrb. (Abt. Syst., Okol. Geogr. Ticre), Richards, H. G., and J. R. Craig
vol. 58, pp. 459480, figs. 1-7. 1963. 11. Pleistocene mcrllusks frorrr the Continental Shell
192911. Cirripeds from the Juan F e r n a n d c ~Islands. IIL The off Argentina. In I'leistocene sedin~entation arid
natural history of Juan Fcrnantl~z ant1 Easter fauna of the Argcn~tinc Shelf, (:. Fray and M.
Jsland, C:. Skottsbcrg, ctl., vol. 111, Zoology, pp. 483- Ewing, eds., Acad. Nat. Sci. Philadelphia, Proc., vol.
492, figs. 1 3. 115, no. 6, pp. 127-147, pls. 1-3.
1930a. Cirripedien von der Stcwart-lnsel und von Siidgeor-
gien. Srrickcnht~rgia~~a, vol. 12, pp. 210-213, I fig. Rodda, P. U., and W. L. Fisher
1930b. Thoracic cirripeds i:ollrcted in 1925 -1927. Discov~.ry 1962. Upper Paleozoic acrothoracic: harr~aclrs from Tcxas.
Rcpt., vol. 2, pp. 223-200, figs. 1-12, pl. I. Texas J. Sci., vol. 14, no. 4, pp. 460-479, l i p . 1 13.
1931. Cirripetls from the Indian Ocean and Malay Archi-
pelago in the British Museum (Nat. Hist.), 1,ontlon. Koss, A.
Arkiv Zool., Stoc~kholrri,vol. 23A, no. 18, pp. 1 12,
1962. Results of the Puritan-American Museum of Natural
figs. 1-3.
History Expedition to Western Mexico. 15. The
1938. Cirripedes from the Indian Ocean ill tlrc collection
littoral balanomorph Cirripedia. Am. Mus. Novitates,
of the Indian Museum, Calcutta. Alem. Indian Mus.,
no. 2084 pp. 1-4, figs. 1-24.
vol. 13, pt. I , pp. 1-81, 28 figs, 3 pls.
1965. Acrothoracicarl barrracle burrows from the Florida
1939. Thoracic: cirripcds collected in 1925-1936. Discovery
Miocene. Crustaceans, vol. 9, pt. 3, pp. 316318,
Rept., vol. 18, pp. 223-238, figs. 1-5.
pl. 2.3.
1955. Cirripcdia. Rept. Swedish Deep-Sea Expedition, vol.
1968. Bredin-Archbold Smithsonian liiolirgical Survey of
2, Zoology, no. 17, pp. 215-2#20.
Dominica. 8. The intertidal halanomorph Cirripcdia.
1957. Thoracic cirripcds from Chile. Rept. Lunds Univ.
Proc. U. S. Natl. Mus., vol. 125, no. 3663, pp. 1-22,
Chile Expedition, 1918-191.9, Lunds Univ. Ars~krift.,
3 figs.
N.F., Avd. 2, vol. 53, no. 9, pp. 3-25.
1970. Studies on the Tetraclitidae (Cirripedia: Thoracica) :
A proposed new genus for the austral species T r t m -
Pantin, C. F. A. (ed.) clita purpurascens brevisrl~tum. Trans. San Dicgo
1960. A discussion on the biology of tlle southern cold Soc. Nat. Hist., vol. 16, no. 1, pp. 1-12, figs. 1-5.
temperate zone. Roy. Soc., Proc. (Lorrtlon), Scr.
B., vol. 152, pp. 429 677. Ross, A., and W. A. Newman
1967. Eocene Balanidae of Florida, including a new genus
Park, J. and species with a unique plan of "turtle-barnacle"
1910. The geology of New Zealand, Christchur~h, New organization. Am. Mus. Novitdtes, no. 2288, pp. 1-21,
Zealand, pp. xx, 1-488, 1 map, text-illus., 1 pl. figs. 1-7.
1969. Cirripcdia, In Distribution of selected groups of
Pilsbry, 11. A marine invertebrates i n waters south of 3 5 " s lati-
tude, J. W. Hcdgpeth, ed., Antarctic Folio Series,
1897. Description of a remarkable Japanese cirripedr.- Amer. Geogr. Soc., vol. 11, pp. 1b3.2, p1. 17.
Scalpellun~sexcornulurn n. sp. Am. Nutur., vol. 31,
pp. 72%7214, fig. 1. Saint-Seine, R. de
1907a. Cirripetlia from the Pacific coast o [ North America.
1954. Existence de cirripades acrothoraciques des le Lias:
Bull. U. S. Bur. Fislicries, vol. 26, no. 617, pp. 193-
Zapfella pattei nov. gen., nov. sp. Soc. Geol. France,
204, figs. 1-4, pls. 6-11.
Bull. and C. K , Ser. 6, vol. 4, pp. 447-451, pls. 18-20.
1907b. The barnacles (Clirripcdia) contained in the collcc-
tions of the IT. S. National Museum. Bull. U. S. Nail.
Sandved, K. G.
Mus., no. 60, 122 pp., figs. 1-36, pls. 1-11.
1 9 0 7 ~ Notes
. on some Pacific cirripetles. l'roc. Acad. Nat. 1966. USNS Eltanin: Four years of research. Antarctic
Sci., Philadelphia, vol. 59, pp. 360-362, pl. 29. J., vol. 1, no. 4, pp. 164-174.
1908. On thC classification of scalpelliform barnacles. Proc.
Acad. Nat. Sci., I'hiladelphia, vol. 60, pp. 1CH-111, Seguenza, G.
fig. 1. 1875-1876. Richcrche paleontologiche intorno ai Cirripedi
1911. Barnacles oI J a l ~ a nand Bering Sea. Bull. I T . S. Bur. Tcrziarii dclla provincia di Messina. Con appendice
Fishcries, vol. 29, pp. 61-81., pls. VlII-XVlI. intorno ai Cirripedi viventi nel Rlediterraneo e sui
1916. The sessile bari1ac1r.s (Cirripetlia) contained in the fossili Terziarii dell'Italia meridionale. pt. i, Balanidi
collections of tlir I T . S. National RTn.scum; including e Verrucidi, 1873; pt. ii, Lepadidi, 1876. Atti Accad.
a monograph of the An~crican spccics. U. S. Natl. l'ontaniana, Napoli, vol. X, pp. 265-4231, pis. A, B,
Mus. l'rull. 93, pp. v-xi, 1-366, figs. 1-99, pls. 1-76. I-X.

Southward, A. J., and E. C. Southward Tokara Islands. IX. Cirripedia. Publ. Seto. Mar.
1958. On the occurrence and behaviour of two little-known Biol. Lab., vol. 4, no. 1, art. 2, pp. 17-26, figs. 1-2.
barnacles, Hexelasma hirsutum and Verruca recta, 1958. Studies on the cirripedian fauna of Japan. VII.
from the continental slope. J. Mar. Biol. Assoc. U.K., Cirripeds from Sagami Bay. Publ. Seto. Mar. Biol.
vol. 37, pp. 633-647, figs. 1-5, pl. 1. Lab., vol. 6, no. 3, pp. 281-311, figs. 1-10.
1965. A giant antarctic barnacle Hexelasma antarcticum
Speden, I. G. Borradaile (Cirripedia, Thoracica) . J.A.R.E. 1956-
1962. Fossiliferous Quaternary marine deposits in the Mc- 1962 Sci. Rept., Ser. E, no. 25, pp. 1-15, figs. 1-4,
Murdo Sound region, Antarctica. New Zealand J. pls. 1-2.
Geol. Geophys., vol. 5, no. 5, pp. 746-777, figs. 1-16.
Stebbing, Rev. T. R. R. 196k On Parascothorax synagogoides gen. n., sp. n.

1910. No. 6, General Catalogue of South African Crustacea, parasitizing Ophiz~ra quadrispina Clark and some
(Part V of S. A. Crustacea, for the Marine In- remarks on geographical distribution of Ascothora-
vestigations in South Africa). Ann. South African cida (Entomostraca) . Akad. Nauk Okeanol. Trud.
Mus., vol. vi, pp. 563-576 (Cirripedia only). Inst. Okeanol., vol. 69, pp. 271-284, figs. 1-10 (in
Russian with English summary).
Stephensen, K.
Weisbord, N. E.
1935. Two crustaceans ( a cirriped and a copepod)
endoparasitic in ophiurids. Danish Ingolf-Expedi- 1965. Two new localities for the barnacle Hexelasma
tion, vol. 3, pt. 12, pp. 3-13, figs. 1-10. antarcticum Borradaile. J. Paleontol., vol. 39, no. 5,
pp. 1015-10'16.
Stubbings, H. G. 1967. The barnacle Hexelasma antarcticum Borradaile-its
1936. Cirripedia. Sci. Repts., The John Murray Expedition, description, distribution, and geologic significance.
1933-1934, Brit. Mns. (Nat. Hist.), vol. 4, no. 1, Crustaceana, vol. 13, pt. 1, pp. 51-60, fig. 1-9.
70 pp., 30 figs.
Weltner, W.
1961. Cirripedia Thoracica from tropical West Africa. At-
lantide Rept. No. 6, Sci. Res. Danish Expedition 189il. Zwei neue Cirripedien aus dem Indischen Ocean
Coasts Tropical West Africa, 1945-1946. pp. 141, (Scalpellum M e g a l a s m a ) . Gesellschaft Natnrf.
11 figs. Freunde, no. 2, Berlin, pp. 80-86, fig. 6.
1967. The cirriped fauna of tropical West Africa. Bull. 1895. Die Cirripedien von Patagonien, Chile und Juan
Brit. Mus. (Nat. Hist.) Zool., vol. 15, no. 6, pp. 229- Fernandez. Arch. Naturg., vol. 1, no. 2, pp. 288-292.
319, 28 figs., pl. 1. 1896. South American Cirripedia, J. Roy. Microscop. Soc.,
Lsndon, Ser. 2, p. 63.
Tarasov, N. I., and G. B. Zevina 1897. Verzeichnis der bisher beschriebenen recenten Cirri-
1957. Usonogie raki (Cirripedia Thoracica) morei SSSR. pedienarten. Arch. Naturg. Jahrg., vol. 1, no. 3, pp.
Fauna SSSR, Zool. Inst. Akad. Nauk, SSSR. New 227-280.
Ser., no. 69, vol. 6, pt. 1, pp. 1-268, figs. 1-106, 1898. Cirripedien. In Ergebnisse der Hamburger magal-
pls. 1-3. haensischen Sammelreise. L. Friederichsen and Co.,
Hamburg, lief IV, pp. 3-15.
Taylor, B. J. 1899. Ergebnisse einer Reise nach dem Pacific (Schau-
1965. Aptian cirripedes from Alexander Island. Brit. insland 1896-1897). Zool. Jahrb., vol. 12, pp. 441447.
Antarctic Surv. Bull., vol. 7, pp. 37-42, figs. 1-3. 1900. Die Cirripedien der Arktis. Fauna Arctica, vol. 1,
pp. 289-312, 1 fig., pl. VIII.
Thomson, W. 1922. Cirripedia der Deutschen Tiefsee-Expedition. Wiss.
1873. Notes from the Challenger. VI. Nature, pp. 347-349, Ergeb. Deutschen Tiefsee-Exped. Valdivia 1898-
figs. 1, 2. 1899, vol. 23, no. 2, pp. 61-112, 3 figs., pls. 11-IV.
Tomliuson, J. T. Withers, T. H.
1955. The morphology of an acrothoraclcdn barnacle, 1913. Some Miocene cirripedes of the genera Hexelasma
Trypelrsa lateralis. J. Morphol., vol. 96, no. I , pp. and Scalpellum from New Zealand. Proc. Zool. Sac.,
97-121, pls. 1-4. London, no. 56, pp. 840-854, figs. 139-14, pls. 85-86.
1960. Cryptophialus coronatus, a new specles of acrothora- 1914. A new cirripede from the Cenomanian Chalk Marl
cican barnacle from Dakar. Bull. l'Inst. Franc. of Cambridge. Geol. Mag. London (dec. vi), vol. I,
d'dfrique Noire, Ser. A, vol. 22, no. 2, pp. 4Q2-410, 3 pp. 4,94497, figs. 1-6.
figs. 1924. The fossil cirripedes of New Zealand. New Zealand
1969. The burrowing barnacles (Cirripedia: Order Tho- Geol. Surv. Branch, Paleontol. B d . , no. 101, pp. 1-47,
racica). U. S. Natl. Mus. Bull. 296, pp. 1-162, figs. figs. 1-8, pls. I-VIII, 2 maps.
145. 1926. A new cirripede from the Upper Cretaceous of West-
ern Australia. J. Roy. Soc. W. Australia, vol. 12,
Utinomi, H. no. 11, pp. 101-104, fig. 2, pl. xi.
1954. Invertebrate fauna of the intertidal zone of the 1928. Catalogue of fossil Cirripedia in the Department of

