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Introduction. A General Statement of the Ideas and the present 
Aims and Status of the Marine Biological Association 

of San Diego, by Wm. E. Eitter. With 2 maps i-xvii 

The Idea of a Marine Biological Survey ii 

The Area to be Surveyed iii 

The Initial Step vii 

Order of Advance on the numerous Lines of Investigation viii 

Knowledge of the Physical Conditions of the Area viii 

Instrumentalities for Prosecuting such a Survey ix 

Necessity of a Salaried Staff x 

Present Status, as to Ways and Means xi 

Historical Note xii 

Eemarks on the present Condition of marine Biology in 

General xiv 

No. 1. The Hydroids of the San Diego Eegion, by Harry Beal 

Torrey. With 23 figures 1-44 

Introduction 1 

Table of Distribution 2 

Key to Families, Genera, and Species 3-5 

Description of Species 6-39 

Bibliography - 40-44 

No. 2. The Ctenophores of the San Diego Eegion, by Harry Beal 

Torrey. Plate 1 45-50 

Description of Species 45-48 

Bibliography - 49-50 

No. 3. The Pelagic Tunicata of the San Diego Eegion, excepting 
the Larvacea, by Wm. E. Eitter. With figures 9-31. 

Plates 2-3 51-112 

Introduction 51-52 

List of Species - 52 

Discussion of Species - 53-101 

Annotated Bibliography 102-107 

Description of Plates 108-112 

No. 4. The Pelagic Copepoda of the San Diego Eegion, by C. O. 

Esterly. With 62 figures 113-233 

Introduction - 113 

Terminology 113-115 

List of Species - 116-117 

Key to Genera 118-121; 205-207 

Description of Species 122-205; 207-226 

Annotated Bibliography 227-233 


No. 5. Non-Incrusting Chilostomatous Bryozoa of the West Coast 
of North America, by Alice Eobertson. With 2 figures 

and Plates 4-16 235-322 

Introduction 235-237 

Terminology 237-241 

List of Species 241 

Description of Species 242-292 

Bibliography 293-296 

Description of Plates 297-322 

No. 6. Differentiation in Hydroid Colonies and the Problem of 

Senescence, by Harry Beal Torrey. With 4 figures 323-332 

Observations 323-331 

General considerations 331-332 

No. 7. The Behavior of Corymorpha, by Barry Beal Torrey. With 

5 figures 333-340 

No. 8. Dinoflagellata of the San Diego Region, I. * >n Eetero- 
dinium, a new Genus of the Peridinidse, by diaries 

Atwood Kofoid. Plates 17-19 341-368 

Introduction and Acknowledgments 341 

Description of Genus Heterodinium 341-342 

Detailed Description of generic Characters 342-351 

Synoptic Key to Species of Heterodinium 350 

Discussion <>t' Species 351 

Bibliography 363 

Description of Plates 364-368 

Index 369-3S2 


Vol. 2, Introduction, pp. i-xvii. April 5, 1905 










ZOOLOGY.— W. E. Ritter, Editor. Price per volume $3.50. Volume I 
(pp. 286) completed. Volume II (in progress): 

Introduction. A General Statement of the Ideas and the Present Aims and 
Status of the Marine Biological Association of San Diego, by 
Wm. E. Ritter. 

No. 1. The Hydroids of the San Diego Region, by Harry Beal Torrey. ] In 

Pages 43, text figures 23. I one 

} cover. 
No. 2. The Ctenophores of the San Diego Region, by Harry Beal j p r j ce 

Torrey. Pages 6, Plate 1. J .60 

No. 3. The Pelagic Tunicata of the San Diego Region, excepting the Lar- 

vacea, by Wm. E. Ritter. Pages 62, text figures 23, Plates 2. 

Price, 65 

The following series in Graeco-Roman Archaeology, Egyptian Archaeology, Ameri- 
can Archaeology and Ethnology and Anthropological Memoirs are publications from the 
Department of Anthropology: 


Vol. 1. The Tebtunis Papyri, Part I. Edited by Bernard P. Grenfell, Arthur 
S. Hunt, and J. Gilbart Smyly. Pages 690, Plates 9, 1903 
Price, $16.00 

Vol. 2. The Tebtunis Papyri, Part 2 (in preparation). 


Vol. 1. The Hearst Medical Papyrus. Edited by G. A. Reisner and A. M. 
Lythgoe (in press). 


Vol. 1. No. I. Life and Culture of the Hupa, by Pliny Earle Goddard. 

Pages 88, Plates 30, September, 1903 . . . Price, 1.25 

No. 2. Hupa Texts, by Pliny Earle Goddard, Pages 290, March, 

1904. Price, 3.00 

Vol. 2. No. 1. The Exploration of the Potter Creek Cave, by William J. 

Sinclair. Pages 27, Plates 14, April, 1904 . . Price, .40 

No. 2. The Languages of the Coast of California South of San 

Francisco, by A. L. Kroeber. Pages 72, June, 1904. Price, .60 
No. 3. Types of Indian Culture in California, by A. L. Kroeber. 

Pages 22, June, 1904 Price, .25 

No. 4. Basket Designs of the Indians of Northwestern California, 

by A. L. Kroeber. Pages 60, Plates 7, January, 1905. Price, .75 
Vol. 3. The Morphology of the Hupa Language, by Pliny Earle Goddard 
(in press). 


Vol. I. Explorations in Peru, by Max Uhle (in preparation). 
No. 1. The Ruins of Moche. 
No. 2. Huamachuco, Chincha, lea. 
No. 3. The Inca Buildings of the Valley of Pisco. 


Vol. 2, Introduction, pp. i-xvii. April 5, 1905 






Director of the Station. 



1. The idea of a Marine Biological Survey ii 

2. The area to be Surveyed iii 

3. The Initial Step vii 

4. Order of Advance on the numerous lines of Investigation viii 

5. Knowledge of the Physical Conditions of the Area viii 

6. Instrumentalities for Prosecuting such a Survey ix 

7. Necessity of a Salaried Staff x 

8. Present Status, as to Ways and Means xi 

9. Historical Note xii 

10. Remarks on the present condition of Marine Biology in General . xiv 

ii University of California Publications. [Zoology 

1.— The Idea of a Marine Biological Survey. 

Volume II of the University of California Publications, 
Zoology, is now either published or in the Editor's hands, waiting 
to be put into type. It is composed exclusively of "Results 
from the Laboratory of the Marine Biological Association of San 
Diego. ' ' 

The investigations on the coast of Southern California now 
having been in progress for several years, and their continuance 
being assured for a few years more at least, it seems fitting that 
this first volume of results should be introduced by some state- 
ment of the general ideas animating the undertaking, and of the 
efforts being made, and means available to realize these ideas. 
Investigations in marine biology, intensiv( rather than extensive 
in character (to borrow a useful agricultural phrase) is the key 
note of the idea. An immediate consequence of the adoption of 
such an idea as a rule of action, has been the necessity of making 
a clear distinction between marint />i<>l<></a, and general />i<>lt><iu 
l>ms< euti d by researches on marim organisms. I have elsewhere 
written as follows of this distinction: 

"The former has for its aim, in the large, the getting of as 
comprehensive an understanding as possible of the life of the sea. 
It, of course, presents itself under a greal variety of secondary 
questions; but the sum total of the phenomena of marint plants 
and animals will never be lost smht of as its real aim. The latter 
makes use of animals and plants that live in the sea in general 
biological researches. That these organisms happen to be marine 
is an incident merely. The investigator turns away from them 
without hesitation when others, from whatever source, come to 
hand that suit his purpose better. Further, the user of marine 
organisms in such investigations is quite indifferent to every- 
thing concerning them that does not bear upon his particular 
problem. He puts aside the marine animal after it has served 
his purpose without having even noticed, perhaps, the major part 
of its traits and qualities and the questions concerning it." 

For this particular undertaking, I believe the ideal, broad 
and general as it is, is eminently useful. It is useful because ii 
uives deli idleness and coordination to action, and furnishes a 
commanding point of view and stimulus. It is justifiable to hold 

Vol. 2] Ritter. — Introduction. iii 

and be guided by it, even though assurance of opportunity to 
carry it out fully is absent. 

The aim as formulated in the articles of incorporation of the 
Association is, "To make a Biological Survey of the waters of 
the Pacific adjacent to the Coast of Southern California." 

2.— Tho Area to be Surveyed. 

The funds available being small, an important and ever- 
present practical question is that of fixing limits. One of the 
first of these was that of limiting the territory to be surveyed. 
The irregularly triangular area extending from Point Concep- 
cion, Lat. 34° 27', at the north, to a base line extending westward 
from the southern boundary of the United States, Lat. 32° 28", 
bounded on the east by the coast line, and on the west by the 
meridian of Point Concepcion, Long. 120° 25', was selected. The 
shore line of this area, exclusive of the islands, is about 280 miles. 
The length of the western side is about 120 miles, and that of its 
southern side about 194 miles. The area contains, therefore, over 
11,600 square miles. 

It is, of course, not to be supposed that a stone wall has been 
built about this area, and that we give no heed to anything out- 
side of it. As a matter of fact, nothing is clearer than that com- 
plete knowledge of it is impossible without extending the explora- 
tions widely beyond it. That it makes a well defined base of 
operations, is about the view we take of it. 

The qualifications of the region are : a. position well to the 
south; a considerable extent of continental shelf, presenting a 
large diversity of bottom, with numerous islands and shoals; 
proximity to oceanic depths and other truly oceanic conditions ; 
a favorable climate; a large variety of shore line; and accessi- 
bility through sea ports and railroads. Two of these advantages, 
that of climate and proximity to oceanic conditions, are held to 
be of very great importance. A fundamental element in investi- 
gations of the sort contemplated is continuoiirsness of the field 
work. Data gathering must go on throughout the year at fre- 
quent intervals. The weather here offers little obstacle to this. 
Heavy storms are rare, and these are practically limited to three 
or four months — January, February, March, and April. For the 


University of California Publications. 


rest of the year there are few days on which, for a portion of the 
day at least, work cannot be carried on anywhere in the area with 
slight interference from heavy seas; and even during the months 
subject to storms only rarely is it interfered with. The practical 
importance of this can hardly be overvalued, as all experienced 
in this sort of work will appreciate. Not only does it make a 
completeness of field observations practicable, that could hardly 
be secured with any kind of a vessel in more storm afflicted 
regions; but it reduces the cost of exploration to the minimum, 
for the work can he done in a vessel much smaller, and hence 
much loss expensive of operation than is ordinarily required for 
such work. Dredging and trawling to a depth of 500 fathoms 
at least from a vessel of 60 foot keel, manned by three men, is 
perfectly feasible; and sounding and various kinds of work on 
surface and intermediate waters can be done at considerably 
greater depths with the same equipment. 

The following table, made up from data contained in the 
Monthly Synopses of the United State Weather Bureau, presents 
information concerning climatic conditions ;it San Diego during 
1904. an entirely typical year. 




O 9 9 

a a a 


Nantucket 34 22 

Key West 73 82 

Farallone 54 49 

San Francisco 56 45 

San Diego 65 47 


Nantucket 32 19 

Key West 76 66 

Farallone 53 49 

San Francisco 55 46 

San Diego 61 48 


I s 

C c 






3.86 15 

1.08 4 

6.13 16 

5.89 16 

1 50 6 





>> a 

■*• c 

S * 
^> « 
> a 

■A 9 





Vol. 2] 

Ritter. — Introduction. 

Temperature. Precipitation. 




Nantucket 39 

Key West 79 

Farallone 60 

San Francisco 57 

San Diego 63 


Nantucket 48 

Key West 80 

Farallone 57 

San Francisco 63 

San Diego 66 


Nantucket 61 

Key West 82 

Farallone 55 

San Francisco 66 

San Diego 65 


Nantucket 65 

Key West 86 

Farallone 55 

San Francisco 66 

San Diego 69 


Nantucket 74 

Key West 87 

Farallone 56 

San Francisco 62 

San Diego 71 


Nantucket 71 

Key West 88 

Farallone 56 

San Francisco 62 

San Diego 76 


Nantucket 67 

Key West 87 

Farallone No 

San Francisco 71 

San Diego 76 



5 5 


H 3 

•A cr> 

23 3 


































































































































































University of California Publications. 


Temperature. Precipitation. 



Nantucket 57 

Key West 82 

Farallone <>1 

San Francisco 68 

San Diego 74 


Xant ucket It; 

Key West 7(5 

Farallone 59 

San Francisco 63 

San Diego 74 


Xannieket 35 

Key Wesi 74 

Farallone 57 

San Francisco 55 

San Diego 66 

*• i 



>. i 

o « 

2 * 















in. 701 



2.3 7 




















1 .58 























.> ...i 














Perhaps the mosl importanl fact, from the presenl poinl of 
view, exhibited by this table is thai pertaining to winds. It will 
be noted, for San Diego, thai the maximum velocity for the year 
was 36 miles an hour, in February. On the basis of the "Beaufort 
Scale" of wind velocities, this is a "Strong Breeze." February 
and March are the climax of the stormy season. For the months 
May to December, inclusive, the maximum velocities run from 
"Gentle Breeze" to "Fresh Breeze." 

La Jolla, the suburb of San Diego at which the laboratory is 
located, is on a rocky point jutting into the open sea with water 
of 200 fathoms attainable inside of five miles: so the ecological 
problems of oceanic plankton, and of bottom-forms can be here 
attacked under peculiarly favorably conditions. 

The western boundary of the area corresponds roughly to 
the edge of the continental shelf in this region, and immediately 
beyond this 2,000 to 2,300 fathoms are reached. While this 
extreme depth is distant about 200 miles from San Diego, by 
making San Nicholas Island a temporary base the 2,000 fathom 

San Miguel 

2I&* fe 08 




Santa. Barbara 

Beqas Rock ° 

San Nicolas Is.^-O 

495- San Clemente Is. 

4I3 657 59 
I08\ ^., 50>9 807 
Tanner Ban'K 


Cortes BciK.K & g2 715 

iSfc.,o '389 

Bishop R'k:...* 7 2 

120 27 

Map 1.— Showing the Area to be surveyed. Modified from United States Coast and Geodetic Survey Chart. 

Vol. 2] Fitter. — Introduction. 


curve is only 65 miles away. Within the area is a wide range of 
depth and great variety of bottom. A basin 40 miles off Point 
Loma has a depth of over 1,000 fathoms. On the other hand, 
the Cortes Banks, just beyond the southern boundary, carry but 
15 feet of water at low tide. 

There can be no doubt that deep sea and 'longshore investiga- 
tions have not yet been brought together to the extent they ought 
to be. 

3. -The Initial Step. 

The first step in such a survey would obviously be to find 
out what plants and animals inhabit the area; to establish a 
speaking acquaintance, as one may say, with the organisms that 
are later to be more intimately known. So far this has absorbed 
most of the effort, and it will of necessity demand the continu- 
ance of much effort for a long time in the future. The ideal 
being kept always in view, the mere description of the new spe- 
cies for the exclusive use of expert taxonomists in the several 
groups, would not be sufficient. The entire fauna and flora must 
be recorded in such a way as to make the records a good founda- 
tion for the broader and deeper studies to follow. These consid- 
erations have determined the character of the faunistic papers 
now published, and that will come hereafter. The present vol- 
ume contains the following contributions to a knowledge of the 
fauna : 

No.l. — The Hydroids of the San Diego Region, by Professor 
H. B. Torrey. 

No. 2. — The Ctenophores of the San Diego Region, by Pro- 
fessor H. B. Torrey. 

No. 3. — The Pelagic Tunicata of the San Diego Region, except- 
ing the Larvacea, by Professor Wm. E. Ritter. 

No. 4. — The Pelagic Copepoda of the San Diego Region, by 
C. 0. Esterly. 

No. 5. — The Nonencrusting cheilostomatous Bryozoa of the 
West Coast of North America, by Dr. Alice Robertson. 

No. 6. — The Dinoflagellata of the San Diego Region, by Pro- 
fessor C. A. Kofoid. 

viii University of California Publications. [Zoology 

Not only are new species described, but all that have thus far 
been found in the area are characterized, and in most cases illus- 
trated by figures, so that these papers will constitute a series of 
hand-books, as far as they go, for the identification of the species 
treated. It is also intended that the bibliographical lists accom- 
panying the papers shall serve as useful guides to the literature 
of the several groups for those who may take them up for the 
study of special problems connected with them. 

4.— Order of Advance on the Numerous Lines of Investigations. 

While there is no reason for attempting a rigorously laid 
out orderof attack on the numerous problems, at natural sequence, 
within certain limits, will establish an order; and where practical 
administrative conditions conveniently adapt themselves to such 
sequence this order will be followed. For example, the species 
representing a given pelagic group having been go1 well in band, 
a natural second step would be the determination of the seasonal 
distribution of the group, since the study of the collections for 
the taxonomy would surely bring together, incidentally, consid- 
erable data on this problem. Following close upon the treatment 
of seasonal distribution would come that of horizontal and ver- 
tical distribution, the chorology ; and inseparably linked with these 
would be the problems of food and reproduction ; and these again 
would lead to problems of migration, with their intimate depend- 
ence upon temperature and other environmental factors. And 
here, completeness of knowledge being ever the watchword, the 
demand would arise for applying experimental and statistical 
methods in the effort to get at the deeper significance of the fads 
observed, and generalizations reached from the observational 
investigations. The chain of questions hanging one to another 
is endless and, of course, completeness of knowledge in a literal 
sense, is an unattainable ideal. 

5.— Knowledge of the Physical Conditions of the Area. 

It does not need to be said, in the light of general biolog- 
ical conceptions reigning in this day, that an aim at comprehen- 
siveness of knowledge cannot for a moment neglect the physical 
conditions under which organisms live. What has to be consid- 





u +4 

Q /Zi**- 






















si 1 


Vol. 2] Fitter. — Introduction. ix 

ered in connection with a marine undertaking like the present, is 
the specific things that must be done, and the means for doing 
them. Oceanography is in position to hand over to the marine 
biologist, ready prepared, a large amount of the information he 
must have ; and, likewise, physics and chemistry have important 
resources that can be drawn upon. But these general sources in 
nowise obviate the necessity for constant and searching studies 
on the sea water in connection with such a survey as that contem- 
plated. Conditions of the water as to temperature, and currents ; 
mineral, gaseous, and albuminoid content, etc., must be known at 
the particular time and place to which the biological studies per- 
tain, and no general knowledge of this character can suffice. 
Physics, chemistry, and hydrography must, therefore, be integral 
parts of such a survey. 

6.— Instrumentalities for Prosecuting such a Survey. 

It is obvious that no small outlay of money would be essen- 
tial for even a good beginning ; and that considerable progress in 
it could be made only with large expenditures for both equipment 
and operation. The ideal laboratory building would not be large, 
but would be constructed with great care. Aquaria would con- 
stitute an important element in the plant for the work on shore. 
From $50,000 to $75,000 should build and equip an ample labora- 
tory and aquaria. 

Equipment for the work at sea would demand the greater 
portion of the capital. For the deep-water work a ship of the 
class of the U. S. Fisheries steamer Albatross would be essential. 
For less depths, say 1,000 fathoms and under, a much smaller 
vessel would be as efficient or even more so, since it can be han- 
dled so much more quickly. As noted above, our area is extremely 
favorable for this purpose. A vessel that could be built and 
made ready for sea (without scientific apparatus) for $10,000 
or $12,000 would be ample. 

Operating expenses would be considerable ; and this leads me 
to speak of the factor most important, but least tried, for the 
successful carrying out of such an idea. 

x University of Calif ornia Publications. [Zoology 

7.— Necessity of a Salaried Staff. 

Obviously, there must be coordinated effort of numerous 
special investigators to make any headway. How is this to be 
secured? In only one way: by paying for it. The diversity of 
talent and training called for, and the prolonged period of service 
requisite, preclude the possibility of success on any other basis. 
Botanists and zoologists there are who would gladly, and without 
thought of money compensation, prepare reports on collections in 
their special groups that might be sent to them; and occasionally 
one would be found not only willing but able to stand the expense 
of a sojourn for a few days or weeks a1 the Station, thai he mighl 
make observations in the field and participate in the collecting. 
But for repeated and long continued work on both living and 
preserved material such as is implied by the range of problems 
contemplated, gratuitous service of this sort cannot be counted 
on. And why should it be expected or asked ! 

So with the other lines of research; a chemisl could easily be 
Pound wlm would be glad to examine water samples that mighl 
be sent to his own laboratory; and geologists there would be who 
under like conditions, from their geological interest, would will- 
ingly reporl on bottom deposits. But where is the chemist, or 
physicist, or geologist, or hydrographer, who would be willing, 

Or could afford, to Undertake such systematic studies, lamely of 

necessity at the Station, as would meet the biological require- 
ments? There is really little more ground for assuming that a 
chemist's scientific interests should be sufficient to induce him to 
enter upon such a task, than that they should be sufficient to 
induce him to do the chemical work at a sugar factory, or a gas 

In short, the only way by which such a survey can be carried 
on with any considerable measure of success is through an organ- 
ized, salaried staff. This, of course, means a large and contin- 
uous expenditure. But the size of the expenditure would be 
fortunately lessened by the circumstance that while the staff 
would be in the aggregate rather large, only a portion, and in the 
main a comparatively small portion, of the time of each member 
would be demanded. In most cases occasional visits to the Sta- 

Vol. 2] Hitter. — Introduction. xi 

tion for brief periods, with most of the work done elsewhere, 
would suffice. So the chief and more permanent members could 
as well as not be persons in regular positions and with regular 
incomes in other institutions. Furthermore, the investigations 
are of such a nature that students in the stage of advancement 
of candidacy for the doctor's degree in a University could, by 
working under the guidance of those more experienced, be of 
much service. 

8.— Present Status, as to Ways and Means. 

An organization incorporated under the laws of Cali- 
fornia, known as the Marine Biological Association of San Diego, 
is at present the structural foundation upon which the survey 
rests: but the Association is prospectively a department of the 
University of California. Provision is made in the articles of 
incorporation that under specified conditions and at the expiration 
of a certain period, all the holdings and undertakings of the Asso- 
ciation shall pass automatically and wholly into the hands and 
under the control of the Board of Regents of the University. 

In the meantime, the University's part in the undertaking 
consists in a measure of cooperation through a committee of 
Regents, with the Managing Board of the Association, in the 
conduct of the business affairs of the survey; in the fact that the 
Director and most of the Scientific Staff are members of the Uni- 
versity Faculty; in the granting to the Association permission 
to take to the Station each year a considerable amount of labora- 
tory equipment and numerous library books; and finally, in 
publishing at its own expense the results of the investigations. 

The assets of the Association at present are : a laboratory 
building at La Jolla, neither large nor of elaborate construction, 
but serviceable for most of the work now in progress ; a schooner 
of nineteen tons register, with auxiliary power, and fitted with 
hoisting engine and gear; collecting apparatus; the nucleus of a 
library; and a definite guaranteed income for three years from 
July 1, 190-4. 

The present officers of the Association are : 

H. IT. Peters, President. 

Dr. Fred Baker, Vice-president. 

xii University of California Publications. [Zoology 

H. P. Wood, Secretary. 

Julius Wangenheim, Treasurer. 

Wm. E. Ritter, Scientific Director. 

E. W. Scripps and Miss Ellen Scripps. members of the Board 
of Directors. 

B. M. Davis, Resident Naturalist, 1904-05. 

Manuel Cabral, Collector. 

The permanent members of the staff since 1901 have been 
Wm. B. Ritter, Ph.D., Professor of Zoology in the University; 
C. A. Kofoid, Ph.D., Associate Professor of Histology and Em- 
bryology; H. B. Torrey, Ph.D., Assistant Professor of Zoology. 
In addition the following, all connected in sonic capacity with 
the University, have been members for Longer or shorter times 
on assignment to particular pieces of work, and for the most 
pari on the pay roll: W. J. Raymond, B.S., Assistant Professor 
of Physics; F. W. Bancroft. Ph.D., Instructor in Physiology; 
Alice Robertson, Ph.D., Assistant in Zoology; C. 0. Esterly, A..B., 
Assistant in Zoology; John F. Bovard, B.S., Assistant in Zool- 
ogy; Margaret Henderson, U.S.: II. M. Evans; L. II. Miller, 
M.S., Assistant in Zoology; Robert Williams. B.S. ; ami Effie -I. 

9.— Historical Note. 

Our work in this area did not begin with the San Diego 
Association, or even witli San DiegO as a base of operations. 
During six weeks of the summer of IS!):? a party of teachers and 
students from the Department of Zoology of the University of 
California, housed in a tent Laboratory at Avalon, Santa Catalina 
Island, made the first dip into these waters. Both the money and 
equipment for this piece of work were supplied by the Regents 
of the University. Another University party, with headquarters ai 
San Pedro, put in several week's of the summer of 1895. Nothing 
further of a formal character was attempted until 1891, though 
individual members of the department made repeated collecting- 
trips to San Pedro throughout the intervening period. All this 
served to prove the great richness in marine life, the advanta- 
geousness as a collecting place, of the San Pedro district. When, 
consequently, it was resolved, in 1901, to make on effort on the 

Vol. 2] Bitter. — Introduction. xiii 

basis of ideas that had beed taking- shape for several years — those, 
in a word, which now animate the undertaking- — San Pedro was 
believed to be the most favorable locus for whatever might be 
done. For this summer it was resolved to aim particularly at 
dredging operations in the shallow waters, made as thorough as 
the time and equipment would permit, with a reconnoissance to 
San Diego if passible. The University being unable to supply 
the money for this, a successful appeal was made to friends of 
the University and of science in Los Angeles and elsewhere. 
Funds to the amount of about $1,800 were secured, with which a 
large though open gasoline launch was hired and fitted for the 
work. She was kept going almost constantly from May 20 to 
August 6. While the dredging and trawling were the chief occu- 
pation, other lines of work were not wholly neglected, particu- 
larly plankton collecting and temperature taking. The proposed 
run to San Diego was made, and from the days devoted to the 
work there a good impression of the biological conditions of that 
region was obtained. 