REFERENCES 205
Geology. Vol. I. Triassic anti Jurassic, Brit. 34us. Isopoda Asellota. Galathea Rept., vol. 6, pp. 7-320,
(Nat. Hist.), London, pp. v-xi, 1-155, figs. 1-92, 184 figs., pls. I-XIX, 21 tables.
pls. 1-12.
1935. Catalogue of fossil Cirripedia in the Department of Yaldwyn, J. C.
Geology. Val. 11. Cretaceous. Brit. Mns. (Nat. Hist.), 1965. Antarctic and subantarctic decapod Crustacea. Mono-
London, pp. 1-534, 64, figs., pls. 1-50. graph. Biol., vol. 15, pp. 324-332.
1936. Miocene cirripedes (Scalpellurn and Tessarelnsrna,
gen. nov.) from India. Ann. Mag. Nat. Hist., Ser. Young, Maxwell W.
10, vol. 18, pp. 589-593, pl. XI. 1929. Marine fauna of the Chatham Islands. Trans. New
1917. Cirripedia, In Paliontologische und geologische Zealand Inst., vol. 60, pp. 136-166, ~ 1 s 16-17.
.
Ergebnisse der Reise von Kohl-Larsen (1928-29)
nach Siid-Georgien, 0.Wilckens, ed., Abhandl., Senc- Zevina, G. B.
kenberg. Naturf. Ges., Abh. 474, pp. 18-19 (Cirri- 1962. Barnacles of the genus Scalpellr~m Leach collected
pedia only), figs. 1-2, pl. 1. by the Soviet Antarctic Expedition on the "Oh'" i n
1951. Cretaceous and Eocene peduncles of the cirripede the Antarctic and Subantarctic. Biol. Rept. Soviet
EuscaZpellum. Bull. Brit. Mus. (Nat. Hist.), Geol., Antarctic Expedition 1955-1958, vol. 2, pp. 252-253.
vol. 1, no. 5, pp. 145L162, figs. 1-6, pls. 11-14. (Translated by: Israel Program for Scientific Trans-
1953. Catalogue of fossil Cirripedia in the Department of lations, Jerusalem, 1966.)
Geology. Vol. 111. Tertiary. Brit. Mus. (Nat. Hist.), 1968. New and rare species of barnacles (Cirripedia
London, pp. vii-xv; 1-396, figs. 1-105, pls. 1-6,.2'. Thoracica) from the Antarctic. Acad. Sci. USSR
Biol. Inst. Invest. Fauna Seas, VI (XIV), Res. Biol.
Wolff, Torben Analyses Soviet Antarctic Expedition 1955-1958, no.
1962. The systemaLics and biology of bathyal and abyssal 4, pp. 85-96, 6 figs.

GLOSSARY OF TERMS
Most of the terms given here are i1lustratc.d in thc text-figures, especially 2, 3 , 69, and 70 on pages I G , 10, 1-1.1,
and 143, respectively.
Acanthopod: Cirri with setdc along lesser curvature larvae and ascothoracicans, and mantle of other
much reduc.ed; setac along greater curvature ar- orders; commonly with calcified plates in thor-
ranged in transverse l o w o f strong, sharp spines acicans.
below articulation (cf. ctenopod, lasiopod, hypo-
lasiopod) . Carina (pl. carinae) : Unpaired posterodorsal plate
possessing alae in balanomorphans (cf. rostrum,
Ala (pl. alae) : Lateral portion of plate in balano- compound rostrum).
morphans, over1appi.d by adjacent plate and dc-
limited by a marked change in the direction of Caudal appendages: A pair of terminal, multiarticu-
growth lines from nearly horizontal to more or late, or uniarticulate uniramous appendages (syn.
less vertical (cf. radius). caudal furca) .
Chela (pl. chelae) : Pincer-like first antenna in as-

Antenniform: Articles more or 1e.s radially sym-
cothoracicans.
metrical. round in cross section and movable
through 360° ; antenna-like.
Cirrus ( 1 . cirri) : Multiarticulate biramous thoracic
limb, extended through aperture in capturing food
Aperture: Opening into mantle cavity (cf. orifice)
(outer ramus: syn. exopocl, anterior ramus; in-
ner ramus: syn. endopod, posterior ramus; cf.
Arthrodial membrane: Cuticular membrane and un-
ctenopod, acanthopod, lasiopod, hypolasiopod) .
clerlyirig tissues forming articulation between
opercular valves arid sheath. Compound rostrum: Tripartite plate overlappins ad-
jacent plates, with or without radii, formed by
Arthropodal: Wiih reference to characteristics of
fusion of rostrum with rostrolaterals (cf. ros-
cirri and trophi in contrast to thosc of platcs.
trum).
Basidorsal point: Conical papilla located posterodor-
Comb collar: Retractable membrane supporting row
sally on hase of penis in balanids.
of uniform setae, located at superior angle of
aperture in acrothoracicans.
Body chamber: See mantle cavily.
Compartment: One of several rigid1 articulated
Branchia (pl. branchiae) : In balanomorphans, leaf- plates forming the wall in verrucomorphans and
like extensions of interior of mantle cavity, pre- balanomorphans (cf. paries) .
sumably respiratory.
Compound eye: One of pair of compound photo-
Bullate : See labrum. receptors commonly appearing in late nauplii
and cyprid larvae; lost in adults (cf. nauplius
Capitulum (pl. capitula) : In lepadomorphans, por- eye).
tion of carapace forming mantle cavity, com-
monly armored with calcareous plates. Complemental male: See male.
Carapace: Fold of integument extending from maxil- Ctenae: Minute comb-like scales, commonly on mouth
lary segment; forming bivalved shell of cyprid parts and appendages.

208 GLOSSARY
Ctenopod: Cirrus with paired setae along lesser cur- Lasiopod: Cirri with setae arranged in brushlike
vature arranged in longitudinal row (cf. acantho- groups at distal end of articles adjacent to articu-
pod, lasiopod, hypolasiopod) . lations (cf. ctenopod, acanthopod, hypolasiopod) .
Cyprid larva (pl. larvae) : Nonfeeding, weak-swim- Lateral bar: In acrothoracicans, particularly Crypto-
ming larval form found in all cirripeds, appear- phialidae, one of a pair of external chitinous
ing after the last riaupliar stage arid attaching thickenings running from opercular bar medially
before undergoing metamorphosis into adult form down each side of mantle sac.
(syn. cypris stage).
Latus (pl. latera) : In lepadomorphans, any of the
Dwarf male: See male.
paired plates forming part of shell, except the
scuta, terga and the unpaired carina, subcarina,
Filamentary appendages : Membranous processes on
rostrum, and subrostrum. ( a ) Carinal latus:
the body of many lepadomorphans and some
located on either side of proximal end of carina.
acrothoracicans; located on dorsal surface, pro-
(b) Inframedian latus: located below upper
soma, and especially at bases of cirri; presum-
latus and between carinal and rostra1 latera. (c)
ably respiratory.
Rostra1 latus: located on either side of rostrum.
( d ) Upper latus : centrally located between
Frena: See ovigerous frena.
inframedian latus, scutum, tergum, carina, and
carinolatus.
Frontolateral horn: One of a pair of tubular, cuti-
cular processes in the nauplius of all orders ex-
cept Ascothoracica; located at the anterolateral Longitudinal canal: Tube formed in compartment of
margins of the head. certain balanomorphans, between longitudinal
septa arid irlrlrr and outer laminae.
Gastric mill: Chitinous triturating apparatus in fore-
gut of Cryptophialidae (Acrothoracica) . Longitudinal seplum ipl. septa) : Wall disposed nor-
mal to inner and outer laminae, separating longi-
Hermaphrodite: Monoecious, sexes combined in one tudinal canals.
individual (cf. male. complemental) .
Male: Always retlucrtl in structure; commonly with-
Hypolasiopod: Cirri with cluster of 3 or more major out Ieeding organs; termed rolnplemental when
setae; at distal ends of articles adjacent to articu- occurring with hermaphrodite.
lations, other setae on articles pairetl (ctenopod).
Mandible: One o r thc thiid pair of head appendages;
Inner lamina (pl. laminae) : Inner shell layers of
hiramous in nauplius; reduced to a blade and
halanomorphan compartments. hen separated
palp in atlults (lost in rhizocephalans) ; second
from outer lamina by longitudinal septa.
behind mouth in nauplius, first in adults. Palp
attached in acrothoracicans hut transferred to
Interlaminate figure: Tn some Iralar~omorphans, sim-
margin of lal~rum in thoracirans.
ple or arborrscerit lines ~ r i r ~ r ~ ifrom
r ~ g outer
lamina across longitudinal :c.pta to inner lamina,
seen when wall is sectioned par,~llelto base. Mandibular palp: Scr mandible.
~ , ~labra) :b ~ ~ ~outgrowth
~ ~ i~ in~ front
~ .Mantle:
~ l Portion of carapacc forming fleshy part of
of mouth, and in naul~liusmore or 1e.s covering capitulum, conimonly armored u itli calcareous
it; in adult thoraricans. suppoiting mandil)ular plates in thora~itans.
palps; commonly bulla~c,,with rour~tlrdcrest, but
flattenrd and cleEt ill higher balanomorphans. M a n t l ~cavity : SIrat e occupied by appendage-bear-
ing portion of bod\, communicating with exterior
- -
Lamina: See inner lamina. by aperture.

Mouth cirri: First pair of cirri in acrothoracicans l'edirel: Basal portion of c i r ~ u s .formed of two seg-
much modilietl and closely associated with trophi nients (syn. pedicle, 1)rtiuncle. protopod).
~al .
(cf. ~ e ~ r r i i rcirri)
Peduricle: Stalk in lepadomorphans, supporting c:api-
Nauplius eye: Unpaired mediar~eyc found in nau- tulurn, attachvd to substratum and commonly
plius and cyprid larvae, and providing elernen~s armed with calcareous scales (also see pedicel).
for unpaired or paired photoreceptors irr certain
adults (syn. median eye).
Plate: T.oosely-, any calcareous part of a thoracican,
hut generally restricted to those plates not modi-
Nauplius (pl. riauplii) : Simplest larval iorrn, with
lied as opercular valves.
three pairs of appendages (first antennae, second
antennae, arid mandibles), commonly passing
Primordial plate: Chitinous plate or valve in lepado-
through six stages before metamorphosis into
morpharis and verrucomorphans, developing at
cyprid larva; sometimes completing development
incipient umhonrs of terga, scuta, and carina dur-
in egg, especially in dcep sea and high latitude
ing metamorphosis.
species.
Occludent margin: Margin of aperture, or of plates Radius (pl. radii) : Lateral part of compartment
or valves surrounding it. overlapping ala of adjacent plate when marked
off from that compartment by an abrupt change
Opercular valve: Any of the movable plates occlud- in direction of growth liries from horizontal to
ing the aperture in I~alanomorphans and verru- more or less vertical.
comorphans.
Rostrum (pl. rostra) : Unpaired anteroverltral plate
Opercular har: One of a pair of chitinous bars paral- in thoracicans, located between rostrolaterals and
leling aperture in acrothoracicans. opposite posterodorsal carina; provided with alae
in lower, but generally compound ifused with
Operculum : Terga, scuta, and associated membranes rostrolaterals and therefore generally provided
(balanomorphans), or movable tergum and scu- with radii) in higher balanomorphans (see com-
tum (verrucomorphans) , forming apparatus oc- pound rostrum).
cluding the aperture.
Sheath: More or less thickened upper internal part
Orifice: Opening in halanomorphan and verruco- of compartments in balanomorphans forming cy-
morphan wall occupied by operculum. lindrical ring from which the opercular parts
are suspended by the arthroclial membrane.
Outer lamina: See inner lamina.
Terminal cirri: Cirri, except first, located at pos-

Ovigerous frena (pl. frenae) : Fleshy ridge or flap
terior end of thorax in acrothoracicans (see
on interior of mantle, holding egg mass (oviger-
mouth cirri).
ous lamella) in certain lepadomorphans.
Palp : See mandible. Umbo (pl. umbones) : Apical portion of a valve

or plate; point from which concentric growth
Paries (pl. parietes) : A compartment, exclusive of lines emanate.
alae and/or radii, of sessile barnacles.
Valve: Any one of the plates forming the operculum
Parietal canal: See longitudinal canal. in sessile barnacles.
Parietal septum: See longitudinal septum. Welting: In balanomorphans, a distinct beading

along upper or superior margin of an ala, ini-
Parietal tube: See longitudinal canal. tially strengthening articulation at alar angle.