For the work on shore on old bath house was rented and con- 
verted into a simple laboratory. The summer of 1902 was like- 
wise spent at San Pedro, but this year nothing was done at sea, 
attention being restricted to the littoral fauna. 

During both these seasons formal courses of instruction in 
Zoology were given as part of the regular University Summer 

Before the next summer the laboratory building and 
best collecting grounds within the small inner harbor at San 
Pedro had been destroyed by the harbor improvements being 
prosecuted there by the U. S. Government. Owing to this and 
to encouraging proposals for financial aid from San Diego, led 
by Dr. Fred Baker, and to the good impression made by the 
experiences there in 1901, it was resolved, in the early spring of 
1903, to move the base of operations to San Diego. During the 
years 1903 and 1901 the boat house at Coronado Beach, given and 
in part fitted up by the Coronado Beach Company, served as a 
laboratory building. 

The work at San Pedro was made possible largely through 
the interest and efforts of Mr. J. A. Graves, Mr. H. W. O'Mel- 

xiv University of California Publications. [Zoology 

veny, and Mr. Jacob Baruch of Los Angeles. The chief contrib- 
utors of money here were: Mr. Jacob Baruch, Mrs. Phoebe A. 
Hearst, Mr. J. A. Graves, Mr. H. W. O'Melveny, Mr. Wm. G. 
Kerchoff, Mr. Wm. B. Bowland, Mr. Van Xuys. The Los Angeles 
Terminal Railroad, Mrs. Margaret Pette, Mr. J. H. Shankland. 
Mr. John E. Plater, and Mr. Charles M. Wright. 

By far the largesl givers to the station since its removal to 
San Diego have been Mr. E. W. Scripps. Miramar; Miss Ellen B. 
Scripps, La Jolla, and Mi-. H. H. Peters. San Diego. In addition, 
the following have contributed substantially: Mr. Win. Clayton, 
for the Coronado Beach Company; Mrs. F. L. Keating. Mr. 
Henry W. Putnam, Mr. (J. W. Marston, and Hon. V. S. Grant. 

10.— Remarks on the Present Status of Marine Biology in General. 

Sil uated as our stal LOB is. mi a biologically almost unknown 
pari of a little known ocean, our firs! concern, chronologically, 
must be with Local conditions and problems. The meagerness of 
knowledge, not only of the fauna ami flora, hut also of the ocean- 
ography of the eastern part of the North Pacific can hardly he 
realized except by the few specialists whose studies have led 
them into immediate contact with it. Sir John Murray, the 
acknowledged prince of oceanographers, when the science is 
regarded as pertaining to the earth as a whole, has recently 
pointed out the urgenl need of further exploration of the Pacific 
from about 150 W. Line-, to the American coast. Our informa- 
tion about the most general facts concerning the currents, for 
instance, is wholly inadequate to constitute a foundation for 
investigations on distribution of organisms. And as to zoology, 
there are whole groups of prime importance for any of the wider 
questions of marine biology, like i he dinoflagellata, the radio- 
laria, and the chaetognatha, about which there is hardly a 
recorded observation. Even the better studied groups, like the 
fishes, the molliisks. and the crustaceans, when ecologically re- 
garded have been hardly more than glanced at. 

But, hemmed in as we are and for a long time must be by 
the limitations of meager local knowledge, we yet venture to look 
somewhat beyond these limits to see where the general idea consti- 
tuting the underpinning of our enterprise stands with reference 

Vol. 2] Bitter. — Introduction. xv 

to the present state of this domain of science ; and in what particu- 
lars, if any, Nature has given us opportunities to be of special 
use in advancing it. Looking over the whole domain, one sees 
that while certain geographical regions, like the Mediterranean, 
the North and Baltic Seas, the environs of the British Islands, 
and, to a less extent, the North American half of the Atlantic, 
have been cultivated, intensely even, in certain particulars, when 
attention is directed to large problems rather than to space areas, 
the thoroughly subjugated portions are exceedingly small. 

Let one go to the Bay of Naples, for instance, perhaps the 
best cultivated locality, and make inquiry about the ecology of 
the most familiar species found there, and see how far from satis- 
factory an answer can be obtained. In the realm of pelagic life, 
no one would contend that the great expeditions of the last half- 
century, even that of the Challenger, of the Blake, and the recent 
more concentrated and betted equipped German Plankton and 
Valdivia Expeditions, and those of the Albatross, have done more 
than to effect a reconnoissance of the field. The most general ques- 
tions of seasonal, vertical, and areal distribution are still topics of 
widest divergence of view, and of lively discussion ; and it is 
obvious that this diversity is in large measure due to the 
mere matter of dearth of readily ascertainable information. 
Beyond the most general truth, important is this is, that the 
bottom of the sea, even in its deeper parts, is inhabited by ani- 
mals, how immediately one comes against a blank wall when he 
begins to ask questions about this life. How abundant is it? 
Does it actually reach into the profoundest depths'? Are we to 
suppose it to be uniformly distributed over the entire ocean floor, 
modified only by local conditions, or as belonging essentially to 
the continental margins, with only an advance guard of strag- 
glers, so to speak, reaching to the localities farthest removed 
from any land? How long, geologically, have the truly abyssal 
depths been inhabited, and when and how did they become inhab- 
itable? From what source did the immigrants to these regions 
come? If from the littoral realms, has there been a general 
movement of approximately equal importance from all shores, 
or has it been chiefly from the polar regions ? What is the signifi- 
cance, biologically, of the continental shelf? What of Murray's 
"mud line"? 

xvi University of California Pubications. [Zoology 

When viewing this whole field of knowledge, and the means 
and methods of investigation, one must he struck by the prevail- 
ing uniformity and inadequacy of the existing' marine stations 
for coping with the situation. This inadequacy is most manifest 
in two particulars; first, in the well nigh complete absence of 
endowment, which is essential for the assurance of that certainty 
and regularity of income by which alone continuous and long 
continued, definitely planned investigations can be prosecuted; 
and secondly, by the fundamental idea on which nearly all these 
institutions are based. They have been and are, with few excep- 
tions, primarily resorts for individual investigators of specific 
biological problems, and not for systematically attacking the 
problems of marine biology proper. 

I would wish to guard myself without fail against being 
understood as passing adverse criticism upon these laboratories. 
They were, most of them, brought into existence by an obvious. 

immediate, and pressing i 1. This they have met, and are 

meeting, magnificently. No oilier instrumentality has contrib- 
uted so Largely to the promotion of genera] biology. The partic- 
ular need which gave them birth was not. however, that here 
considered. Only in the course of natural progress has this need 
come pressingly into existence. We are able now to formulate 
more definitely than has hitherto been possible, the problems in 
this tield. and to see more clearly what methods and instruments 
must be used in their prosecution. 

We are in position to appreciate, tor example, as never before 
the importance of knowing the complete life-histories of animals. 
We are becoming ever more impressed as knowledge advances, 
with the truth that no segmenl of the phenomena presented by 
an animal, morphological or physiological, is fully understood 
until it is regarded in the light of the entire life career of that 
animal. We are likewise in position to see as never before what 
must be done to attain to this fullness of knowledge. We must, in 
the first place, learn by observation all the facts of the life- 
history of the animal. In the second place, we must make use 
at every point possible of a combination of observation and 
experimentation for the interpretation of these facts. 

VoL - -] Hitter. — Introduction. xvii 

I verily believe the value of the experimental and statistical 
methods now so largely nsed in biology is not fully appreciated 
even by some of the most skilful and constant experimenters 
themselves, nor will it be until these methods are better coordi- 
nated with observation in Nature. The problems of animal 
migration, to be specific, we now know depend largely, at least 
so far as the simpler aquatic forms are concerned, on purely 
physiological reactions to temperature, light, sex relations, food, 
etc. ; and we are already in possession of important clues to the 
way these questions must be studied ; but we must leam, through 
careful and extended observation of the animals in nature, just 
what it is we have to interpret. Need for a kind of marine bio- 
logical research not specially felt a few years ago is now becom- 
ing urgent. 

The laboratory of the Liverpool Marine Biology Committee 
on the Isle of Man, under the directorship of Professor W. A. 
Herdman, and the proposals of the International Commission 
for the Investigation of the Sea, are distinctly in the direction of 
what the future must have for carrying on such researches. 

The portions of Nature unsubjugated by science are vast — 
it almost seems as though they grow vaster the longer we work 
at them; and one of the great questions science has ever before 
her is that of making such effort as she is able to put forth count 
for the most. One way of doing this is by giving good heed, not 
alone to the talents and tastes of workers, and money endow- 
ments, but as well to the opportunities held out by Nature herself. 

The conditions placed by Nature before us mark unmistak- 
ably the road we ought to take. 


Vol. 2, Nos. 1 and 2, pp. 1-51, PI. 1. ' December 21, 1904 











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The San Diego Region, as here defined, extends along the 
coast from LaJolla, ten miles north of Pt. Loma, to the Coronado 
Is., fifteen miles south of the same point. In climate, topography 
and faunal characters, it is in many respects similar to the 
region about San Pedro, Cal., which lies some ninety miles to the 
northwest. The hydroids about San Pedro have been considered 
in a former paper (:02). The present work is concerned only 
with the local hydroids, 1 of which there are 42 known species, 
8 being new ; and all are represented in the collections of the 
University of California or the Marine Biological -Association 
of San Diego. The accompanying table will show their recog- 
nized distribution. 

No attempt has been made to give complete specific synony- 
mies. The plan adopted gives (1) the original name of the spe- 
ck's, (2) the permanent name if some change has been necessi- 
tated, and (3) all synonyms in papers dealing with Pacific Coast 

1 With the single exception of S. pedrensis. 



University of California Publications. 

[ Zoology 

Table of Distribution. 


o ir 






A 1 * 






■a = 
































1. Bimeria gracilis Chirk 

2. Bougainvillia glorietta 


3. Eudendrium rameum (Pallas) 

4. ramosum (Linn. | 

5. Bp 


('). Hydractinia californica . 


7. Corymorpha palmaT. 

8. Tubularia eroeea (Ag.) 



9. Haleeium annulatumT 

Id. kofoidi T. 

1 1 . Washington] Nutting 
» lampanulariidae 

12. Campanularia edwardsi Nutting 
everta Clark 


liincksi Alder 

vdluliilis (Linn. ).. 
Obelia corona 

dichotoma (Linn.) .... 

geniculate ( Linn. ) .. 
Clytia bakeri 

eompressa I Clark i 



Calycella syringa (Linn.) 

25. Lafoea duniosa (Flem.) 


Sertularella group 

26. Sertularella halecina T 

27. pedrensis 
tenella (Alder) 
turgida (Trask) 

Dynamena group 

31. Dynamena cornicina McC. . 

32. St- rt nl aria desmoidis T 

33. Eurcata Trask 

Thuiaria group 

34. Sertularia filicula E. & S 


35. Aglaophenia diegensis T 

inconspicua T 

pluma (Linn.) 

struthionides (M.) 

Diplocheilus allmani 

Plumularia alieia T. 

megaloeephala A... 
plumularioides(C. ) 
setacea (Ellis) 










Totals 7 






















11 211 42 14 






X E 

+ NE 







Vol. 2] Torrey. — Hydroids of the San Diego Region. 


1. No true hydrotheeae or gonangia Gymnoblastea 2 

1. True hydrotheeae and gonangia present.. .., Calyptoblastea 13 

2. Tentacles in proximal and distal sets: hydranth abruptly set off 

from stem Pennariidae 3 

2. Tentacles in one circlet, filiform (5 

3. Solitary nutritive polyp, rooted in sand; perisarc rudimentary 

Corymorpha 4 

3. Several nutritive polyps from common stolon; perisarc well 

developed; gonophores fixed Tubularia 5 

4. Not more than 30 proximal tentacles; gonophores without tentacles 

C. palma (p. 9) 

5. About 25 proximal tentacles; gonophores with (3-10 flattened pro- 

cesses T. crocea (p. 10) 

(3. Colony encrusting; hydranths clavate ; spiral zooids; spines on 

hydrorhiza Hydractiniidae, Hydractinia 7 

6. Colony branching 8 

7. Sterile hydranths with (3-10 tentacles; blastystyles with knoblike 

tentacles H. californica (p. 19) 

8. Proboscis trumpet shaped; hydranth not fusiform 

Eudendriidae, Eudendrium 9 

8. Proboscis conical ; hydranth fusiform Bougainvilliidae 10 

9. Stem and principle branches polysiphonic E. rameum (p. 8) 

9. Stem polysiphonic at base only ; branches simple 

E. ramosum (p. 8) 

10. Gonophores fixed sporosacs Bimeria 11 

10. Gonophores liberated as medusae, with four pairs of tentacles 

Bougainvillia 12 

11. Stems simple, only partially annulated; hydranth with 10-12 

tentacles B. gracilis (p. (3) 

12. Stem and branches without annulae, often twined about each other.. 

B. glorietta (p. 7) 

13. Nematophores absent 14 

13. Nematophores present; hydrotheeae sessile, on one side of stem or 

branches only Plumulariidae 38 

14. Hydrotheeae with definite basal septa; gonangia not aggregated 15 

14. Hydrotheeae without definite basal septa, tubular; smooth margined ; 

gonangia aggregated Lafoeidae 28 

1."). Hydrotheeae much wider than deep Haleciidae l(i 

15. Hydrotheeae campanulate, usually deeper than wide, never adnate 

nor immersed Campanulariidae 19 

15. Hydrotheeae sessile, adnate or immersed, in 2 to (5 rows.Sertulariidae 30 

16. Gonangia with sporosacs Halecium 17 

17. Stem not fascicled 18 

17. Stem fascicled; ultimate branches geniculate.. H. washingtoni (p. 11) 

18. Annulated; hydrathecae about half as deep as broad, margin everted 

H. annulatum (p. 10) 

4 University of California Publications. [Zoology 

18. Not annulated; hydrothecae shallow, margin not everted 

H. kofoidi (p. 11) 

L9. Operculum of numerous small triangular pieces; unbranched ; 
hydrothecae tubular Calycella, C. syringa (p. 20) 

19. No operculum 20 

20. G-onophores sessile, sporosacs Campanularia 21 

20. Gonophores free medusae 23 

21. Stims branched, non- fascicled; hydrotheca with 12-14 marginal 

teeth C. edwardsi (p. 11) 

21. Stems unbranched 22 

22. Hydrotheca! margin with 12-15 crenations; gonangium compressed, 

small aperture C. everta (p. 12) 

22. Hydrotheca with 10-12 Bquare-topped teeth C. hincksi (p. 13) 

22. Hydrotheca deep, with 11-12 short, sharp teeth....C. hesperia (p. 12) 

22. Hydrotheca small, tubular, with !» low blunl teeth 

C. volubilis (p. 13) 

23. Medusae liberated with at least 16 tentacles Obelia 24 

23. Medusae liberated with 4 tentacles Clytia 26 

24. Hydrotheca] margin smooth 25 

24. Hydrotheca] margin bidentate O. corona (p. 14) 

25. Branching irregular; stem non -geniculate O. dichotoma (p. 15) 

25. Usually unbranched, except for pedicels; latter on shoulder pro- 

cesses of geniculate stem O. geniculata (p. 15) 

26. Stems simple 21 

26. Stems ami branches polysiphonic ; colonies Large and bushy, 

hydrotheca] margin with 12-15 teeth C. universitatis (p. 19) 

27. Branched; hydrotheca delicate, with about 14 carinate teeth 

C. hendersoni (p. 18) 

27. Unbranched; hydrotheca usually with thick wall, crenate margin: 

gonangium compressed, wide aperture C. compressa (p. 17) 

27. No branches except hydranth pedicels; hitter usually with but 

single annulus; hydrotheca] margin smooth C. bakeri (p. 16) 

28. Hydrothecae straight Lafoea 29 

29. Hydrothecae sessile : stem creeping or erect and polysiphonic 

L. dumosa (p. 20) 

30. Hydrothecae in two rows 31 

31. Hydrothecae alternate, one to an internode ..... Sertularella group 32 
31. Hydrothecae opposite, one pair to an internode Dynamena group 35 

31. Hydrothecae subopposite to alternate, more than two to an internode 

Thuiaria group 37 

32. Hydrotheca] margin dentate 33 

32. Hydrotheca] margin entire, everted, hydrotheca tubular; branches 

and gonangia rising within hydrothecae S. halecina (p. 26) 

33. Three marginal teeth 34 

:V.'y. Four marginal teeth: hydrothecae and stems annulated 

S. tenella (p. 28) 

34. Hydrothecae corrugated; internodes long, slender; gonangia 

spinulose S. pedrensis (p. 27) 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 5 

34. Hydrothecae smooth; internodes moderate; gonangia with high 
narrow transverse corrugations S. tricuspidata (p. 28) 

34. Hydrothecae smooth or roughened; internodes stout, geniculate; 

gonangia with distal spines or annulae or both..S. turgida (p. 29) 

35. Margin of hydrotheca without teeth S. desmoides (p. 30) 

35. Margin of hydrotheca with two teeth 36 

36. Teeth prominent; mouth of hydrotheca large S. furcata (p. 31) 

36. Teeth moderate, mouth of hydrotheca small D. cornicina (p. 30) 

37. Aperture of hydrotheca round; hydrotheca abruptly narrowed 

distally S. filicula (p. 32) 

38. Nematophores fixed by a broad base 39 

38. Nematophores fixed by a narrow base 44 

39. Stem simple; hydrotheca with dentate margin, no anterior intra- 

thecal ridge; corbulae without hydrothecae at bases of leaflets 

Aglaophenia 40 

39. Stem simple; hydrotheca with anterior intrathecal ridge; single 

sapracalycine sarcostyle ; no corbulae Diplocheilus 43 

40. Hydrotheca with 11 irregular teeth A. struthionides (p. 35) 

40. Hydrotheca with 9 teeth 41 

41. Median tooth recurved 42 

41. Median tooth not recurved A. pluma (p. 34) 

42. Hydrocladial nodes well marked: mesial nematophore reaches mouth 

of hydrotheca A. inconspicua (p. 34) 

42. Hydrocladial nodes weak ; mesial nematophore does not reach mouth 

of hydrotheca A. diegensis (p. 33) 

43. Margin of hydi'otheca smooth; mesial nematophore very short and 

broad D. allmani (p. 36) 

44. Hydroeladia pinnately disposed on erect stem without thorny pro- 

cesses or nematophorous branehlets; no corbulae Plumularia 45 

45. Hydrocladial internodes thecate and non-thecate 46 

45. Hydrocladial internodes thecate only P. plumularioides (p. 38) 

46. Nodal septa transverse 47 

46. Nodal septa alternately transverse and oblique; thecate internodes 

twice as long as non-thecate P. alicia (p. 37) 

47. Hydroeladia borne on short processes of stem, always alternate 

P. setacea (p. 39) 

47. Hydroeladia borne on long processes of stem, very slender, opposite 
distally P. megalocephala (p. 37) 

University of California Publications. 




(Jen. Bimeria, Wright, 1859. 
1. Bimeria gracilis Clark. 

Bimeria gracilis, dark, 1876a, p. 252, pi. :'>s. fi^. 3. 

T/iophosome. Stems rising from creeping hydrorhiza to height of 
Lin id :',u mm., with numerous short branches. Eydranths borne alternately 
mi latter, on moderate pedicels; 9-13 rather stout tentacles. Stem usually 
smooth, occasionally wrinkled. Pedicels with 2 to 5 more or less indistincl 
annulae at the base. Perisarc opaque, extending to bases of tentacles. 

Gonosome. Sporosacs ovate, borne on branches singly or in pairs. 
Pedicel short ami smooth; spadix branched. 


Fig. 1. — Bimeria r/rticilis. (Jonopliores. 

Distribution. Dredged near the mouth of San Diego Bay. in 
3 fathoms. July. 1903; La Jolla, at low water. July. 1903. San 
Diego I Clark). 

There can be little doubt that this is Clark's species. The 
perisarc was covered with minute adhering particles. Male 
sporosacs are slightly smaller than female. 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 

Gen. Bougainvillia, Lesson, 1836. 
2. Bougainvillia glorietta, n. sp. 

Trophosome. Stems branched, rising from a creeping hydrorhiza in 
clusters to the height of 20 to 30 cm. Stems, or stem and branches fre- 
quently twine about each other. Perisarc smooth, without aunulae, occa- 
sionally wavy, unusually adhesive, covered with particles of dirt and diat- 
oms, reaching bases of tentacles. Terminal hydranths largest, with 20 
1o 25 tentacles carried in two or three irregular whorls, the outermost 
shortest. Tentacles highly contractile, held stiffly when at rest. 

Gonosome. Gonophores each on a short pedicel, in groups of two or 
three on branches or hydranth stalks. 

Fig. 2. — Bougainvillia glorietta. 
Hydranth from below. 

Fig. 3. — B. glorietta. 
Hydranth with tentacles 
partly retracted; edge of 
perisarc visible. 

Distribution. San Diego Bay, Cal. 

This is the second species of the genus to be found on this 
coast. The first was collected in San Francisco Bay by A. Agas- 
siz ('65) and referred to B. mertensi. It was taken again by 
myself in Oakland Harbor, Cal., and is mentioned in a footnote 
on p. 1 of my former paper ( :02). The present species resembles 
B. superciliaris, yet differs in lacking annulae in the perisarc 
and in the twining habit of stem and branches. Medusae with 
4 pairs of tentacles, 4 simple mouth processes and 8 ocelli were 
still within the perisarc, July 5, 1903. 

8 University of California Publications. [Zoology 

Gen. Eudendrium, Ehrenberg, 1834. 

3. Eudendrium rameum (Pallas). 

Tubularia ramea, Pallas, 1766, p. 83. 

Eudendrium rameum, Johnston, 1847, p. 45, pi. 5, figs. 1, 2. 

Eudendrium rameum, Torvcy, 1902, p. 33. 
Trophosome. "Hydrocaulus profusely branched, attaining a height 
of from three to six inches, fascicled in the main stem and principal 
branches; main stem attaining a thickness of more than a quarter of an 
inch, and as well as the principal branches, very irregularly ramified; 
branches alternately losing their fasciculation and then consisting of single 
capillary tubes, which may continue to branch before the emission of the 
ultimates or hydranth-bearing ramuli, which are regularly alternate in their 
disposition; perisarc rigid, occasionally marked with nearly obsolete ampl- 
iations on the smaller branches. Bydranths with about twenty tentacles, 
frequently atrophied Li the male after the production of gonophores. ' ' 

Gonosome. Male sporosacs two-chambered, borne upon the body of 
the hydranth in a verticil immediately below the tentacles; female sporo- 
sacs oval, scattered on the hydrocaulus for some distance below the 

hydranth." (Allman. 71.) 

Distribution. Mouth of San Diego Bay, between tides (June 
26, 1903: no gonosome); San Pedro. C;\\.. on float at surface 
(Dec, 1901; do gonosome). Mediterranean, Norway, (It. Britain 
(Allman). Jan Mayen (Markt.) Helgoland (Hartlaub). Green- 
Land (Levinsen). Northern Asia (Thompson). 

4. Eudendrium ramosum (Linn). 

Tubularia ramosa, Linnaeus, 1767, p. L302. 

Eudendrium ramosum, Ehrenberg, L834, p. 296. 

Eudendrium ramosum, Torrey, 1902, p. 34. 
Trophosome. Hydrocaulus much branched, fascicled at the base 
and attaining a height of four inches or more; primary ramifications irreg- 
ular, after which the branches become regularly alternate and mostly dis- 
tichous in their arrangement, giving off all along their length, from their 
upper or distal sides, short, usually simple ramuli. which support the 
hydranths on their summits; perisarc firm, annulated at the origin of the 
branches, or even along the entire length of the smaller branches. 
Hydranths usually with about twenty tentacles. 

Gonosome. Male sporosacs two chambered, borne on body of hydranth 
below tentacles which often atrophy. Female sporosacs piriform, scattered 
on body of hydranth and stalk immediately below it. 

Distribution. Mouth of San Diego Bay, in 3 fathoms; Pacific 
Grove, Cal. Gt, Britain (Allman). Jan Mayen, Adria, Rovigno 
(Markt.). Helgoland (Hartlaub). Greenland (Beteneourt). 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 9 

The San Diego material consists of three stem fragments 
which are provisionally placed in this species. The definition is 
Allman's ('71), with a few slight verbal changes. 

5. Eudendrium sp. 

A fragment of a colony, consisting of hydrorhiza and a few 
very short stems, was taken at Point Loma, June 27, 1903. The 
perisarc is sparsely and wavily annulated. Female blastostyles 
with tentacles. 