PLATES

PLATE I

PLATE -11

PLATE I11

PLATE IV
PLATE V

PLATE VI
PLATE VII

PLATE VIII
PLATE IX

PLATE X

PLATE XI
PLATE XI1

PLATE XI11
PLATE XIV

PLATE XV
PLATE XVI

PLATE XVII
PLATE XVIII

PLATE XIX
PLATE XX


PLATE XXIII

PLATE XXIV
PLATE XXV

PLATE XXVI
PLATE XXVII

PLATE XXVIII
PLATE XXIX

PLATE XXX
PLATE XXXI

PLATE XXXII

"Pi 5
*
PLATE XXXIII

PLATE XXXIV

PLATE XXXV
PLATE XXXVI

PLATE XXXVII
PLATE XXXVIII

PLATE XXXIX

PLATE XL

PLATE XLI
PLATE XLII

PLATE XLIII
PLATE XLIV

PLATE XLV
PLATE XLVI

PLATE XLVII
PLATE XLVIII

PLATE LEGENDS
Plate I A. Scalpellid harnacles (indicated hy long arrows) ant1 anerriones (one indicatcd hy upper short arrow), growing on
sten1 of whip coral (lowc-r short arrow), inhahiting apparently firm irregular bottom covered l ~ yfine sediments marked
by animal tracks in the vicinity of thr organisrris, autl scatterrtl rocks. 3493 meters, off South Shetland Islands (Ellanin
Cruise 6, Jan. 7, 1963, Sta. 41, Camera Sta. 41 ( 3 6 ) , 1;rame 10, 5&"93'30"'S, 59"08'W).
1J. Scalpellid barnacles (horizontal long arrows) growing on arborescent organisms inhahiting soft hottom marked hy
fecal casts (vertical short arrows), anirnal tracks and trails, and burrows indicated by mounds. An elasipod sea eucurl~ber
(lower horizontal short arrow) appears hrading towards clump of I,arnaclrs, and brittle starfish (upper short arrow)
seen holding arms abovc the substratum and prcsurnahly fccding in a gentle current. 4773 meters, off South Shetland
Islands (Ellanin Cruise 4, August 2, 1962, Caliirru Sta. 8, Frame 21, 61°41'S, 6l01O'W).
Plate I1 A. Numerous scalpellid 1,nrnaclcs (several groups indicated by long arrows), anemones (several indicated hy short
arrows directed downward), and galatheidcan anomurous crabs (several indicated by short arrows directed upward).
inhahiting nearly sediment-free volcanic rock. 2377 rnelers, off South Sandwich Islands.
B. Same situation and faunal complex as A. Note scalpellids growing along depressions in rocks much as do forms
living in high energy environments (examples indicated by oblique long arrows). Individual barnaclrs (horizontal long
arrows) appear leaning and extending cirral nets to the left, anti tentacles on the right side of the anemone (short
arrow directed downward) appear similarly oriented, suggesting a current progressing from left to right, carrying sedi-
ments and food with it. 2377 meters, off South Sandwich Islands (Eltanin Cruise 22, Feb. 17, 1966, Sta. 29, Camera Sta. 18,
Frame 4, 60°03'S, 29"58'W) .
Plate I11 Whale harnacles: Conchoderma auritum (LinuB) on Coronuln diadema (Linni.) from humphack whale, California,
X 1.3 (John Nordback photo, SIO) .
Plate IV Brachyzapfes elliptica gigantea Taylor: A, cast of hurrows in a belemnite rostrum, ~ 3 B,; casts of interior of burrow
viewed from right and "rostral" sides, X 5 ; C, as R, viewed from rostral and left sides, X 5 (rephotographed from Taylor,
1965, text-fig. 3 ) .
Plate V A-E, Lrpadidae; F, S(:alpellidar: A, Lepns (Lepas) austrnlis Darwin, X2.1 (Eltanin Sta. 1185) ; R, L. (L.) auslralis
Darwin, ~3 (Eltanin, Sta. 1162) ; C , L. (L.) (inatifera Linni., ~6 (Eltanin, Sta. 122) ; L), L. (Dosimal fascicularis Ellis
and Solander, X2.5 (Eltanin, Sta. 117) ; E, Concho~lerrr~a ziirgatum ( S ~ c n g l e r ) ,X 2 (Eltanin, Sta. 117) ; F, Smilium
acutum (Hoek) , X4.2 (Eltanin, Sta. 1420).
Plate VI A-E, Euscalpellum antarcticum Withers; F-H, Cretiscalpellunz aptiensis antarcticum Taylor: A, peduncle (part
of, broken longitudinally), outer vicw; B, inner view of A showing shape of plates; C, ~ e d u n c l e (part of), outer view;
I), peduncle (part of), top view; E, outer view of D (A-E, X 0 . 9 ) ; F, latex cast of natural mold showing scutum,
X6.2; G, latex cast showing internal and external impression of tcrga, X2.2; H, latex cast showing impression of
carinci, ahove renter, ~ 2 . 2(rephotographed from Withers, 1951, pl. 12, figs. 2, 3, and 4 ; Taylor, 1965, text-figs. 1 and 2 ) .
Plate VII %rugnzato/rpns georgiensis Withers: A, incomplete rapituluni, X4.2. Ttie left plate is a tergum; the middle and
right are scuta. The rostrum lies heneath and to the right of the right scutum. Both small plates lying over the tergum
and left scutum are lower latera (Senckcnberg. Mus. Cat. No. 328). B, many sc:uta, X7.4 (Senckenberg. Mus. Cat.
No. 404) (rephotographed from Withers, 1947, pl. 1, figs. 1, 2 ) .
Platc VIII Arcoscczlpellum: A, A. 1atu.sc.ulum sp. nov., holotype, X1.9 (Eltanin, Sta. 1150) : B, A. wc,ltnc,ri (Gruvel), X63 (El-
tanin, Sta. 1084) ; C, A. acicularunz sp. nov., holotypc, X2.7 (Eltanin, Sta. 112) ; D, A. africanum (Hock), ~ 5 . 5(Eltanin,
Sta. 289) ; E, A. uitreum (Hoek), adult, X2.8 (Eltanin, Sta. 18) ; F, A. uitreun~(Hoek), juvenile, X5.5 (Eltanin, Sta. 18) ;
C , A. jorrr~o.sum (IIock), ~ 5 . 5(Eltunin, Sta. 268) ; H, Arcoscnlprllunz sp., ~ 4 . 3(Eltanin, Stn. 18) ; I, A. huccinum sp.
nov., holotype, X2.4 (Eltanin, Sta. 1429) .

Plate IX Arcoscalpellunr: A. A. irnbricotec/urn sp. nov., holotype, ~ 5 . 6(Eltanin, Sla. 372) ; B, A. rrzichelottianum (Seguenza),
~ 5 . 7(Eltanin, Sta. 18) ; C, A. parallelogrornrna (Hock), ~ 2 . 3(Eltanin, Sta. 558) ; D, Arcoscalpellum sp., cf. A. sinuntum
(Pilsbry), ~ 7 . 5IEltanin, Stci. 18) ; E, A. darwinii ( H o c k ) , X0.7 (Eltanin, Sta. 945) ; 17, A. mr~ltico.statum sp. nov.,
holotype, ~5 iEltanin, Sta. 071) ; G , A. czngulcrre (Nilsson-Cantell), X8.3 (Eltanin, Sta. 410) ; H, A. liberum (Nilsson-
Cantell), X8.3 (Eltanin, Sta. 410).
Plate X Scalpellidae: A, Neo,sccrlpellurn schizop/acinurn sp. nov., holotype, ~2 (Eltanin, Sta. 1150) ; B, Neoscalpellum eltaninae
sp. nov., holotypc, ~ 2 . 1(Eltrrnin, Sra. 43) ; C, Litoscalpellum fissicarinulurr~gcn. et sp. nov., paratype, X2.1 (Eltanin,
Sta. 1084) ; D, Lito.scalpel1um simplex gcn. et sp. nov., hnlotype, ~2 (Eltcrnin, Sta. 410) ; E, Litoscalpellum walleni gen.
et s p nov., holotype, ~3 (Eltar~in,Sta. 948) ; F, Litoscolpellun~cliscoveryi (Cruvcl), X12 (Eltanin, Sta. 439).
Plate XI Gyrnnosc~l~ellr~nz
tarasovi grn. ct sp. nov.: A-C, paratypes, X3.7, X3, X2.3, respectively; D, holotype, ~2 (Eltanin,
Sta. 430).
Plate XI1 Scal~ellidae:A, Scalpellurr~gibberz~nz Aurivillius, ~ 1 . 1(Eltanin, Sta. 217) ; B, Brochia bulata gen. et sp. nov.,
holotype, ~ 9 . 7(Eltanir~,Sta. 557) ; C, Scalpellurn vunhoffrni Gruvel, X7.3 (Eltanin, Sta. 1084) ; W E , Australscalpellum
schiznzatoplacinum gen. ct sp. nov., paratypes, X11.2 (Eltanin, Sta. 410).
Plate XI11 Scalpellum gibbrrurrz Aurivillius (horizontal long arrow) on a stylasterinc hydrocoral, Errina sp.; Balanus laeuis
laevis Brugui$re (vertical long arrow) ; cctoproct hrycjzoan (upper short arrow) ; articulate brachiopod (middle short
arrow) ; and spiorhid polychacte wnrrn (lowrr slrort arrow) on a scallop, X1.5 (Eltanin, Sta. 969, approx. 250 meters: off
Tierra del Fuego) .
Plate XIV Verruca gibbosa Hoek: A, E, side views of cornpletc shell, X5.8; 13, apical view, X5.8; C, 11, external and internal
views, respcctively, of movable scutum and tergum, ~ 7 . 5(Eltunin, Sta. 1067).
Plate XV Buthvlasrna corolliiorn~c~(FIork) : A-C. entire specimen, attached to a rostral plate, ~ 3 . 3 ;A, B, viewed from
right side; C. picwed Irom above; I), E, spccimerr attdchcd to volcanic rock, X1.9; D, viewed from rostral e n d ; E,
viewed frorn above (NZARP Endeavour, Sta. A-463).
I'latc XVl Rathylasmn corollijornze (Hoek) : Exterior view of wall plates. ~ 2 . 7 ;A, rostrum; 11, right lateral; C, right
c.arinolatera1; D, carina (NZARP Endeavour, Sta. A-463).
Plate XVll Ra~hylasnrn corollijorrr~r t Hoek) : A-D, opercular valves, ~ 3 . 9 ;A, K, cxtrrit~r,lcft scutum and tergum, respcc-
tively; C, D, intcrior left tcrgum and scutuni, respectively; E-H, i n t ~ r i o rof wall plates, ~ 1 . 5 ;E, rostrum; F, right
lateral; G, right c:arinolatcral; H, carina (NZARP Endeavo:~r,Sta. A-463).
Plate XVIII Bathylccsma corollijorme (Hoek) : A, B, rxtcrior of lcft and right latcral, respectively, ~ 2 C,; D, extrrior ant1
interior of right sc,uturn, ~ 2 . 3 .Arrow indic,ates inconipletc boring 1,y a mollusc: ( J A R E Urr~itcika-Maru,Sta. 4 ) .
corollijormr ( H c ~ e k:) C:arin,r, X2.1; A, exterior; I$, sitic view; C, interior (JAKE Un~rtaka-'Wnru,
I'larc XIX Rath~lusrr~n
Sta. 4 ) .
Plate XX Bathylasmu corollrjormr (Hoek) : A, whole s p c ~ i m e nviewcd from right srdr, X2.1 (BANZARE Discovery, Sta. 30) ;
H , C, longitudinal thin section of shell; R, from Ross Sea, X4.8 (NZARP Endeavour, Sta. A-456) ; C, from "Taylor
Fornration," McMurdo Sound, X34 (Geol. Survey (N.Z.) GS-7505).
Plate XXI Bathylasma corolliformr (Hoek) : Longitudinal thin sections of shell; A, from Ross Sea, X36 (NZARP Endeavour,
Sta. A-625) ; B, from upper third of shell, "Scallop Hill Formation," McMurdo Sound, ~ 4 (Geol.9 Survey (N.Z.)
GS-7512).