Fam. Hydractiniidae. 

Gen. Hydractinia, Van Beneden, 1844. 

6. Hydractinia calif ornica, n.sp. 

Trophosomc. Sterile hydranths 2 to 2.5 mm. long in extension, with 
(5 to 10 tentacles, usually in 2 recognizable whorls ; proboscis domed to con- 
ical. Spines .5 to .9 mm. long, conical, often with truncated tops and 
irregular protuberances; with about 10 longitudinal dentate ridges. 

Gonosome. Sporosacs, with 1 or 2 eggs in female, borne in clusters 
of 2 to 10 or more about half way from the base of the blastostyle. Latter 
with 5 to 10 knobdike clusters of nematocysts representing tentacles; 1 to 
1.3 mm. long. 

Colors. Perisarc deep brown, fleshy parts white. 

Distribution. Off San Diego, in 50 fathoms. Covering the 
shells of Dentalium polygonum inhabited by hermit crab. 

This species is very closely allied with H. echinata (Flem.) of 
Europe and TI. polyclina Ag. of the eastern United States, which 
are themselves almost indistinguishable. It appears to differ 
from them in its much smaller number of tentacles. The latter 
arise in threes, fours, or fives, or irregularly; there is no single 
typical method. 

Fam. Pennariidae. 

Gen. Corymorpha, Sars, 1835. 

7. Corymorpha palma. 

Corymorpha palma, Torrey, 1902, p. 37; 1902a, p. 987. 
Trophosome. Stems each 6 to 14 cm. long, rooted in sand by a 
dense tangle of filamentous processes, and covered by perisarc proximally 

10 University of California Publications. [Zoology 

lor one third or one fourth of its length; thickest near proximal end, taper- 
ing gradually into a narrow neck which supports the hydranth. Latter 
with 18 to 30 proximal tentacles in one whorl, with a span of about 2.5 
cm.; distal tentacles more than twice as numerous, more or less irregularly 
placed around the mouth in several whorls. 

Gonosome. Gonophores medusoid, permanently fixed to peduncles 
springing from the base of the proboscis just within the whorl of proximal 
tentacles, each with a ring and 4 radial canals, ami a manubrium at leas! 
twice as long as the hell, without a mouth; tentacles wanting; velum may 
be present or absent. 

Distribution. San Diego and s.-m Pedro, Cal., throughout the 
year, between tides, on sand tints. Eggs laid in May, June, July. 

(ifi). Tubularia, Linnaeus, 1767. 
s. Tubularia crocea (Ag.). 

Parypha crocea, L. Agassi/, 1862, 111. p. 249, pis. 23, 23a, figs. 1-7. 
Parypha microcephalia, A. Agassiz, 1865, p. 195. 

Tubularia crocea (Ag.), Allmaii, ls71. p. 41<i. 
Tubularia elegans, Clark, 1876a, p. 253, pi. :'. y . fig. 2. 
Tubularia crocea, Torrey, 1902, p. 4::. pi. -1. Pigs. 22, 23. 
Trophosome. Colony usually a bushy mass of stems, tangled below, 
which may be In .-in. long and may occasionally branch. Eydranths with 

not more than 25 proximal tentacles. 

Gonosome. Gonophores borae in pendulous clusters en peduncles 
arising between proximal and distal tentacles; with <i t" 10 flattened pro- 
cesses, varying in size, more prominent in female, sometimes hardly visible 
in male. 

Distribution. S;m Diego liny. S;m Pedro Harbor, and San 
Francisco Bay, Cal. Eastern United Stntcs (Agassiz). 


Fnin. Haleciidae. 

den. Halecium, Oken, 1816. 

!). Halecium annulatum. 

Halecium annulatum, Torrey, 1902, p. 49, pi. 3, figs. 30, 31. 

Trophosome. Stems rising from a creeping hydrorhiza to a height 
of 7 mm.; the longei have - regularly alternating branches. Stem and 
branches more or less regularly annulated throughout. Hydrothecae may 
be half as deep as broad; margin everted. Sessile hydrothecae alternately 
on either side of stem or branch; peduncles arising within these carry 
other hydrothecae which may also give rise to other peduncles. 

Gonosome. Female gonangia broadly ovate, excessively compressed, 
with terminal aperture. Single gonophore with numerous ova, surrounded 
by blastostylar processes reaching to gonangial wall. 

a^ol. 2] Torrey. — Hydroids of the San Diego Region. 11 

Distribution. Coronado Is., Mexico (July. 1903), and Coro- 
nado, Cal. (July, 1901). Growing on seaweed. 

10. Halecium kofoidi. 

Halecium kofoidi, Torrey, 1902, p. 49, pi. 3, figs. 32, 33. 

Trophosome. Stems rising from creeping hyclrorhiza, branching 
irregularly; largest colonies with thick trunk may reach 5 mm. in height. 
Branches arise just below hydrothecae; divided obliquely into internodes 
of approximately equal length. Each internode usually bears on a distal 
shoulder process a sessile hydrotheca which does not reach beyond the 
distnl node. Within this hydrotheca another may arise, and another within 
the latter, both on short stalks somewhat constricted at the base and bent 
slightly away from the stem. Hydrothecal wall especially thick. 

Gonosome. Male gonangia long, oval, smooth, 3 or 4 times as long 
as broad; may be waved proximally; small terminal aperture. 

Distribution. Coronado Is., Mexico (July 25, 1903, on kelp 
at surface); Pt. Loma ; mouth of San Diego Bay, 5 fathoms; 
( Jatalina Is., 42 fathoms. 

11. Halecium washingtoni Nutting. 

Halecvum geniculatum, Nutting, 1899, p. 744, pi. 63, fig. 1. 

Halecium washingtoni, Nutting, 1901, p. 789. 

Halecium mittingi, Torrey, 1902, p. 50. 
Trophosome. Colony fascicled, branching in much the same plane, 
branches approximately alternate; non-fascicled branches more or less regu- 
larly annulated at their bases, with long internodes which are arranged in 
zigzags distally. Hydrothecae at the distal end of each internode, singly 
or in pairs, margins everted. Hydranths with 16 to 24 tentacles. 

Gonosome. "Gonangia borne singly in the axils of the branches 
and branchlets, regularly ovoid in one view, barnacle-shaped in the other; 
aperture large, terminal" (Nutting, '99). 

Distribution. Pt. Loma, July, 1901. Puget Sound (Nut- 

Fam. Campanulariidae. 

Gen. Campanularia, Lamarck, 1816. 

12. Campanularia edwardsi Nutting. 

Campanularia edwardsi, Nutting, 1901, p. 346, fig. 28. 

Trophosome. "Colony attaining a height of over an inch, branch- 
ing somewhat irregularly, but with a distinct tendency to send off pedicels 
from the main stem in sub-opposite pairs. Stems, branches and pedicels 
exceedingly long and slender, with the annulation confined to the proximal 
portions, except the few just below the hydrothecae. Hydrothecae very large, 
deeply eampanulate, with 12 to 14 exceedingly sharp, slender teeth. 
Hydranth with about 28 tentacles. 

Gonosome. "Unknown" (Nutting). 

12 University of California Publications. [Zoology 

Distribution. Mouth of San Diego Bay, in :? fathoms. Woods 
Hole. Mass., between tides (Nutting). 

There is much variation in the size and proportions of the 
hydrothecae, correlated apparently with mode of growth. The 
stems arc usually densely clustered. Some of the larger stems. 
however, grow more freely above the rest. These longer stems 
hear the longest hydrothecae, which may be .92 mm. long. On 
the crowded si ems. the hydrothecae may he no longer than .(10 
mm. Short scattered stems produce the largest hydrothecae of 
all. The diameter does not vary with the length, as the following 
measurements in mm. show: ,625x.26; .<>7 x .Ml : .68x.45; 
,70x.45; .72x39; .92x.45. As a rule, however, the diameter 
is less than half the length. The thecae growing in the clusters 
are relatively broader. 

The gonosome was qo1 present, July 15, 1903. 

13. Campanularia everta Clark. 

Campanularia everta, Clark, L876o, p. 253, pi. :'.!». tig. 4. 
Campanularia everta, Torrey, L902, p. 51, pi. 4. figs. :'>~>-:;7. 
Tropho8ome. Pedicels <>t' variable length, smooth, wavy or Lrregii 
Larly annulated, arising directly from a creeping hydrorhiza; spherical 
annula immediately below each hydrotheca. Wall of latter varies greatly, 
from excessive thickness to thin; straight or convex in profile; margin 
usually crenate. 

Gonosome. Gonangia somewhat compressed, ovate, with small round 
terminal aperture. Aerocysts may lie present in female, which are some- 

w hat Larger than male. 

Distribution. San Diego, low water to 24 fathoms; Catalina 
I., 42 fathoms; Pacific Grove, Cal. San Diego Clark). 

Gonosome present. June 26, 1903. 

Transitions between all the forms of hydrothecae have been 
traced in the same colony. ('. everta can he distinguished from 
CI yt in compressa by the gonosome; the gonangia have a much 
narrower aperture and the gonophores are fixed sporosacs. 

14. Campanularia hesperia, n. sp. 

Trophosome. Stems simple, unbranehed, from a creeping hydrorhiza, 
terminating in hydranths; with about 10 rings at base, 2 to 4 just below 
hydrothecae, and usually 3 or 4 others in the distal half of the stem. Hydro- 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 13 

thecae less than half as broad as long (.5x.22 mm.; .6 x .25 mm.), with 
11 or 12 short, sharp marginal teeth. Hydraxrth with 22 to 24 tentacles. 
Gonosome absent, July 3 3, 1903. 

Fig. 4. — Campanularia hesperia. Hydrotheea and pedicel. 

Distribution. La Jolla, Cal., between tides, on the tests of 

This species closely resembles Clytia cylindrica Ag. in habit 
and skeletal features, but the latter species has but 16 tentacles, 
and the absence of the gonosome makes it desirable to distinguish 
between the two for the present. 

15. Campanularia hincksi Alder. 

Campanularia Hi/nlcsii, Alder, 1857, p. 127. 
Campanularia Einlsii, Hineks, 1868, p. 162, pi. 24, fig. 3. 
Campanularia hinlisi, Torrey, 1902, p. 53. 
Trophosome. Pedicels arise directly from hydrorhiza. Hydrothecae 
large and deep, with 11 to 14 flat-topped teeth which may have rounded 
corners or be slightly hollowed out above. Wall very thin, with delicate 
longitudinal lines from the margin between the teeth. 

Gonosome. Gonangia much elongated, slightly tapering distalward, 
to truncate end ; 10 to 18 wavy annulations. 

Distribution. Mouth of San Diego Bay, in 3 fathoms; off 
San Diego in 40 to 75 fathoms. Newport, R. I. (Nutting). Brit- 
ish coasts, from 10 to 20 fathoms to deep water (Hincks). 

Gonosome present, June 29, 1903. 

16. Campanularia volubilis (Linn.). 

Sertularia volubilis, Linnaeus, 1767, p. 1311. 
Campanularia volubilis, Alder, 1856, p. 358, pi. 13, fig. 7. 
Campanularia volubilis, Hartlaub, 1901, p. 357. 
Campanularia volubilis, Torrey, 1902, p. 54, pi. 5, fig. 48. 
Trophosome. Pedicels long, annulated, springing from hydrorhiza. 
Hydrothecae small, broadly tubular; margin with 9 to 10 short blunt teeth, 
frequently reduplicated. 

Gonosome. Gonangia smooth, flask-shaped, somewhat compressed, 
with a long narrow neck and small circular aperture. 

Distribution. San Diego, shore rocks; San Pedro. Cal., 9 
fathoms; Tomales Bay, Cal., shore rocks. Near Vancouver. B. C. 
(Hartlaub). Gulf of St. Lawrence (Packard). Massachusetts 


University of California Publications. 

[ Zoology 

(Agassiz). Iceland, British coasts (Hincks). Norway (Sars). 
Helgoland (Hartlaub). White Sea (Mereschkowsky) . 

No gonosome in the San Diego specimens, June 26, 1903. 

Gen. Obelia, P. et L., 1809. 
17. Obelia corona, n. sp. 

Tropho8ome. Colonies very low; stems simple, short, slightly flex- 
uous, from a creeping Btolon, with 3-6 annulae distal to each pedicel. 
Hydranths long, narrow, tapering, margin with 8-10 teeth, each with two 
sharp cusps, pedicel short, completely annulated, with 2-4 annulae. 
Hydranth with about -J4 tentacles. 

Gonosome. Gonangia about 3 times as long as broad, with wide 
aperture; pedicel slender, with 2-4 annulae. Numerous medusae, largest 
with 24 tentacles. 

Dimensions. Bydrotheca: ,44x.20; .4:; x .IS mm. 

Gonangium: .76x.20 mm. (including pedicel). 


Pig. 5. — Obelia corona. 

Pig. 6. — 0. corona. Gonangia. 

Distribution. San Diego Bay, on piles tinder wharves at low 
tide. July 15, 1904. Creeping over sponges. 

Hydranths and gonangia are frequently borne on pedicels 

springing directly from the stolon. Stems and stolons transform 
with readiness into each other in the colonies at hand, owing 
doubtless to the many opportunities offered by life on a growing- 
sponge for variations in the contact stimulus. 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 15 

18. Obelia dichotoma (Linn). 

Sertularia dichotoma, Linnaeus, 1767, p. 1312. 

Obelia dichotoma, Hincks, 1868, p. 156, pi. 28, fig. 1. 

Obelia dichotoma, Calkins, 1899, p. 356, pi. 3, fig. 16. 

Obelia dichotoma, Torrey, 1902, p. 57. 
Trophosome. ' ' Stem filiform, slender, nearly straight, irregularly 
branched, ringed above the origin of the branches, of a deep horn color: 
branches suberect, often very long, and more or less ramified, ringed at 
intervals, a single calycle in the axils; hydrothecae alternate, broadly com- 
panulate and deep, polyhedral above, each side corresponding with a very 
slight sinuation of the margin, borne on ringed pedicels, which vary in 
length from 4 or 5 to as many as 16 rings. ' ' 

Gonosome. ' ' Gonotheeae axillary, slender, smooth, widening from 
the base upwards, and terminating above in a raised, somewhat conical 
aperture." (Hincks). Medusae liberated with 16 tentacles (Hincks), 20 
to 24 (Southern California specimens). 

Distribution. San Diego ; San Pedro, Cal. Puget Sound 
(Calkins). Alaska (Nutting). Eastern United States (Nut- 
ting). Helgoland (Schulze). N. Asia (Thompson). 

It is possible that the California species is the stock which 
produces an undescribecl medusa of the genus Obelia that is very- 
abundant in the neighborhood of San Diego. If this prove to be 
the case, the hydroid, though identical with 0. dichotoma as 
regards the trophosome, will become a new species. 

19. Obelia geniculata (Linn.). 

Sertularia geniculata, Linnaeus, 1767, p. 1312. 

Obelia geniculata, Allman, 1864, p. 372. 

Obelia geniculata, Torrey, 1902, p. 58. 
Trophosome. "Stem zigzag, sometimes sparingly branched, jointed 
at each of the flexures, and thickened immediately below them, so as to 
form a series of projections or rests, from which the pedicels arise, hydro- 
thecae somewhat obconical, rather short, the length slightly exceeding the 
width, with a plain margin, borne on short, annulated stalks (rings (4-6), 
which are suberect and taper slightly upwards." ' 

Gonosome. "Gonotheeae axillary, urnshaped, attached by a short 
ringed stalk (3-4 rings)." (Hincks.) Medusae at time of liberation with 
24 tentacles. 

Distribution. Coronado, Cal., at surface; Catalina I., 42 fath- 
oms; San Francisco, between tides. Eastern United States 
(Agassiz, Nutting). Europe (Hincks). White sea (Meresch- 
kowsky). New Zealand (Hartlaub). 

Gonosome present, July 1, 1903. 


University of California Publications. 


The geniculation varies in different parts of the colony and 
may be absent in some regions. The colonies from Coronado are 
unusually low, and branched. Otherwise they are not distinguish- 
able from the typical forms. 

Gen. Clytia, Lamouroux, 181G. 

20. Clytia bakeri, n. 


Trophosome. Stems clustered, about i!<* mm. long, without branches. 
Each stem free, for 2 to 5 mm., from pedicels of hydrothecae, which then 
follow each other alternately in quick succession; closely annulated at base, 
annulae increasing gradually in length, ultimately becoming internodes of 

stem, st, 'in Internodes usually '■'■ to I times as long as broad, each bearing 
:i pedicel mi a shoulder process from distal end. Hydrothecae small, con- 
ical, without marginal teeth; pedicel consisting usually of bul one annuls 
almost as long as broad. 

Gonosome. Gonangia lone, narrow, with bottlenose apertures, taper 
ing gradually to short peduncles; Kerne usually in pairs at the bases of 
hydrothecal pedicels. Sporosacs abundant, 12 to 20. 

Fig. 7. — Clytia bakeri. 
Proximal portion of 
stem, showing charac- 
teristic aiinulation. 

Fig. 8.— C. bakeri. 

Portion of stem in distal 
half, with three typical 
hydrothecae, one of 
which is sessile. 

Pig. 9. — C. bakeri. 

Proximal region of 
stem with gonangia 
and hydrotheca with 
exceptionally long 

Vol. 2] Torreij. — Hydroids of the San Diego Region. 17 

Distribution. Pacific Beach, in the surf, attached in tufts to 
the posterior region of both valves of the beach clam Donax, a 
most unusual situation for a hydroid, which may acount for the 
irregular, damaged margins of the hydrothecae. The beach was 
literally covered with the colonies, Jan. 2, 1904. Mouth of San 
Diego Bay, under similar circumstances, July, 1904. 

There is considerable variation in the length of the annul ae 
or internodes on the stem and the annulation of the pedicels. 
Occasionally a pedicel has several annulae (fig. 9), but this is the 
case only in the lower portion of the stem — e.g., the pedicel 
referred to is the lowest in the colony. More often the hydro- 
thecae are sessile ; this condition is found in the distal portion 
of the colony. As a general rule, the annulation disapears from 
base to tip. 

The dimensions of the hydrothecae in four average specimens, 
are as follows, length first, in mm. : .69 x .45 ; .68 x .50 ; .50 x .52 ; 
.48 x .37. 

Dimensions of gonangia : .98 x .33 ; 1.14 x .28. 

This species is named in honor of that tireless friend of edu- 
cation and public spirited citizen of San Diego, Dr. Fred Baker. 

21. Clytia compressa (Clark). 

Campanularia compressa, Clark, 1876o, p. 214, pi. 8, figs. 5, 6. 

Clytia compressa, Nutting, 1901, p. 170, pi. 17, figs. 3, 4. 

Cytia compressa, Torrey, 1902, p. 58, pi. 6, fig. 49. 
Trophosome. As in C. everta. 

Gonosome. Gonangia compressed, broadly ovate, with truncated 
top and large aperture. 

Distribution. San Diego, 5 fathoms; San Pedro, CaL, 3 
fathoms. Oca, Al. (Nutting), Shumagin Is., Al., 6-20 fathoms, 
on Lamina ria (Clark). 
Gonosome present. May 23 and July 13, 1901. 


University of California Publications. 


22. Clytia hendersoni, n. sp. 

Trophosome. Colonies branching, 3 to 5 cm. high. Internodes of 
the stem flexuous, with a pronounced knee at the base of each, and run- 
ning parallel with hydranth pedicels for nearly half their length; above 
each knee, 3 to 8 annulae. Pedicels completely annulated, with 6-14 annulae. 
Hydrothecae large, deep, 1.00-1.2 mm. long by .40.-.60 mm. in diameter, 
tapering gradually, with very thin and easily collapsible walls, bordered by 
about 14 very sharp, keeled teeth. 

Gonosome. Gonangia with wide mouths, widest in distal half, 
tapering, 3 times as long as broad, with wavy contours but not annulated. 
Pedicels short, with 3 or 4 annulae. Usually 3 or 4 medusae in each gonan- 
gium, each with 4 tentacles and without gonads. 

Fig. 10. — Clytia hendersoni. Stem 
with hydrothecae. 

hendersoni. Stem 

Fig. 11.— C. 
with gonangia. 

Distribution. San Diego Bay, 3 fathoms, July 15, 1903. 
Growing rather thickly on sponges. The flexuous character of 
the stem is more pronounced distally. 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 


This species is named for Miss Margaret Henderson as a slight 
mark of appreciation of her efficient assistance in the prepara- 
tion of this paper. 

23. Clytia universitatis, n. sp. 

Campanularia, denticulate,, Torrey, 1902, p. 51, pi. 4, fig. 34. 

Trophosome. Stem long, branching irregularly, forming bushy tufts 
often exceeding 200 mm. in length. Stem and branches polysiphonie. 
Hydranth pedicels long, almost completely annulated; hydrothecae deep, 
tapering, with 12-15 marginal teeth ; hydranths with about 28 tentacles. 

Gonosome. Gonangia borne on stem, branches or hydranth pedicels, 
less than 3 times as long as broad, with short pedicel which may or may 
not contain a single annulus; with wide aperture. Contour somewhat irreg- 
ular, occasionally I or 2 annulations distally. Medusae numerous, oldest 
with 4 tentacles. 

Dimensions. Hydrothecae, in mm.: .70x.37; .78x.40; .81x.39; 
.83x42. Gonangium: .95x39; 1.00 x .41. 

Fig. 12. — Clytia uni- 
versitatis. Hydrotheca. 

Fig. 13. — C. nniver 
sitatis. Gonangium. 

Distribution. San Diego Bay, on piles of wharves at low 
tide, July 15, 1904; San Pedro Bay, Cal., Dec, 1901. 

The species bears a general resemblance to Campanularia ver- 
licillata. The pedicils of the hydranths, however, are not arranged 
in verticils, and the gonophores are not sporosacs. Young colo- 
nies taken in San Pedro Harbor, December, 1901, were previ- 
ously identified with C. denticulata Clark, though certain differ- 
ences were noted and the immaturity of the colonies prevented 
an accurate determination. 

•_'() University of California Publications. r ZoOLuGY 

Gen. Calycella, Hincks. 1861. 
24. Calycella syringa (Linn). 

Sertularia syringa, Linnaeus, 1767, p. 1311. 

Calicella syringa, Hincks, 1861, p. 294. 

Calycella .syringa, Hincks, 1868, p. 206, pi. 39, fig. 2. 

Calycella syringa, Calkins, 1899, p. 358, pi. 4, fig. 20. 

Calycella syringa, Clark. Is7(i, p. 217, pi. 12, fig. 25. 

Calycella syringa, Hartlaub, 1901, p. 358. 

Calycella syringa, Nutting, 1899, p. 741; 1901, p. 176. 

Calycella syringa, Torrey, 1902, p. 59, pi. 6, fig. 50. 
Trophosome. Pedicels borne on stolon, shorter than hydrothecae. 
Margins of hydrothecae frequently reduplicated. 

Gonosome. Gonangia on stolon, with acrocysts at maturity; ovate, 

Distribution. San Diego Bay, 1 to 5 fathoms. Puget Sound; 
Berg Inlet .-Hid Kadiak, Al. (Nutting). CoaJ Harbor and Shum- 
agin Is.. Al. (Clark). White Sen (Mereschkowsky). Northern 

Asi;i i 'I'll pson). Kara Sea (Berg). British Coasts, [celand 

in loo fathoms (Hincks). Greenland (Levinsen). Helgoland 
i Hartlaub). 

No 'j: )soiiic in San Diego colonies, July 15, 1903. 

Pam: Lafoeidae. 

(i<ii. Lafoea, Lamouroux, L821. 

25. Lafoea dumosa (Flem.). 

Sertularia dumosa, Fleming, 1828, p. s -' ; . 

La [ova dumosa, Sars, l s 'i'-'. 

Lafoea dumosa. Hincks. L868, p. 200, pL 41, fig. 1. 

Lafoea dumosa, Clark, 1876, p. 216, pi. 12. fig. 215. 

Lafoea dumosa, Nutting, L899, p. 747. pi. 64. 

Lafoea dumosa. Torrey, 1902, p. 59. 
Trophosome. Stem simple ami creeping or fascicled ami erect. 
II yilrothecae strong, oarrowed toward the base, with little or no pedicels. 

Gonosome. Gonangia columnar, with bottle necks, crowded together 
in encrusting masses. 

Distribution. San Diego Bay. in 6 fathoms: Port Orchard, 
Puget Sound. California Coast (Clark). Alaska (Clark, Nut- 
ting). White Sea (Mereschkowsky). New En eland Coast (Ver- 
rill. Nutting). West Indies. 450 fathoms (Allman). British 
Coasts (Hincks). Spitzbergen (Marktanner-T.). North Cape, 
Norway (Sars). Helgoland (Hartlaub). 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 21 

Both the erect and creeping forms were found at San Diego. 
Overgrown with G. hincksi. No gonosome, June 29, July 15, 

Fain. Sertulariidae. 