PLATE LEGENDS 245
Plate XXII Bathylasma corolliforme (Hoek) : Longitudinal sections, same shell illustrated in pl. XXI B; A, from middle region,
X6O; B, from basal region, X 4 1 (Geol. Survey (N.Z.) GS-7512).
Plate XXIII Bathylasma hirsutum (Hoek) : A-D, opercular valves, ~ 2 A,; B, exterior, C, D, interior of right scutum and ter-
gum, respectively; E, F , whole specimen, x1.1; E, from right side; F, from carinal end; G, H, entire specimens; arrows in-
dicate antenniform rami of cirrus I11 (photographed alive by Alan Southward, Mar. Lab., Plymouth).
Plate XXIV Bathylasma hirsutum (Hoek) : A-C, carina; A, interior; E, exterior; C, left side, X 1.8 (MEAUK Sarsia, Sta.
P1/62) ; D, carina from left side, X l . 6 (MBAUK Sarsia, Sta. P 4 / 9 ) (photo 11y Alan Southward, Mar. Lab., Plymouth).
Upper arrow indicates "welting"; next arrow, line separating internal and external outer alar surfaces; next, internal
surface; lowest arrow, inferior alar rriargin.
Plate XXV Bathylasma aucklandicum (Hector) : A, longitudinal section, X44; B, external view of right lateral, ~ 3 . 3(Loc.
No. N381.524, Coll. No. GS-695) ; C, external view of plaster cast of lateral plate made by T. H. Withers (Brit. Mus.), X2.9
(photo by Alan Southward, Mar. Lab., Plymouth). Arrow on B indicating line separating welting from external alar
surface; on C indicating internal outer alar surface.
Plate XXVI Tetrachaelasma southwardi gen. et sp. nov., holotype: A-D, entire specimen; A, carinal, B, rostra1 and C, right
lateral views, X1.7; D, viewed from above, X1.3; E, F, exterior and G, H, interior views of left and right terga and scuta,
respectively, X2.6 (Eltanin-SOSC Sta. 6 ) .
Plate XXVII Tetrachaelasma southwardi gen. et sp. nov., holotype: Carina; A, interior, B, exterior and C, right oblique views,
X2.5 (Eltanin-SOSC Sta. 6 ) .
Plate XXVIII Tetrachaelasma southwardi gen. et sp. nov.: A, entire specimen viewed from above, X2.4; B-D, holotype;
B, internal view of rostrum; C, D, exterior view left lateral and carina, ~ 2 . 2(Eltanin-SOSC Sta. 6 ) .
Plate XXIX Tetrachaelasma southwardi gen. et sp. nov.: Dead shells viewed from exterior; A-E, carinae; F-H, left laterals;
ILK, right laterals; L, rostrum, x 1.3 (Eltanin, Sta. 376).
Plate XXX Tetrachaelasma southwardi gen. et sp. nov.: Carina; A, exterior; B, from right side; C, interior, ~ 1 . 7(Eltanin,
Sta. 225).
Plate XXXI Tetrachaelasma southwardi gen. et sp. nov.: Thin sections; A, transverse, X32; B, longitudinal (top at left), X 4 1
(Eltanin, Sta. 376).
Plate XXXII Transverse thin sections of parietes: A, B, Aaptolasma americanum (Pilsbry) ; C, D, Aaptolasma callistoderma
(Pilsbry) (A, C, X52; B, D, enlarged) (see pl. XLIII and text for explanation).
Plate XXXIII Transverse thin sections cf parietes: A, B, Aaptolasma leptoderma gen. et sp. nov. (A, ~ 8 4 B, ; enlarged) ;
C, D, Aaptolasma triderma gen. et sp. nov. (C, ~ 3 3 D, ; enlarged portion, laminae forced apart revealing calcareous
hridges) ; E, F, Aaptolasma brintoni gen. ct sp. nov. (E, ~ 5 5 F,
; enlarged).
Plate XXXIV Transverse thin sections of parietes: A, Balanus (Balanus) regalis Pilsbry, X28; B, Balanus (Balanusl pacificus
Pilsbry, ~ 3 3 C,
; D, Elminius simplex Darwin (C, X92; D, enlarged) ; E-G, Tetraclita squamosa rubescens Darwin
(E, X41; F, outer lamina, X78; G, radius, X133).
Plate XXXV Aaptolasma callistoderma (Pilshry) : A, E, rostrum; B, F, left lateral; C, G, left carinolateral; D, H, carina,
interior and exterior, respectively, ~ 4 I,; J, exterior right, K, L, interior left scuta and terga, respectively, of a larger
specimen, X3.7 (Albatross, Sta. 3741).

Plate XXXVI Aaptolasma americanum (Pilsbry) : A, B, interior left and C, D, exterior right scuta and terga, respectively,
X6.5 (Albatross, Sta. 2663, holotype, USNM Cat. No. 14559).
Plate XXXVII Aaptolasma americanum (Pilsbry) : A, transverse thin section of wall (upper arrow, chitin-filled longitudinal
tube; middle arrow, "interlaminate figure" marking position of tooth along basal margin of wall; lower arrow, secondary
layers of inner lamina), X42; B, chitinous filling between outer and inner laminae (upper horizontal arrow) and nodular
calcareous extensions embedded in chitin (lower arrow) (enlarged, from A) (Albatross, Sta. 2662; USNM Cat. No. 48093).
Plate XXXVIII Aaptolasma brintoni gen. et sp. nov., holot~pe:A, B, interior and C, D, exterior terga and scuta, respectively,
X8.1; E, entire specimen on articulate brachiopod, X4.1; F, I, interior rostrum, left lateral, left carinolateral and carina
(hole in carina grinding artifact), X4.2 (SIO Naga Expedition, Sta. 60-212).
Plate XXXIX Aaptolasma: A-E, A. brintoni gen. et sp. nov., holotype; A, B, exterior of left lateral and carinolateral, X10.4,
and C, D, rostrum, X7.2 and carina, X 8 ; E, transverse section of wall, X 4 l (SIO Naga Expedition, Sta. 60-212) ; F, G, A.
triderma gen. et sp. nov., holotype: F, entire shell from right side, X3.1; G, transverse section of wall, ~ 3 (SIO
4 Jyn I11
Expedition, Sta. 51). Upper arrow, inner lamina; middle arrow, chitinous filling; lower arrow, outer lamina.
Plate XL Aaptolasma leptoderma gen. et sp. nov., holotype: A-D, opercular valves; A, B, exterior and C, D, interior right
scutum and tergum, respectively, X17; E-J, wall plates; E, interior right lateral, ~ 6 . 7 ;F, exterior left lateral; G, H,
interior and exterior right carinolateral; I, interior carina; J, interior rostrum, ~ 7 . 5 . Arrows indicate, in descending
order: C, articular ridge, articular furrow, adductor muscle pit, lateral depressor muscle crests, occludent margin, rostra1
depressor muscle crests; G, welting, alar angle, boundary between alar and parietal portion of wall, interior alar margin,
overlapping margin without radii; H, sheath, depression receiving ala of adjacent plate; J, portion of calcareous basis
adhering to rostrum (Amboina Danish Expedition, Sta. 58).
Plate XLI Balanus (Austrobalanus) vestitus Darwin: A, wall, X5.8; B, C, exterior right and D, E, interior left scuta and
terga, respectively, x 8.9 (Eltanin, Sta. 1431).
Plate XLII Balanus (Bathybalanz~s) pentacrini Hoek, paratype: A, interior of rostrum, X25; B, wall from right side,
X17; C, D, exterior right and E, F, interior of left scuta and terga, respectively, ~ 2 (Siboga,
2 Sta. 253).
Plate XLIII Transverse thin sections of wall: A, Bathylasma corolliforme (Hoek), X35 (NZARP Endeavour, Sta. 463) ; B,
Balanus (Bathybalanus) pentacrini Hoek, X133 (upper arrow, "intralaminate figure" indicating position of basal tooth;
middle arrow, fibers; lower arrow, secondary internal calcification) (Siboga, Sta. 253) ; C, Aaptolasma callistoderma
(Pilsbry), X41 (short arrows pointing to left, "interlaminate figures"; short arrows pointing to right, chitinous filling of
tubes separating inner and outer lamina; long arrows, calcareous bridges between inner and outer lamina that were basal
teeth when formed; vertical short arrows, fibers) (Albatross, Sta. 3741) .
Plate XLIV Balanus (Balanus) laevis laevis Bruguihre: A, wall viewed from above (broad transverse ribs due to substrate
replication), X3.5; B, C, exterior left and D, E, interior right scuta and terga, respectively, X6.2 (Eltanin, Sta. 969).
Plate XLV Balanus (Megabalanus) deccrus Darwin: A, wall viewed from right side, ~ 2 B,; C, interior left and D, E, exterior
right scuta and terga, respectively (Eltanin, Sta. 1431) .
Plate XLVI Balanus spp.: A, cf. Balanus flosculus sordidus Darwin(?), empty shells on Chlamys mawsoni Fletcher (rephoto-
graphed from Fletcher, 1938, pl. 10, fig. 1 ) ; B, C, Balanus sp. Hennig (?spp.) (rephotographed from Hennig, 1911, pl. 2,
figs. 3-7) ; B, columnar individual with well-developed radii, from left side, X2.7; C, balanid bases, marginally thickened
x
but without ribs, 1 (see pl. XLVII).
Plate XLVII Balanus sp. Hennig ( ? spp.) : A, B, bases of the balanid type, radiating lines indicating well-developed teeth
in margins of probably porous wall, X0.9; C, individual viewed from left side, appearing to have 8 plates ( ! ) , apparently
without radii, X2.7 (rephotographed from Hennig, 1911, pl. 2, figs. 3-7).
Plate XLVIII Xenobalanus globicipitis Steenstrup: Two specimens on margin of porpoise (Tursiops sp.) tail fluke, near
Isla de Guadalupe, Baja California. Note calcareous parietes anchoring the barnacles to the skin of the porpoise, X5.