Anyone who has had occasion to work among the Sertulariidae 
will admire the masterly way in which Nutting ( :04) has dealt 
with the perplexing questions of classification in that family. I 
am not yet prepared, however, to abandon Schneider's plan of 
segregating the species into typical groups which shall take the 
places of genera. These groups do not necessarily give their 
names to the species which they include. Thus they discourage 
the growth of synonyms, offer no awkward bars to the free pass- 
age of any species from one group to nearer relatives, and at the 
same time lessen the confusion which the present unsettled state 
of opinion regarding the relationships of existing species tends to 

When it is not easy to define groups clearly, owing either to 
the uncertain values of diagnostic characters or to baffling tran- 
sitional forms, it is plainly desirable to have as few groups as 
convenience will permit. Marktanner-Turneresteher ('90) dis- 
tinguished 18 genera. Nutting has reduced this unusual total 
to 12 ; but that number, I am convinced, is still too large. The 
distinction between Thuiaria and Abietinaria hardly seems of 
enough service to overweigh the practical difficulties which it 
invites ; and although Nutting has put forth every effort to make 
it useful, he has only succeeded in distinguishing the genera by 
relying now on one, now on another combination of characters, 
not an attractive makeshift. But however desirable or undesir- 
able this procedure may be, I make no reservations in condemn- 
ing Allman's genera Thecocladium and Synthecium as Nutting- 
has defined them. According to Allman, Thecocladium is distin- 
guishable by the intrathecal origin of its branches, Synthecium 
by the intrathecal origin of its gonangia. The justice of my 
objections ( :02, pp. 61, 62) to genera founded on single charac- 
ters of such a sort is admitted by Nutting, who then attempts 
to strengthen both genera, but particularly Synthecium, which 
alone ocurs in American waters, by supporting them on combi- 
nations of characters. Synthecium is accordingly based upon a 

■2~2 University of California Publications. [Zoolooy 

combination of strictly opposite branches, smooth ma ruined hy- 
drothecae, absence of opercula, as well as the intrathecal origin of 
the gonangia. 

Analysis, however, dues not reveal the strength which is 
claimed for this structure. In the first place, combinations are 
of little value unless the characters selected for combination 
vary independently of each other, which is obviously not true 
of margin and operculum, as Nutting is aware. In the second 
place, it is well known that at some stage in the development of 
all hydroids. a perisarcaJ membrane blocks the exit of the 
hydranth from the hydrotheca, and that this membrane becomes 
the one-, two-, three-, or four-parted adult operculum, according 
In the character of the margin, or may be wanting altogether. 
Among sertularians with smooth round margins, it is often deli- 
cate, and is commonly lost. In Sertularella formosa, according 
to Nutting, it is usually wanting, but occasionally appears as a 
"thin membrane stretched like a drumhead across the aperture." 
In Sertularella hartlaubi, according to the same authority, the 
operculum is "in some cases an adeauline flap, in others appar- 
ently an irregularly ruptured mbrane stretched straigb.1 

across the aperture like ,-i drumhead." Sertularella halecina (a 
Synthecium according to Nutting) possesses ;i thin drumhead 
operculum before the hydranth emerges for the firsl time, bul 
is non-operculate in the adult. Such tacts only Lead inevitably 
to the conclusion of llartlaub > :(l(). p. Si that the absence of an 
operculum is of no taxonomic consequence. In the third place. 
Nutting does not appeal 1 to insist that Synthecium shall exhibit 
the opposite branching which his definition demands, when he 
places S. halecina in that genus. It is possible to assume that 
he was heedless of the mode of branching in this species, but 
this assumption is hardly applicable to the alternately branching 
Synthecium alti rnans Allman. It is more probable that Nut- 
ting included S. halecina in spite of its branching. Yet in thus 
escaping the responsibility of removing it from Synthecium to 
Sertularella or to an entirely new genus, he abandons opposite 
branching as a distinguishing mark of Synthecium. It would 
appear, then, that there are but two instead of four characters 
on whose association the genus is really based: the smooth round 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 23 

margin of the hydrothecae and the intrathecal origin of the 

Are these characters of equal rank? Evidently Nutting pre- 
fers the latter, since he does not hesitate to align such a smooth 
round margined species as 8. formosa with the typical dentate 
operculate species of Sertularella. Here I can by no means agree 
with his judgment, but must take the ground which I previously 
occupied ( :02, p. 62). The mere location of gonangia, whether 
arising within or outside of hydrothecae, cannot to my mind be 
of such taxonomic importance as the striking differences of the 
trophosomes in species like S. halecina and S. tubitheca; though 
I am far from refusing its aid as a means of distinguishing 

The general grounds which I formerly urged against Syn- 
thecium need not be repeated here. I am still in hearty accord 
with the position then taken, but a review of actual conditions 
in 8. halecina may prove more convincing. In the majority of 
cases, the gonophores of this species arise within hydrothecae. 
Occasionally, however, they are borne directly on the stolon 
(fig. 14), as in Dynamena cornicina. When such a difference in 
the position of the gonangia exists in different species, it has 
been held by Allman and Nutting to indicate generic distinction. 
For instance, Sertularella integritheca, with smooth round mar- 
gined hydrothecae and extrathecal gonangia, is said to be gene- 
rically distinct from such a form as Synthecium alternans All- 
man, with intrathecal gonangia but otherwise similar to S. integ- 
ritheca. Occasional conditions such as the intrathecal origin of 
gonangia in species in which the gonangia are usually extra- 
thecal, are held by the same authors to be abnormalities which 
may be disregarded in classification. The extrathecal origin of 
the gonangia of 8. halecina 1 being but occasional, would natu- 
rally find a place in the same category. These views do not 
appear to me to be justifiable, for the occasional presence of 
extrathecal gonangia must lessen the importance of the usual 
condition, and should not be disregarded for this very reason. 

1 If it be objected that the extrathecal gonangia in S. halecina arise on 
the stolon instead of the stem, it may be remembered that stem and stolon 
are fundamentally the same structure, and transform into each other with 
the utmost readiness. 


University of California Publications. 

r Zoology 

According to Nutting- ( :04, p. 42), "it occurs not infrequently 
in several widery different forms among the Sertularidae that a 
gonangium will occasionally have its origin within the lumen 
of the hydrotheca, although these species normally produce gon- 
angia in the ordinary position." The very fact that what is 
typical of S. halecina is atypical of most other sertularians, 
though not infrequent, and what is occasional in S. halecina is 
usually typical of the others, only leads to a rejection of the 
view which lays more than specific importance on the difference 
in the typical position of gonangia in different species. 

Pig. 1 1. — Serlulart lla 
halecina. Portion of 
stolon with gonan- 
gium (s) : base of one 
stem, with intrathecal 
gonangium { i ). 

Pig. 15; — S. halecina. 
Portion of stem show- 
ing origin nt' branch 
just below hydrotheca 


Pig. 16. — S. halecina. 

Portion of colony. Show- 
ing :m extrathecal branch 


If the branches of S. halecina be now considered, it will be 
seen that they, as well as the gonangia, emerge typically from 
hydrotheeae. In this respect, then, 8. halecina is a Thecocla- 
dium. Yet this character is not invariable. Occasionally 
branches arise independently of hydrotheeae. Four such cases 
are shown in figs. 15, 16, 17, 18. In figs. 15. 17, 18, the branches 
arise on the bulging stem just beneath hydrotheeae, a familiar 
origin of branches in the Seiiulariidae. In fig. 16, the branch 
is not associated with a hydrotheca in any way. All these cases 
(except fig. 17) were found on the same colony: and it is inter- 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 

esting that of three successive branches from a short section of 
the same stem (fig. 18), the origin of the first is typical of The- 
cocladinm, the origin of the second and third is typical of Ser- 
tularella. Without denying the usefulness of the usual manner 

Fig. 17.— S. halecina. A younger stem than Fig. 18.— S. halecina.— Stem with three 
is shown in fig. 18 with two extrathecal branches two of which are extrathecal, 
branches. the third intrathecal, in origin. The 

stern is old, with damaged hydrothecae. 

of branching, or any other typical character, in defining species, 
the facts which have just been enumerated hardly supply the 
idea of stability which is commonly associated with the concep- 
tion of a genus. 

26 University of California Publications. [Zoology 

Sertularella halecina is at once a typical Syntkeciwm in the 
character of the hydrothecal margin and the intrathecal origin 
of the gonangia, a typical Thecocladium in the intrathecal origin 
of the branches, a typical St rtulart ll<i in the manner of the origin 
of both branches and gonangia. Allman refers to the intrathecal 
origin of two branches in Synthecium campylocarpum as an 
abnormality, and Nutting speaks similarly of Hie intrathecal 
origin of the only two branches which were presenl in the mate- 
rial from which I described Sertularella dentifera. Both cases 
may be abnormal, in the sense of unusual; but in the light of 
conditions in S. halecina, is it wise to dismiss them forthwith as 
taxonomieally insignificant .' Which are the abnormal, the insig- 
nificant characters in S. h<il< <in<i .' I must confess my inability 
to decide. Until such a decision be reached. I do not think better 
can be done than to consider the species a member of the Sertu- 
larella group, in which it was originally placed. 

Sertularella group. 
2G. Sertularella halecina. 

Sertularella halecina, Torrey, 1m<il!, y. in, p. u, fig. 55. 

Trophosome. steins from a creeping stolon rise to height of 30 mm., 
with few branches which originate either within hydrothecae or just below 
them. Nodal < li \ isi i his faint, often wanting. Sydrotheeae adnate at base 
(inly, cylindrical, with slighl swelling en Lower Bide of base, wide aperture 
with smooth, everted rim. 

Gonosome. Gonangia arise within hydrothecae or from stolon, long, 
tubular; single tubular gonophore. 

Distribution. San Diego Bay, 3 to 12 fat I is: growing on 

kelp and among bryozoa. Gonosome present, -Inly. 1901: dune. 
July. 1903. 

My reasons for withdrawing this species from Synthecium, 
where Nutting placed it. have been given above. I am not yet 
prepared to consider it identical with S. cylindrica Bale ('Nut- 
ting, :04), because there is no record of the method of origin 
of branches and gonangia in the latter, and the ma ruins of the 
hydrothecae are not so distinctly or characteristically everted. 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 


27. Sertularella pedrensis, n. sp. 

Sertularella conica, Torrey, 1902, p. 60. 

Trophosome. Stems from creeping stolon, longest 35 mm., with 
occasional branches; stems and branches divided into slender internodes 
of variable length. Hydrothecae distant, borne at distal ends of inter- 
nodes, free for two thirds their length, narrowing to tridentate apertures, 
which are often reduplicated, with tripartite opercula; each hydroheca 
with 3 to 6 transverse rugae which are stronger on adeauline side. 

Gonosome. Gonangia ovate, covered thickly and completely with 
slender spines. 

Figs. 19, 20. — Sertularia pedrensis. Hydrothecae. 

Fig. 21. — S. pedrensis. 

Distribution. San Pedro, Cal. 

The trophosome of this species so closely resembles Allman's 
descriptions and figures of S. conica, that I formerly identified it 
with the latter. Nutting ( :04) has since pointed out that the 
hydrothecae of S. conica have four marginal teeth, a fact which 
at once distinguishes the two species. The recent discovery of 
two gonangia on the San Pedro colonies affords an unmistak- 
able diagnostic character. 

28 University of California Publications. [Zoology 

28. Sertularella tenella Alder. 

Sertularia tenella, Alder, 1856, p. 357, pi. 13, figs. 3-6. 

Sertularella tenella, Alder, 1857, p. 113. 

Serularella tenella, Hincks, 1868, p. 242, pi. 47, fig. 3. 

Sertularella tenella, Hartlaub, 1901, p. 360, pi. 21, figs. 12, 20, 21. 

Sertularella tenella, Torrey, 1902, p. 64. 
Trophosorae. "Zoophyte minute; stems short, slender, simple or 
slightly branched, zigzagged and jointed and twisted above each calycle; 
hydrothecae rather distant, elongate, barrel shaped, finely ribbed across, 
the aperture erect, patent, squared, 4-toothed, and closed by a four sided 
operculum. ' ' 

Gonosome. "Gonothecae ovate, slender, ringed transversely, produced 
above into a short, tubular orifice" (Hincks). 

Distribution. La Jolla, Cal., between tides; San Diego, 9 
fathoms. Bare I. (Hartlaub). Gt. Britain, between tides to 
deep water (Hincks). New Zealand (Hartlaub). 

Growing on rocks and Fums. No gonosome, July 16, 1901, 
July 13, 1903. Longest stem, 4 mm.; length of hydrotheca, .4 to 
.5 i ., breadth, .25 mm. 

29. Sertularella tricuspidata (Alder). 

Sertularia tricuspidata, Aider, 1856, p. 356, pi. K'>. figs, 1, 2. 

Sertularella tricuspidata, Hincks, lsiis. p. l':'.!», pi. 47, fig. 1. 

Sertularella tricuspidata, Clark, L876, p. 224, pi. 12, figs. 26, 27. 