SYSTEMATIC INDEX
Names in b o l d f a c e indicate new taxa. Species a n d / o r subspecies in italics are synonyms; genera and
higher categories in italics are synonyms or have been changed. Numerals in italics indicate the page on
which the description or primary citation appears. Roman numerals indicate plates.
Aaptolasma, 14, 140 (chart), 141 (fig.), 142 (key), 149, 150, angulare, 10, 43 (key), 48, 49 (fig.), 184, 194, 195, IX
158, 159, 165, 167, 168 annandalei, 60
americanum, 142, 158 ( k e y ) , 159, 161, 162, 164, XXXII, antarcticum, 10, 43 (key), 50 (fig.), 84, 184
XXXVI, XXXVII atlanticum, 84
brintoni, 142, 158 ( k e y ) , 160, 162, 163 (fig.), 164, 168, aurorae, 111, 184
XXXIII, XXXVIII, X X X I X australicum, 58
callistoderma, 142, 149, 158 ( k e y ) , 159, 160 (fig.), 161, b ellum, 91
162, 164, 168, X X X I I , X X X V , XLIII berndti, 11, 43 (key), 57, 184
leptoderma, 142, 155, 157, 158 (key), 159, 164, 765 (fig.), bouveti, 11, 43 ( k e y ) , 52 (fig.), 184
166 (fig.), 167, 168, XXXIII, X L bouvieri, 11, 43 ( k e y ) , 53, 54 (fig.), 92, 93, 184, 185
triderma, 142, 158 ( k e y ) , 159, 162, 164, 168, XXXIII, brevecarinatum, 10, 43 (key), 53, 55 (fig.), 185
XXXIX buccinum, 55, 56 (fig.), 57 (fig.), 58, 194, 195, VIII
Abathescalpellum, 13, 14, 93 (key), 94 (chart), 104, 105 capense, 76
koreanum, 94, 104 compactum, 10, 11, 43 (key), 58 (fig.), 185
Acasta, 141 (fig.), 150 condensum, 76
accumulation, Arcoscalpellum, Scalpellum, 42 convexum, 185
acicularum, Arcoscalpellum, 43 cornutum, 76
Acrothoracica, 5, 9, 17, 25, 26 (fig.), 28, 29, 184, IV curvatum, 91
acutum, Smilium, 38 darwinii, 43 (key), 58, 59 (fig.), 60, 194, 196, IX
africanum, Arcoscalpellum, Scalpellum, 47 discoveryi, 118
alatum, Scalpellum, 72 dubium, 46
alboranense, Neoscalpellum, Scalpellum, 96, 99 elegans, 58
alcockianum, Arcoscalpellum, 60 elongatum, 58
Alepas, 36 eumitos, 76
Altiverruca, 135 (key) fissum, 130
americanum, Aaptolasma, Hexelasma, 161 formosum, 10, 43 (key), 60, 61 (fig.), 62, 91, 194, 196, VIII
anatifera, Lepas, 31 formosum, 87
Anelasma, 36 gaussi, 11, 43 (key), 62 (fig.), 185
anglicum, Scalpellum, 124 giganteum, 60
angulare, Arcoscalpellum, Scalpellum, 48 gigas, 60
annandalei, Arcoscalpellum, 60 hirsutum, 62, 63 (fig.), 194, 196
Annandaleum, 13, 21, 96 (key), 121, 722, 123 (chart) imbricotectum, 64, 65 (fig.), 66, 194, 196, IX
gruvelii, 722, 123 (chart) improvisum, 186
japonicum, 722, 123 (chart) incisum, 84
lambda, 722, 123 (chart) juddi, 60
subflavum, 722, 123 (chart) latusculum, 11, 43 (key), 66, 67 (fig.), 68 (fig.), 194, 196,
antarcticum, Arcoscalpellum, Scalpellum, 50 VIII
antarcticum, Euscalpellum, 40 liberum, 11, 43 (key), 66, 68, 69 (fig.), 186, 194, 195, 196,
antarcticum, Bathylasma, Hexelasma, 143 IX
aptiensis, Cretiscalpellum, 42 magnaecarinae, 11, 43 ( k e y ) , 70 (fig.), 186
aptiensis antarcticum, Cretiscalpellum, 41 michelottianum, 42, 60, 64, 77 (fig.), 73, 108, 194, 196, IX
arafurae, Hexelasma, 755 moluccanum, 60
Arcoscalpellum, 10, 12, 13, 14, 37 (key), 42 (chart), 43, multicostatum, 11, 43 (key), 73, 74 (fig.), 75 (fig.), 76,
58, 96, 108, 125, 130 194, 195, IX
accumulatum, 42 nymphonis, 118
acicularum, 10, 43 ( k e y ) , 44 (fig.), 45 (fig.), 194, 196, parallelogramma, 76, 11 (fig.), 78 (fig.), 79, 194, 195, 196,
VIII IX
africanum, 43 (key), 47, 47 (fig.), 194, 196, VIII praeceps, 58
alcockianum, 60 recurvirostrum, 11, 43 (key), 79, 80 (fig.), 186
247
248 SYSTEMATIC INDEX
regina, 60 flosculus, 9, 10, 169 (chart), 171

regium, 60 flosculus sordidus, 168, 170, 171 (fig.), 189, XLVI
regium latidorsum, 60 hameri, 171
regium ovale, 60 hirsutus, 138, 149
regulus, 60 hoekianus, 138, 142, 171, 172, 173
rigidum, 58, 91 imperator, 10, 768 (chart), 169
semisculptum, 67, 68 laevis, 10, 14, 169 (chart), 775 (fig.), 176 (fig.), 194, 195,
species, cf. sinuatum, 87 (fig.), 82, 194, 196, IX XIII, XLIV
species, 82, 83 (fig.), 194, VIII laevis coquimbensis, 175
talismani, 91 laevis fossilis, 175
tenue, 67, 68 laevis nitidus, 175, 176
triangulare, 43 (key), 84, 85 (fig.), 186 pentacrini, 169 (chart), 773, 174, XLII, XLIII
tritonis, 60 sordidus, 170
utinomii, 86, 91 species, 10, 149, 169 (chart), 777, 190, XLVI, XLVII
velutinum, 42, 73 tintinnabulum, 776
ventricosum, 43 (key), 86 (fig.), 186 vestitus, 9, 168, 169 (chart), 170 (fig.), 177, 194, 195, XLI
vitreum, 10, 43 (key), 58, 60, 86, 87, 88 (fig.), 89 (fig.), Balanus (Austrobalanus), 9, 10, 12, 139, 168 (chart), 169, 178
90 (fig.), 91, 118, 194, 196, VIII Balanus (Balanus), 775
weltneri, 10, 11, 43 (key), 92, 93 (fig.), 186, 194, 195, VIII Balanus (Bathybalanus), 773
arcuatum, Scalpellum, 86 Balanus (Chirona), 777
arcuatum, Scalpellum, 86 Balanus (Hesperibalanus), 155, 174, 175
aries, Smilium, 37 Balanus (Hexaereusia), 141 (fig.)
Ascothoracica, 5, 11, 12, 23, 184 Balanus (Megabalanus), 776
Ascothorax, 15, 23 species, 10
bulbosus, 10, 11, 23 (key), 24 (fig.), 184 Balanus (Metabalanus), 142, 172
ophioctenis, 23, 24 Balanus (Solidobalanus), 139, 173, 174, 175
species, 10 bartonianum, Scalpellum, 724
atlanticum, Arcoscalpellum, 84 Bathybalanus, IX, 10, 139, 142, 773, 174, 175, XLVI
aucklandicum, Bathylasma, Scalpellum, Pollicipes, Hexelasma, pentacrini, 142, 773
151 Bathylasmatidae, 14, 137 (key), 738, 140, 141 (fig.), 142,
auritum, Conchoderma, 36 143 (fig.), 149, 151, 154, 155, 160, 172, 173, 174, XLIII
aurorae, Litoscalpellum, Scalpellum, Arcoscalpellum, 111 Bathylasma, 7, 8, 10, 14, 26, 138, 141 (fig.), 142 ( k e y ) , 743,
australicum, Arcoscalpellum, 58 149, 152, 154, 155, 158, 168, 177
australis, Lepas, 31 antarcticum, 143, 151, 189
australis var. weltneri, Lepas, 31 aucklandicum, 140 (chart), 142, 143 (key), 149, 150, 757,
Australophialus, 11, 25 154, X X V
Australscalpellum, 12, 13, 14, 37 ( k e y ) , 124 (chart), 130, corolliforme, 8 (fig.), 9, 10, 11, 14, 25, 140 (chart), 142, 743
133 ( k e y ) , 144 (fig.), 145 (fig.), 149, 150, 151, 154, 194, 195,
schizmatoplacinum, 11, 124 (chart), 130, 131, 132 (fig.), 196, X V , XVI, XVII, XLIII
194, 195, XII hirsutum, 8, 12, 14, 140 (chart), 142, 143 (key), 749, 150
Austrobalanus, 9, 10, 12, 139, 168, 169, 178 (fig.), 151, 154, XXII, XXIV
flosculus sordidus, 168, 170, 171 (fig.), 189, XLVI helium, Arcoscalpellum, Scalpellum, 87
bengalense, Smilium, Euscalpellum, 37
vestitus, 9, 168, 169, 170 (fig.), 177, 194, 195, XLI
benthophila, Neoscalpellum, 96
avenionense, Scalpellum, 725
berndti, Arcoscalpellum, Scalpellum, 57
bicornuta, Verruca, 136
balaenaris, Coronula, 178 bigelovii, Scalpellum, 119
Balanidae, 12, 23, 137 (key), 138, 139, 140, 141 (fig.), 142, bouveti, Arcoscalpellum, Scalpellum, 52
143, 168, 169, 173, X X X I V bouvieri, Arcoscalpellum, Scalpellum, 53
Balaninae, 140, 141 (fig.), 142, 182 Brachyzapfes, 9, 11, 25, 28
balanoides, Scalpellum, 54 eiliptica, 9, 25 (chart), 29
Balanomorpha, 7, 8, 9, 12, 14, 17, 18, 20, 21, 23, 137, 138, 139, eiliptica gigantea, 9, 28, 184, IV
141 (fig.), 143, 146, 154, 155, 158, 189 brevecarinatum, Arcoscalpellum, Scalpellum, 53
Balanus, 10, 18 (fig.), 138, 141 (fig.), 142, 143, 150, 158, 159, Briarosaccus callosus, 10, 11, 23 (key), 783 (fig.), 184
168, 171, 172, 173, 775, 178 brintoni, Aaptolasma, 762
balanus, 775 Brochia bulata, 14, 37 (key), 124 (chart), 130, 733, 134
campbelli, 177 (fig.), 194, 196, XII
corolliformis, 138, 143, 148, 159 bulbosus, Ascothorax, 23
corolliformis, 159 buccinum, Arcoscalpellum, 55
decorus, 170, 776 (fig.), 177 (fig.), 178, 194, 195, XLV bulata, Brochia, 733
evermanni, 142, 169 (chart), 777, 172 (fig.), 173 burdigalense, Scalpellum, 724
SYSTEMATIC INDEX 249
Calantica, 37, 38, 40 diadema, Coronula, 779

pedunculostriata, 37 dicheloplax, Neoscalpellum, Scalpellum, 96
pilsbryi, 37 dicheloplax benthophila, Neoscalpellum, Scalpellum, 96
calcaratum, Scalpellum, 125 discoveryi, Litoscalpellum, Scalpellum, Arcoscalpellum, 117
callistoderma, Aaptolasma, Balanus, Hexelasma, 159 Dosima, 32
callosus, Briarosaccus, 183 dubium, Arcoscalpellum, 46
Cameraverruca, 135 (key)
campbelli, Balanus, 177 edwardsi, Neoscalpellum, Scalpellum, 96
capense, Arcoscalpellum, 76 elegans, Arcoscalpellum, 58
Catophragmus, 139, 141 (fig.) Elminius, 138 (key), 139, 140, 141 (fig.), i58, 159, X X X I V
Chamaesipho, 139, 141 (fig.) elongata, Simonizapfes, 28
Chelonibia, 139, 141 (fig.) elongatum, Arcoscalpellum, 58
Chelonibiinae, 141 (fig.), V 2 elliptica, Brachyzapfes, 25
Chionelasmus, 139, 141 (fig J elliptica gigantea, Brachyzapfes, 28
Chirona, 139, 171, 172, 173 eltaninae, Neoscalpellum, 103
evermanni, 142, 169, 171, 172 (fig.), 173 Emersoniinae, 141 (fig.), 142
chitinosum, Scalpellum, 122 erectum, Scalpellum, 72
Chthamalidae, 137 (key), 138, 139, 141 (fig.), 142, 143 (fig.), eumitos, Arcoscalpellum, 76
155, 173, 174 Euscalpellum, 9, 12, 40
Chthamalus, 139, 141 (fig.) antarcticum, 9, 40, 186, VI
scabrosus, 171 bengalense, 37
cochlearium, Mesoscalpellum, Scalpellum, 119 rostra turn, 40
compactum, Arcoscalpellum, Scalpellum, 58 evermanni, Balanus (Chirona), 777
comptum, Scalpellum (Arcoscalpellum), 42 eximium, Scalpellum, 72
Conchoderma, 29 ( k e y ) , 32, 34
auritum, 10, 30 ( k e y ) , 36, 181, 186, 187, III, XLVIII
fascicularis, Lepas, 33
virgatum, 7, 29 (chart), 30 (key), 35 (fig.), 36, 37, 194,
faurei, Scalpellum, 124
195, V
fischeri, Scalpellum, 124
concinna, Zeugmatolepas, 42
fischeri costatum, Scalpellum, 724
condensum, Arcoscalpellum, 76
fissicarinatum, Litoscalpellum, 108
convexum, Litoscalpellum, Scalpellum, Arcoscalpellum, 111
fissum, Arcoscalpellum, 130
coquimbensis, Balanus, 175
flosculus, Balanus (Austrobalanus), 170
cornutum, Arcoscalpellum, 76
flosculus sordidus, Balanus (Austrobalanus), 170
corolliforme, Bathylasma, Hexelasma, 143
formae, Scalpellum, 724
corolliformis, Balanus, 143
formosum, Arcoscalpellum, Scalpellum, 60
Coronula, 10, 23 (key), 168, 178, 181, 182, 190, III, XLVIII
formosum, Scalpellum, 87
balaenaris, 178, 179, 180
fossilis, Balanus, 175
diadema, 779, 180, 190, III
fosteri, Hexelasma, 755
reginae, 178, 179 (fig.), 180, 190
Coronulinae, 141 (fig.), 142
gaussi, Arcoscalpellum, Scalpellum, 62
Cretiscalpellum, 9, 12, 41
georgiensis, Zeugmatolepas, 42
aptiensis, 42
gibberum, Scalpellum, 725
aptiensis antarcticum, 9, 41, 186, VI
gibbosa, Verruca (Altiverruca), 735
unguis, 41
gibbosa somaliensis, Verruca, 136
Creusia, 141 (fig.)
gibbum, Scalpellum, 123
Cryptophialidae, 25
giganteum, Arcoscalpellum, 60
Cryptophialus, 8, 9, 10, 11, 13, 25 (chart) ( k e y ) , 26 (fig.),
gigas, Arcoscalpellum, Scalpellum, 60
27, 28
globicipitis, Xenobalanus, 180
melampygos, 25 (chart) (key), 26 (fig.), 27, 28
gracile, Scalpellum, 87
minutus, 25
gruvelii, Annandaleum, Scalpellum, 122
tomlinsoni, 10, 23 (key), 25 (chart) (key), 26 (fig.), 27,
gruvelii var. quadratum, Annandaleum, Scalpellum, 122
28, 194, 195
Gymnoscalpellum, 12, 13, 37 (key), 93 ( k e y ) , 94 (chart),
turbonis, 25 (chart) (key), 26 (fig.), 28 105, 107, 115, 119, 125
curiosum, Scalpellum, 122
insigne, 94 (chart), 105, 107, 108
curvatum, Arcoscalpellum, 91
larvale, 94 (chart), 105, 107, 108
leoni, 94 (chart), 105, 108, 186
dalpiazi, Scalpellum, 124 tarasovi, 11, 34, 94 (chart), 105, 106 (fig.), 107, 108, 194,
darwini, Verruca, 136 195, 196, X I
darwinii, Arcoscalpellum, Scalpellum, 58
debile, Neoscalpellum, Scalpellum, 96 hameri, Balanus, 171
decorus, Balanus (Mebagalanus), 776 hastatum, Scalpellum, 39
hausmanni, Zeugmatolepas, 42 anatifera, 29 (chart), 30 (fig.) ( k e y ) , 37, 32, 33, 194, 195, V
Hesperibalanus, 156, 174, 175 australis, 7, 10, 29 (chart), 30 ( k e y ) , 37, 32 (fig.), 33 (fig.),
Hexacreusia, 141 (fig.) 187, 188, 194, 195, V
Hexelasma, 10, 14, 138, 140, 142 (key), 143, 148, 149, 154, 755, australis var. weltneri, 31, 188
156, 158, 159, 165, 166, 167, 168, 172, 173 fascicularis, 7, 29 (chart) ( k e y ) , 33, 34 (fig.), 194, 195, V
americanum, 138, 139, 161 hillii, 188
antarcticum, 5, 10, 139, 143, 144, 148, 152, 154, 189 scalpellum, 123
arafurae, 138, 142, 755 ( k e y ) , 156, 157 Lepas (Dosima), 32
aucklandicum, IX, 10, 138, 148, 149, 151 Lepas (Lepas), 37 (key)
callistoderma, 138, 139, 159 leptoderma, Aaptolasma, 765
corolliforme, IX, 138, 143, 148, 149, 189 liberum, Arcoscalpellum, Scalpellum, 68
fosteri, 142, 755 ( k e y ) , 156 (fig.), 157 (fig.), 158 Litoscalpellum, 12, 13, 14, 21, 37 (key), 95 (chart), 96
hirsutum, IX, 138, 143, 148, 149 (key), 108, 110, 115, 119, 122
hoekianus, 138 aurorae, 11, 95 (chart), 108, 777 (fig.), 184
velutinum, 138, 142, 755 (key), 156, 157, 158, 165, 166, 167 convexum, 11, 95 (chart), 108, 777, 112 (fig.), 185
velutinum, 165 discoveryi, 10, 11, 95 (chart), 108, 777, 118 (fig.), 185, 194,
hillii, Lepas, 188 195, X
hirsutum, Arcoscalpellum, Scalpellum, 62 fissicarinatum, 11, 95 (chart), 108, 109 (fig.), 110, 115,
hirsutum, Bathylasma, Hexelasma, 149 117, 194, 195, X
hirsutus, Balanus, 149 intermedium, 95 (chart), 108
hoeki, Verruca, 735 intermedium, 113
hoekianus, Balanus (Metabalanus), 138 korotkevitshae, 108, 186
hoekianus, Hexelasma, 138 laccadivicum, 95 (chart), 108
hungaricum, Scalpellum, 724 nipponense, 95 (chart), 108, 772, 113 (fig.)
hypocrites, Smilium, 37 simplex, 11, 95 (chart), 108, 110, 774 (fig.), 115, 117,
194, 195, X
imbricotectum, Arcoscalpellum, 64 walleni, 11, 95 (chart), 108, 110, 776 (fig.), 117, 194, 196, X
imperator, Balanus, 168 longirostrum, Scalpellum, 39
imperfectum, Mesoscalpellum, Scalpellum, 119 longius, Scalpellum, 122
improvisum, Arcoscalpellum, 186 lovisatoi, Scalpellum, 124
incisum, Arcoscalpellum, 84
insigne, Gymnoscalpellum, Scalpellum, 105 magnaecarinae, Arcoscalpellum, Scalpellum, 70
intermedium, Litoscalpellum, Scalpellum, 108 magnum, Scalpellum, 724
intermedium, Scalpellum, 113 marginatum, Neoscalpellum, 96
Megabalanus, 175, 776, 177
japonicum, Annandaleum, 722 decorus, 170, 776 (fig.), 177 (fig.), 178, 194, 195, XLV
japonicum, Neoscalpellum, Scalpellum, 122 tintinnabulum, 776
japonicum biramosum, Scalpellum, 122 Megalasma, 155
japonicum metapleurum, Scalpellum, 122 melampygos, Cryptophialus, 25
javanicum, Mesoscalpellum, Scalpellum, 119 Mesoscalpellum, 12, 13, 37, 95 (chart), 96 (key), 115, 778, 119,
juddi, Arcoscalpellum, 60 122, 125
cochlearium, 95 (chart), 779
kaempferi, Trilasmis, 155 imperfectum, 13, 95 (chart), 779, 120 (fig.), 121, 122
kampeni, Smilium, 37 javanicum, 95 (chart), 779
Kentrogonida, 783 sanctaebarbarae, 95 (chart), 779, 121 (fig.), 122
koreanum, Abathescalpellum, Scalpellum, 104 Metabalanus, 138, 139, 171, 172, 173
korotkevitshae, Litoscalpellum, Scalpellum, 108 hoekianus, 138
Metaverruca, 735 (key)
minutus, Cryptophialus, 25
laccadivicum, Litoscalpellum, 108
mitra, Verruca, 136
laccadivicum var. investigatoris, Scalpellum, 108
michelottianum, Arcoscalpellum, Scalpellum, 71
laevis coquimbensis, Balanus, 175
mockleri, Zeugmatolepas, 42
laevis fossilis, Balanus, 175
molinianum, Scalpellum, 124
laevis laevis, Balanus, 775
molliculum, Scalpellum, 122
laevis nitidus, Balanus, 175
lambda, Annandaleum, 722 moluccanum, Arcoscalpellum, Scalpellum, 60
larvale, Gymnoscalpellum, Scalpellum, 105 moraviense, Scalpellum, 724
latusculum, Arcoscalpellum, 66 multicostatum, Arcoscalpellum, 73
leoni, Gymnoscalpellum, Scalpellum, 105
Lepadidae, 23, 29 nauckanum, Scalpellum, 724
Lepadomorpha, 10, 12, 17, 18, 20, 23, 29, 143, 184 Neoscalpellum, 12, 13, 15, 21, 37 ( k e y ) , 93 (key), 94 (chart),
Lepas, 18 (fig.), 29 (key), 30, 37, 32, 34, 36 95, 96, 102, 104, 105, 115, 119, 125, 130
alboranense, 96 Scalpellidae, 12, 20, 23, 37, 119, 130, 133

benthophila, 96 scalpellum, Scalpellum, Lepas, 123
debile, 94 (chart), 96, 97 (fig.), 98 (fig.), 99, 100, 102 Scalpellum, 12, 13, 14, 18 (fig.), 20, 37 (key), 42, 53, 96, 111
dicheloplax, 96 122, 123 (chart), 124 (chart), 125, 128, 130, 131
eltaninae, 94 (chart), 96, 103 (fig.), 104, 194, 196, X accumulatum, 42
japonicum, 99 acutum, 38
marginatum, 94 (chart), 96 africanum, 47
phantasma, 94 (chart), 96, 99, 100 (fig.), 102 alatum, 72
schizoplacinum, 11, 13, 94 (chart), 96, 101 (fig.), 102, alboranense, 96, 99
194, 196, X anglicum, 123 (key), 124, 125
nettlebladti, Scalpellum, 124 anglicum, group of, 124
nipponense, Litoscalpellum, Scalpellum, 112 angulare, 48
nitidus, Balanus, 175 antarcticum, 50
nudipes, Smilium, 37 arcuatum, 86
nymphonis, Arcoscalpellum, Scalpellum, 118 arcuatum, 86, 87
aucklandicum, 151
Octolasmis, 130 aurorae, 111, 184
Octomeris, 139, 141 (fig.) avenionense, 125
ophioctenis, Ascothorax, 23, 24 balanoides, 54
ornatum, Scalpellum, 124 bartonianum, 124
ovatum, Scalpellum, 96 bellum, 60, 87
berndti, 51
Pachylasma, 138, 139, 141 (fig.), 151, 154, 155 benthophila, 96
parallelogramma, Arcoscalpellum, Scalpellum, 76 bigelovii, 119, 122
parvulum, Smilium, 37 bouveti, 52
patagonicum, Scalpellum, 125 bouvieri, 53, 112
pedunculatum, Scalpellum, 51 brevecarinatum, 51, 53, 63
pedunculostriata, Calantica, 37 burdigalense, 124
Peltogastridae, 183 calcaratum, 125
pentacrinatum, Scalpellum, 51 chitinosum, 122
pentacrini, Balanus (Bathybalanus), 173 cochlearium, 119
peronii, Smilium, 37 compactum, 53, 58
pfeifferi, Scalpellum, 123 comptum, 42
phantasma, Neoscalpellum, Scalpellum, 99 convexum, 53, 111, 112
pilsbryi, Calantica, 37 curiosum, 122
pilsbryi, Scalpellum, 37 dalpiazi, 124
pilsbryi, Tessarelasma, 155 darwinii, 58
Platylepas, 182 debile, 96
pollicipedoides, Smilium, 37 dicheloplax, 96, 97, 99
Pollicipes, 41, 151 dicheloplax benthophila, 96, 97, 99
aucklandicum, 151 discoveryi, 58, 118, 185, 186
unguis, 41 edwardsi, 96
polymorphum, Scalpellum, 122 erectum, 72
praeceps, Arcoscalpellum, 58 eximium, 72
Pygophora, 25 faurei, 124
fischeri, 124
rathbuniana, Verruca, 136
fischeri costatum, 124
recurvirostrum, Arcoscalpellum, Scalpellum, 79
formae, 123 ( k e y ) , 124, 125
regina, Arcoscalpellum, 60
formae, group of, 124
reginae, Coronula, 178
formosum, 60, 86
regium, Arcoscalpellum, Scalpellum, 60
formosum, 87
regium latidorsum, Arcoscalpellum, 60
gaussi, 62
regium ovale, Arcoscalpellum, 60
gibberum, 10, 53, 111, 124 (chart), 125, 127 (fig.), 128, 194,
regulus, Arcoscalpellum, 60
195, 196, XII, XIII
Rhizocephala, 5, 11, 12, 23, 183, 184
gibbum, 123 (key)
rigidum, Arcoscalpellum, 58
robustum, Scalpellum, 124 gibbum, group of, 123
Rostratoverruca, 135 (key) gigas, 71
rostratum, Euscalpellum, Scalpellum, 40 gracile, 87
gruvelii, 121, 122
sanctaebarbarae, Mesoscalpellum, Scalpellum, 119 gruvelii var. quadratum, 122
scabrosus, Chthamalus, 171 hastatum, 39
hirsutum, 62 stroemii substroemii, 124

hungaricum, 124 studeri, 124
imperfectum, 119 subnavum, 122
insigne, 105 talismani, 87, 91
intermedium, 80 triangulare, 84
intermedium, 113 vanhoffeni, 10, 11, 13, 124 (chart), 128 (fig.), 129, 186,
japonicum, 99, 100, 101 194, 195, VII
japonicum biramosum, 122 velutinum, 42, 51, 71, 72
japonicum metapleurum, 122 ventricosum, 53, 86
javanicum, 119 verticillatum, 37
koreanum, 104 vitreum, 86, 87
korotkevitshae, 110, 186 vulgare, 123
laccadivicum, 108 weltneri, 92, 112
laccadivicum var. investigatoris, 108 schizmatoplacinum, Australscalpellum, 131
lambda, 122 schizoplacinum, Neoscalpellum, 101
larvale, 105 scorpio, Smilium, 37
leoni, 108, 186 semisculptum, Arcoscalpellum, 67, 68
liberum, 68, 71 sessile, Scalpellum, 51
longirostrum, 39 sexcornutum, Scalpellum, 37
longius, 122 sigmoideum, Scalpellum, 124
lovisatoi, 124 simplex, Litoscalpellum, Scalpellum, Arcoscalpellum, 114
magnaecarinae, 70, 71 Simonzapfes elongata, 28
magnum, 123 ( k e y ) , 124 sinense, Smilium, 37
magnum, group of, 124 sinuatum, cf. Arcoscalpellum species, Scalpellum, 81
michelottianum, 42, 71 Smilium, 18, 37 (key), 38, 40
molinianum, 124 acutum, 37, 38 (fig.), 39, 40, 124 (chart), 194, 195, V
molliculum, 122 aries, 37
moluccanum, 71 bengalense, 37
moraviense, 124 hypocrites, 37
nauckanum, 124 kampeni, 37
nettlebladti, 124 nudipes, 37
nipponense, 112 parvulum, 37
nymphonis, 58, 118, 186 peronii, 37
ornatum, 124 pollicipedoides, 37
ovatum, 96 scorpio, 37
parallelogramma, 76 sinense, 37
patagonicum, 125 squamuliferum, 37
pedunculatum, 51 uncus, 37
pentacrinatum, 51 Solidobalanus, 139, 173, 174, 175
pfeifferi, 123 somaliensis, Verruca, 136
phantasma, 99 sordidum, Scalpellum, 72
pilsbryi, 37 sordidus, Balanus, 170
polymorphum, 122 southwardi, Tetrachaelasma, 152
recurvirostrum, 79 squamuliferum, Smilium, 37
regium, 71 stearnsi, Scalpellum, 123, 130
robustum, 124 stearnsi inerme, Scalpellum, 123
ro stratum, 40 striatum, Megalasma, 155
sanctaebarbarae, 119 stroemii, Scalpellum, 123, 124, 129, 133
scalpellum, 123 (key) stroemii latirostrum, Scalpellum, 124
scalpellum, group of, 123 stroemii substroemii, Scalpellum, 124
sessile, 51 studeri, Scalpellum, 124
sexcornutum, 37 subflavum, Annandaleum, Scalpellum, 122
sigmoideum, 124 sulcata, Verruca, 136
Synagogidae, 23
simplex, 42
sinuatum, 81
sordidum, 72 talismani, Arcoscalpellum, Scalpellum, 91
stearnsi, 123, 130 tarasovi, Gymnoscalpellum, 105
stearnsi inerme, 123 tenue, Arcoscalpellum, 67, 68
stroemii, 123 (key), 124, 129, 133 Tessarelasma, 14, 139, 141 (fig.), 142 ( k e y ) , 149, 155
stroemii, group of, 123 pilsbryi, 140 (chart), 142, 155
Stroemii latirostrum, 124 Tetrabalanus, 141 (fig.)
Tetrachaelasma, 8, 14, 141 (fig.), 142 ( k e y ) , 143, 149, 151, gibbosa somaliensis, 136, 188, 189
752, 154, 158, XXVI hoeki, 735
southwardi, 8, 10, 12, 14, 140 (chart), 142, 152, 153 (fig.), mitra, 136
194, 196, XXVI, XXVII, XXVIII, XXIX, X X X , X X X I rathbuniana, 136
Tetrachthamalus, 141 (fig.) somaliensis, 135
Tetraclita, 139, 141 (fig.), 158, 159, XXXIV sulcata, 136
Tetraclitidae, 137 (key), 139, 140, 747 (fig.), 143, X X X I V Verruca (Altiverruca), 735 (key)
Tetraclitinae, 168 Verruca (Cameraverruca), 735 (key)
Thoracica, 7, 11, 12, 79 (fig.), 20, 29, 184 Verruca (Metaverruca), 735 (key)
tintinnabulum, Balanus (Megabalanus), 176
Verruca (Rostratoverruca), 735 (key)
tomlinsoni, Cryptophialus, 25
Verruca (Verruca), 735 (key)
triangulare, Arcoscalpellum, Scalpellum, 84
Verrucidae, 135
triderma, Aaptolasma, 764
Verrucomorpha, 7, 12, 14, 18, 23, 735, 188
Trilasmis, 130
verticillatum, Scalpellum, 37
kaempferi, 155
tritonis, Arcoscalpellum, 60 vestitus, Balanus, 768
Trypetesa, 28 virgatum, Conchoderma, 35
Tubicinella, 181, 182 vitreum, Arcoscalpellum, Scalpellum, 87
turbonis, Cryptophialus, 25 vulgare, Scalpellum, 123
uncus, Smilium, 37 walleni, Litoscalpellum, 776

unguis, Cretiscalpellum, Pollicipes, 41 weltneri, Arcoscalpellum, Scalpellum, 92
utinomii, Arcoscalpellum, 86
Xenobalanus, 10, 23 ( k e y ) , 137, 146, 168, 180, 182, III, XLVIII

vanhoffeni, Scalpellum, 128 globicipitis, 180, 190, XLVIII
velutinum, Arcoscalpellum, Scalpellum, 73
velutinum, Hexelasma, 755
velutinum, Hexelasma, 165 Zapfella, 28, 29
ventricosum, Arcoscalpellum, Scalpellum, 86 Zapfellidae, 28
Verruca, 12, 13, 18, 135 (key) Zeugmatolepas, 9, 12, 42
bicornuta, 136 concinna, 42
darwini, 136 georgiensis, 9, 41, 42, 186, VII
gibbosa, 10, 14, 23 (key), 124 (chart), 135, 136 (fig.), hausmanni, 42
137, 188, 189, 194, 196, XIV mockleri, 42
SUBJECT INDEX
Africa determinate, 140, 162
Cape Town, 31 diametric, 140, 159, 164
Equatorial Guinea, 119 malformation, 57
North, 14, 175 pattern, 8, 146
South, 11, 26, 28, 31, 33, 36, 76, 82, 119, 187 rearmament during, 130, 133
Tanzania, Dar es Salaam, 99 replication during, 148
anatomy and biology longitudinal canal (tube), 18, 138, 139, 140, 141
abdomen, 23 chitinous, 158
ala, alar angle, 143 male
appendages, head, 18, 19, 20, 23, 137, 138, 173 complemental, 38, 42
appendages, thoracic depression for, 43, 66
acanthopod, 7, 19, 29, 32, 34 dwarf, 8, 17, 18, 23, 29, 42, 46, 132
antenniform ramus, 19, 140, 150, 153 nonfeeding, 8, 25
card (grapple), 19, 137 muscles, 17-19
cirral counts, 21 nauplius, 8, 9, 11, 19, 84, 183
cirral types, 19 opercular plates, valves, parts, bars, 17, 18, 21, 23, 26-28,
cirrus, 17, 21 139, 140, 141, 145
number of articles, growth, 147, second-fourth, 19, 137 ovigerous lamella, 17, 60
139 parasitic, 23, 183
ctenopod, 7, 19, 30 peduncle, 18, 23
lasiopod, 7, 19 radius, 18, 138-142, 161, 172
maxilliped, 24 pseudo, 181
setae, types, 19 rearmament, 130, 133
vestigial cirri, 67 replication, substrate, 148
armament, 20 rostrum, 18, 139-141
rearmament, 130, 133 sexes, separate, 8
reduction, 112, 126 sheath, 17, 143, 146
basidorsal point, 19, 137, 174 shells, thickness and age, 146
basis, 137, 138 spawning, seasonal, 8
calcareous, 10, 155, 159; inflection of wall, 155, 158 thin section, 18, 146
membranous, 158 upper latus, abnormal loss of, 56, 74
bivalved carapace, 19, 23 welting, 143, 146, 151, 160
branchia, 17, 147 Antarctica
brood chamber, 8, 9, 128 Ellsworth Land, 88, 91
burrowing, 7, 11, 17, 20, 23, 26, 27, 28 Gaussberg, 92, 129
capitulum, 18, 21, 23 George V Land, 148
carapace, bivalved, 23 Graham Land, 10, 168, 186, 190
caudal appendages, 17, 19, 21, 23, 30, 139-141 Kaiser-Wilhelm II Land, 51, 62
complemental male (see male) MacRobertson Land, 144
compound eye, 19, 26, 27 Marie Byrd Land, 144
compound rostrum, 141 Naze, The, 40
cyprid larva, 8, 11, 19, 132 Palmer Archipelago, 48, 68, 69, 70, 118, 186
dwarf male (see male) Victoria Land, 26, 144
feeding, 7, 8, 25 Australia, 1, 9, 46, 168
filter, 7, 20, 23
piercing, biting, 23 bank
filamentary appendages, 17, 19, 25, 26, 30, 36, 128 Burdwood, 76
float, gas filled, 29, 33 Gunnerus, 189
fouling, 31, 171, 187 Sars, 149, 152, 154
gastric mill, 25
Scotia, 144
growth
basin
allometry, 173 Labrador, 72, 81, 82, 88
dearmament during, 112, 126
Southeast Pacific, 2, 58, 66, 88, 101, 152
254
SUBJECT INDEX 255
bathymetry, currents, and substratum depauperate, 14

abyssal, 7, 9, 96 diversity, 7, 11, 12, 14, 154
Antarctic Convergence, 1, 5, 6, 7, 9, 10, 11, 12, 14, 40, 168, endemism, 8, 10, 11, 13, 14, 177
184 extinction, 9, 12
bathya], 9 fauna, reduction of, 7
bathymetry, 3, 184, 191, 195, 197 Glacial Age, 7
benthonic, 1, 7, 10, 23 hiatus, bathymetric, 13, 14
currents, 1, 7, 28, 31 invasion, 9
nekton, 29, 36 monotypic, 14, 149
pelagic, 1, 7, 10, 23, 29, 33 polyphyletic, 143
sediments, 7 predation, 130, 159
subantarctic waters, 1, 12, 14 ratio, Balanomorpha/Lepadomorpha, 11
substrate species-groups, 12, 27, 123, 168
basalt, 59 West Afro-Caribbean, 9
calcareous, 7, 9, 17 expeditions
limestone, 17, 29 Antarctique Allemande du Gauss, 5
pumice, 33 Australian Antarctic, 148, 193
size, 7, 8 British Antarctic, 2, 5
solid, 7 British-Australian-New Zealand Antarctic, 2, 193
replication (see organisms), 148 Danish, 1922, 165, 167, 193
bay Deutsche Siidpolar, 5
Bengal, 40 Japanese Antarctic Research, 6, 148, 192
Chesapeake, 119 Jyn III, 164
Commonwealth, 5, 185 Naga, 162, 164, 193
Cumberland, 111 National Antarctic, 5
Terra Nova, 144, 188 New Zealand Antarctic Research, 2, 193
West Cumberland, 185 Swedish South Polar, 111
Tui, 155, 193
Burma, 143, 155
cape Falkland Trough, 64, 125

Good Hope, 119, 187
Hatteras, 119 geologic history
Horn, 1, 31, 33, 152, 171, 175, 188 Cenozoic, 7, 9, 168
Central America, 13 continental drift, 1
channel Cretaceous, 1, 6, 9, 11, 14, 15, 28, 29, 37, 40, 41, 42, 184,
Faeroe, 138 186
Neumayer, 69, 70 Eocene, 9
coast fossil, 6, 10, 148, 189
Adelie, 111 freezing, 1, 15
Alexander, 9, 28, 29, 41, 184, 186 glaciation, 1, 7, 12, 15
Bryan, 88 glacier, 5, 148, 188
Gondwanaland, 1
Ingrid Christensen, 51, 84
ice, 1
Prince Harald, 144, 148
interglacial, 10
Wilhelm II, 92, 129
Jurassic, 9, 42
land connections, 1
Drake Passage, 43, 152, 187
Mesozoic, 9, 139
Miocene, 9, 10, 14, 143, 148, 151, 155, 175, 190
Europe, 9, 14, 175 Oligocene, 9, 10, 14, 138, 143, 168
Britain, 9, 149, 186 Paleocene, 15
Denmark, 9 Pleistocene, 7, 9, 10, 14, 143, 149, 168, 175, 177, 188
Hungary, 175 Pliocene, 10
Ireland, 149 pre-Pleistocene, 148
Northern, 14 Quaternary, 6
evolution Scallop Hill Formation, 10
austral group, 11, 21, 25 sea-floor spreading, 1
bipolar, 10, 11, 15 subfossil, 5
burrowing, 20 Tertiary, 9, 10, 14, 15, 138
circumantarctic (circumpolar), 7, 10, 13, 42, 187 gulf
competitive exclusion, 154 Aden, 40
Coral Seas, 7 Mexico, 67
256 SUBJECT INDEX
harbor Santa Catalina, 99

Granite, 58, 185 Sao Miguel (Ilha d e ) , 8 4
Stromness, 185 Scott, 189
Hatteras Plain, 72, 88 Seymour, 185
Snow Hill, 185
island (s) Solomon, 185
Adelaide, 47, 48 South Georgia, 5, 9, 23, 74, 111, 118, 137, 144, 184, 185,
Alexander, 9, 28, 29, 41, 184, 186 186, 187, 190
Andaman, 40 South Orkney, 61, 92, 108, 129
Antipodes, 39 South Sandwich, 144
Aspland, 48, 68, 105, 114, 131 South Shetland, 48, 50, 68, 105, 114, 131, 148, 149, 185, 186,
Auckland, 39, 55 189, 190
Azores, 40, 84, 99, 107 Stewart, 5, 187
Balleny, 186 Sumatra, 40
Bermuda, 96, 97 Tasmania, 187
Briscoe, 61, 62 Tristan da Cunha, 48
Borneo, 64 Zanzibar, 62
Bounty, 55
Bouvet, 52, 184 Japan, 12, 40, 62, 76, 86, 91, 113, 138, 149, 156, 160, 164
Brabant, 48, 68, 118
Bridgeman, 105, 131 Museums
Campbell, 170, 177, 187 Australian (Sydney), 145
Celebes, 64 British Natural History (London), 2, 71, 145
Chatham, 31, 168 Dominion (Wellington), 155
Clarence, 50, 184, 185 United States National (Washington, D.C.), 2, 173
Cockburn, 10, 177, 190 Zoological (Amsterdam), 173
Coronation, 92, 129 Zoological (Copenhagen), 165, 166
Crozet, 54, 67
Cuba, 9 New Zealand, 1, 8, 9, 10, 14, 26, 28, 33, 36, 143, 148, 151, 155,
Deception, 190 157, 168, 170, 177, 187
Elephant, 48, 68, 114, 131
Falkland, 5, 11, 32, 43, 76, 77, 125, 133, 152, 175, 184 ocean
Galapagos, 119 Arctic, 11, 13, 133
Graciosa ( I l h a ) , 8 4 Atlantic, 11, 15, 40, 99, 122
Greenland, 72, 81, 82, 88, 91 North, 12, 13, 15, 21, 40, 143, 149, 148
Hawaii, 11, 31, 175, 187 Northeast, 149
Heard, 80 South, 5
Humps, 40 Western, 119, 138, 158, 161, 162, 184
Iceland, 119 Indian, 5, 40
Inaccessible, 108 Indo-Pacific, 1, 14, 15, 159
Juan Fernandez (Islas), 31, 188 Indo-West Pacific, 9, 11, 13, 21, 138, 155, 156, 157, 158, 162
Japan (see Japan) Pacific, 15
Joinville, 68, 74, 131 East, 12, 13, 21, 122
Kepulauan Kai (Kei Is.), 165, 167, 193 North, 11, 138
Kerguelen, 9, 80, 138, 144, 148, 168, 186, 187, 189 Northwest, 14, 15
Kermadec, 31, 40, 188 South, 5, 40, 117, 152
King George, 105 Southeast, 2, 15, 31, 33
Larsen, 108 organisms
Malpelo (Isla), 122 brachiopods, 29, 162, 165
Moluccas, 40, 62, 130 bryozoans, 25, 186
Motutapu, 151 coelenterates
Nantucket, 96, 119 antipatharians, 23, 69
New Zealand (see New Zealand) corals, 7, 17, 25, 26, 29
Nightingale, 48 gorgonians, 186
O'Brien, 105, 131 hydroids, 33, 184-186
Palau Taam, 173 parasites of, 11, 23
Peter I, 58, 117 scyphomedusae, 36
Prince Edward, 54, 67 zoantharians, 86
Renaud, 61 cetacean epifauna, 5, 10, 23, 36, 168
Saint Helena, 187 crustaceans,
Santa Barbara, 120 crabs, and parasites of, 11, 23, 184
SUBJECT INDEX 257
echinoderms Coral, 46
crinoids, 23, 173 Mediterranean, 11, 14
echinoids, 7, 17, 71, 189 Okhotsk, 62
ophiuroids, 23, 184 Ross, 1, 5, 9, 11, 26, 118, 144, 145, 184, 185, 186
parasites of, 11, 23 Scotia, 61, 62
feathers, 33 South China, 162
fish, 11, 36, 187 Timor, 86
molluscs Weddell, 1
belemnites, 28, 29, 184 ships
gastropods, 11, 102, 130, 154, 159, 170, 177 Albatross, 2, 96, 97, 99, 107, 113, 119, 120, 160, 161, 171, 192
lamellibranchs, 28, 171, 190 Amboina, 165, 167, 193
shells of, 17, 20, 28, 29 Atlantis, 2, 97, 99, 193
polychaete, 185 Aurora, 5, 11, 144, 145, 193
pycnogonids, 50, 118, 185, 186 Beagle, 5, 187
seaweed (kelp), 11, 187, 188 Belgica, 5
sponges, clionid, 28 Chain, 2, 96, 97, 99, 193
tunicate, 50 Challenger, 2, 40, 46, 48, 49, 54, 64, 67, 71, 79, 85, 91, 107,
whales, 10, 186, 187, 190 138, 144, 145, 148, 193
Xiphosura, 159 Discovery, 5, 23, 48, 50, 52, 148, 149, 193
Eastward, 2, 72, 193
peninsula Eltanin, 2, 3, 191, 192, 194, 195, 196, 197
Antarctic, 1, 9, 25, 27, 28, 41, 47, 61, 74, 131, 144 Endeavour, 2, 9, 26, 144, 145, 193
Cape York, 46 Gauss, 5, 51, 62, 92, 129, 184, 185, 186
Otago, 177 Kainan Maru, 5
Riiser-Larsen, 188 Ob\ 6
Peru-Chile Trench, 60, 63 Pourquoi Pas?, 5
plateau Sarsia, 149, 193
Blake, 161 Siboga, 2, 62, 86, 138, 155, 156, 165, 173, 193
Campbell, 39, 55 Soya, 2, 6, 144, 192
Kerguelen, 2 Terra Nova, 2, 5, 144, 145, 148, 185, 193
point Triton, 138
Arenas (Punta d e ) , 125 Umitaka-Maru, 2, 6, 144, 192
Dungeness (Punta), 125 William Scoresby, 5, 23
Flagon, 184 sound
Larsen, 185 Choiseul, 133
program McMurdo, 1, 10, 58, 118, 144, 184, 185, 186
Japanese Antarctic Research, 2 strait
New Zealand Antarctic Research, 2, 193 Bismarck, 70
United States Antarctic Research, 2, 191, 192 Branfield, 149
Magellan, 171, 175, 184
reagents and materials, 17, 18 Makassar, 64
research institutions (also see museums) South America, 5, 9, 14, 15, 36, 119, 168, 175, 189
Auckland University, 2, 193 Argentina (Mar del Plata), 79, 85
Biological Laboratory of the Imperial Household, 86 Brazil, 125, 175
Duke University Marine Biological Laboratory, 2, 193 Chile, 31, 60, 63, 152, 175, 188
Marine Biological Association, United Kingdom, 149, 193 Colombia, 122
New Zealand Geological Survey, 144, 145, 151, 192 Patagonia, 125, 187
New Zealand Oceanographic Institute, 2, 9, 26, 145 Peru, 9, 103, 175
Scripps Institution of Oceanography, 2, 163, 164, 193 Tierra del Fuego, 9, 30, 35, 125, 168, 171, 175
Tokyo University of Fisheries, 145
United States Bureau of Commercial Fisheries, 2, 192 United States
Woods Hole Oceanographic Institution, 2, 193 California, 99, 107, 120, 176
ridges and rises, 1, 3, 54, 67, 137, 157 Florida, 67, 184
Maryland, 82, 119
sampling devices, 191-197 New Jersey, 96
sea North Carolina, 119
Banda, 62, 156 South Carolina, 72, 88
Bellinghausen, 58
Bering, 11, 171, 184 Vietnam, 162

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ANTARCTIC

American Geophysical Union

American Geophysical Vnion

Monographic account based on specimens

William A. Newman and Arnold Ross

AMERICAN GEOPHYSICAL UNION

National Academy of Science-National Research Council

Cop3right @ 1970 bj- the American Geophysical Union

Library of Congress Catalogue Card No. 71.129339

List Price, $25.00

WILLIAMA. NEWMAN ARNOLDROSS

Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Order Ascothoracica Lacaze-Duthiers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Family Synagogidae Gruvel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Genus Ascothorax Djakanov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Order Acrothoracica Gruvel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Suborder Pygophora Berndt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Suborder Lepadomorpha Pilsbry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Key to Antarctic Lepadidae .................................

Genus Euscalpellum Hoek . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Key to Antarctic Species of Arcoscalpellum . . . . . . . . . . . . . . . . . .

Plates .................... ...... . . . . . . . . . . . . . . . . . . . . . . . . ...

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1 Scotia Ridge 7 Albatross Cordillera 12 Mid-Indian Ridge

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Auckland University AU Ncw Zraland Antarctic Rcscarch Expedition NZARP

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by widely flaring the apertural end in the manner so

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A. berrulti (Gruvel, 1907b) Antarctic faunal communications and ascothoracicans

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t Extinct genera and species.

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14. ANTARCTIC CIRRIPEDIA

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MATERIALS AND METHODS

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Lepos Sm111um Sca/pellum s I.

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MATERIALS AND METHODS

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MATERIALS AND METHODS 21

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A Crypfophiolus melompygos Ber ndt Crypfophialus furhnis Bar nard

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tachment disc and slender neck extending to aper-

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I h e anterior projects dorsally as an excentric apex

3.6 0.8 7.2 1.8 4.0

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Lepos austrolis Darwin v Lepos fasciculoris Ell is & Solander

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