Sertularella tricuspidata, Nutting, 1899, p. 741. 

Sertularella tricuspidata, Hartlaub. 1901, p. 359. 

Sertularella tricuspidata, Nutting, 1901, p. l v ."'. 

Sertularella hespfiria, Torrey, 1902, p. 63, pi. 7, tiys. 57, 58. 
Trophosome. "Colony a matted mass of shoots and twigs some 
times attaining a height of 5 or 6 inches. Stem not fascicled, slender, 
divided into Lnternodes, each of which bears a hydrotheca or a branch with 
its axillary hydrotheca. Branches irregularly alternate, often branching 
profusely either alternately or dichotomously, divided into regular inter- 
nodes each of which bears a hydrotheca. some of the nodes being double 
and oblique, which gives a twisted appearance to the branch. Hydro- 
thecae distant, small, cylindrical, without corrugations, the distal half 
or more being free; margin with three strong, equal and equidistant teeth." 
Gonosome. "Gonangia borne profusely on the main stem and 
branches, large, oblong-ovate, marked throughout with very prominent 
compressed annular ridges, the uppermost of which forms a bowl-shaped 
structure from the center of which arises the tubular neck which ends in 
a slightly everted margin and round aperture. ' ' 

Distribution. San Diego Bay. 1 to 9 fathoms. "Abundant 
throughout the north polar and north temperate regions of the 
world" (Nutting, :04). 

Vol. 2] Torrey. — Hydroids of the Son Diego Region. 


30. Sertularella turgida (Trask). 

Sertularia turgida, Trask, 1854, p. 113, pi. 4, fig. 1. 
Sertularella turgida, Clark, 1876«, p. 259, pi. 38, figs. 4, 5. 
Sertularella conica, Calkins, 1899, p. 359, pi. 4, fig. 22. 
Sertularella nodulosa, Calkins, 1899, p. 360, pi. 5, fig. 29. 
Sertularella turgida, Hartlaub, 1901, p. 360, pi. 21, figs. 5, 6. 
Sertularella turgida, Torrey, 1902, p. 64, pi. 7, figs. 59-62; pi. 8, 
figs. 63-69. 
Trophosome. Stems stout, from creeping stolon, about 30 mm. long, 
seldom branching; divided into short geniculate internodes. Hydrothecae 
large, stout, free for about half their length ; aperture large, with 3 
strong teeth. 

Gonosome. Gonangia large, ovate, distally spinose or annulated or 
both ; aperture small. 

Figs. 22, 23. — Sertularella turgida. Gonangia. 

Distribution. Pacific Coast, from Coronado Is. to 54° N. lat. 
Off Japan (Albatross hydrographic station 3775). Between 
tides to 204 fathoms. 

This is by far the most variable species on the coast, a char- 
acteristic to which reference was made in my former paper 
( :02, p. 65). Figs. 22 and 23 make more complete the transi- 
tions between forms of gonangia there figured. From a spiny 
type an annnlated type is reached through a spiny-annnlated 
condition. The hydrothecae are extremely variable, as regards 
shape, wrinkling and immersion. The internodes vary much in 
length and thickness. 

30 University of California Publications. [Zoology 

Dynamena group. 
31. Dynamena cornicina McCrady. 

Dynamena cornicina, McCrady, 1858, p. 204. 

Sertiilaria complexa, (lark, ls79, p. 245, pi. 4, figs. 26-8. 

Sertularia complexa, Bale, 1888, p. 769, pi. 18, figs. 1-4. 

Sertularia cornicina, Nutting, 1901, p. 359, fig. 56. 

Sertularia complexa, Nutting, 1901, p. 360, fig. 57. 

Sertularia cornicina. Nutting, 1904, p. 58, pi. 4, figs. 1-5. 
Trophosome. Stems snort, slender, unbranched, rising from a creep- 
ing stolon to height of 10 to 20 mm.; divided into regular internodes each 
with a pair of opposite hydrothecae distally. Eydrothecae tubular, annate 
in front for two thirds their length; margin with two teeth. 

Gonosome. Gonangia borne at base of stems, broadly ovate, annu- 
lated throughout, with broad aperture. 

Distribution. Coronado Is.. Cal., on seaweed at the surface. 

Charleston, S. C. (McCrady). W Is Bole, Mass. (Nutting). 

Pourlalcs Plateau (Nutting). Yucatan Coast (Clark). Aus- 
tralia (Bale). 

The colonics from the Coronado Islands were identical with 
S. complexa, though there were easy transitions to the typical 
trophosome of D. cornicina. I have followed Nutting in consid- 
ering the two species synoliy nious. 

1). cornicina is very close to 8. desmoides, Erom which it 
appears to differ in the absence of branches, the sharply toothed 
aperture, and the position of the nodal constrictions immedi- 
ately above rather than immediately below the hydrothecae. 

32. Sertularia desmoides. 

Sertularia desmoiMs, Torrey, 1902, p. 65, pi. 8, figs. 70--72. 
Sertularia desmoides, Nutting, L904, p. "><;. pi. •">. figs. 1-3. 

Trophosome. Stems from creeping stolon, rising to height of 30-50 
mm., branching sparely and irregularly. Internodes vary in length, but 
the portion distal to the hydrothecae is never longer than the rest of the 
internode. Two hydrothecae on the proximal portion of each internode, 
opposite and contiguous on one side of the stem for one half their length, 
bending sharply outward in distal half and narrowing to a smooth or some- 
what bilabiate operculate aperture. 

Gonosome. Gonangia sessile, ovate, half as broad as long, with a 
wavy outline and broad round aperture. 

Distribution. San Diego, 1-25 fathoms; San Clemente 1., 42 
fathoms: San Pedro. Cal., 13 fathoms. Albatross station 2939, 
lat. N. 33° 36', long. AY. 118° 09' 30", 27 fathoms (Nutting). 

Gonosome present, July. 1901. June 27, 1903. Both robust 
and attenuated varieties were obtained. 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 31 

33. Sertularia furcata Trask. 

Sertularia furcata, Trask, 1854, p. 312, pi. 5, fig. 2. 

Sertularia furcata, Agassiz, 1865, p. 145. 

Sertularia furcata, Clark, 1876a, p. 258, pi. 39, fig. 3. 

Sertularia furcata, Torrey, 1902, p. 66, pi. 8, figs. 73-5. 
Trophosome. Stems short, unbranched, rising from a creeping 
stolon to height of 10-15 mm.; divided into short internodes, each with 
a pair of hydrothecae opposite and in contact on one side of the stem for 
half their length. Two strong marginal teeth and a large aperture. 

Gonosome. Gonangia broadly ovate, compressed, with moderate 
terminal aperture. 

Distribution. San Diego Bay, 5 fathoms; Coronado Is., Mex., 
18-24 fathoms; San Pedro, Cal., 9 fathoms; San Francisco, shore 
rocks. Farallone Is., Cal. (Trask). Santa Barbara and Santa 
Cruz, Cal. (Clark). 

Nutting ( :04) has identified this species with the Dynamena 
pulchella of d'Orbigny from Patagonia, quoting Clark's descrip- 
tion of S. furcata, however. I am unable to follow him because 
his reproduction of d'Orbigny 's figures does not show the con- 
tact of the members of each pair of hydrothecae, which is a 
marked character of the species, the internodes are longer and 
more slender than the constantly short internodes of S. furcata, 
and the two species are widely separated geographically as well. 
It is true that Clark's figure does not show the contact of the 
hydrothecae, but that is because he has probably drawn the 
reverse rather than the face of the stem. This view is supported 
by the position of the gonangia, which ordinarily occur on the 
face of the stem, and by the similarity of Clark's figure to fig. 
73 of my former paper, representing the reverse of one inter- 
node of the stem. 

Gonangia were present in colonies collected in November, 
1897, and July, 1901. By a .strange confusion which I came upon 
in the preparation of the present paper, I laid claim in my 
former paper to the discovery of the gonosome of the species, 
though it was well known to me that Trask, as well as Clark, 1 
had described and figured both trophosome and gonosome. 

'Nutting (:04), who calls attention to this blunder, has himself erred 
in ascribing the first complete description, including gonosome, to Clark. 
1 do not see Trask 's paper in Nutting's bibliography, and infer that he 
was not acquainted with it at first hand. 

32 University of California Publications. [Zoology 

Thuiaria group. 
34. Sertularia filicula E. & S. 

Sertularia filicula, Ellis and Solander, 1786, p. 57, pi. 6. 

Sertularia anguina, Trask, 1854, p. 112, pi. 5, fig. 1. 

Sertularia labrata, Murray, I860, p. 250, pi. 11, fig. 2. 

Sertularia filicula, Hincks, 1868, p. 264, pl.53, fig. 3. 

Sertularia anguina, Clark, 1876a, p. 255, pi. 40, figs. 1, 2. 

Sertularia anguina var. robust u, Chuk. Is7(w. p. 256, pi. -40, 
figs. 3, 4, 5. 

Sertularia filicula, Torrey, 1902, p. 68, pi. 9. fig. 80. 

Sertularia filicula, Nutting, 1904, p. 117, pi. 34, fig. 1. 
Trophosomt . Stems with alternating branches, [.innately disposed; 
divided into internodes each of which usually bears a branch and three 
hydrothecae, two sub-opposite, the third axillar. Branches may themselves 
branch: divided into unequal internodes, each bearing several hydrothecae, 
sub-opposite, in pairs. Hydrothecae flaskshaped, adnate for more than half 
their length, apertures small, round, opening upward. 

Gonosonn . Gronangia pearshaped, produced somewhal distally, end- 
ing with small round aperture. 

Distribution. Sac Diego, 1~)-'2~) fathoms; San Pedro, San 
Francisco, CaL, shore rocks. Monterey to Point Reyes, Cal. 
(Trask). Vancouver I. (Dawson). Alaska, 10 fathoms; San 
Miguel T., Cal. (Clark). Whit.- Sea (Mereschkowsky). New 
England coast (Verrill). Grand Manan, 20 fathoms (Stimp- 
son). Labrador (Packard"). Greenland (Levinson). North 
Atlantic (Bonnevie). Norway I Marktanner-Turneretscher) . 
British shores (Hincks . 

Nutting's treatmenl of Sertularia <ni<iiiiii<t Trask is unfor- 
tunate. To begin with, the figures of his Abietinaria anguina 
(Trask) are so far from typical of S. <ni<itii>i<i Trask. judging 
either from Trask 's figures, Clark's figures or ;ill of my own 
material, some of which was collected at the entrance of San 
Francisco Bay, where Trask also obtained the species, that I 
suspect they really represent a distinct species, lie says the 
specimens he has seen "are from Santa Barbara, Cal., and Ber- 
ing Sea, and they all agree well with Dr. Clark's description of 
Sertularia anguina var. robusta." His figures, however, resemble 
Clark's variety less than the typical S. anguina as show-n by 
Clark's own figures. Yet in his synonymy there appear S. 
anguina Trask, S. labrata Murray (a correct synonym) and S. 
anguina var. robusta Clark, but not the 8. anguina Trask of 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 33 

Clark, the figures of which the without the slightest shadow of 
doubt typical of Trask's species. There is as little question that 
my S. filicula E. & S. ( :04, p. 68, pi. 9, fig. 80) belongs with 
Trasks's and Clark's S. anguina; so I fail to see why Nutting 
placed it instead, though dubiously, with his Abietinaria filicula 
(Ellis and Solander). 

After expressing his inability to agree with me "in consid- 
ering this species identical with A. filicula," Nutting at once 
adds in a footnote : " It is possible that the name anguina should 
be retained for the var. robusta of Clark, which is apparently 
distinct." I take these statements to mean that if 8. anguina 
and 8. filicula prove to be identical, the var. robusta should 
remain under Trask's old name. Why Nutting is unable to see 
the identity of the two species he does not say and I am at a loss 
to discover. My reasons for uniting them lie in the fact that 
Trask's figure of 8. anguina, though crude and containing an 
error in showing 4 rather than 3 hydrothecae on the stem 
between the bases of successive branches, Murray's figure of 8. 
labrata, Clark's figures of 8. anguina, Hincks' figures of S. fili- 
cula and my own observations of both trophosome and gonosome 
refer unmistakably to the same species; and they agree with 
Nutting's figure (PI. 34, fig. 1) of Abietinaria filicula (Ellis and 
Solander) and not, curiously enough, with his figures of Abieti- 
naria anguina (Trask) on the same plate, figs. 5-7. They agree 
also with the var. robusta of Clark in all details save stoutness 
of the stem, a difference which is probably referable to differ- 
ences of environment, not heredity. 

Fam. Plumulariidae. 

Gen. Aglaophenia, Lamouroux, 1812. 

35. Aglaophenia diegensis. 

Aglaophenia diegensis, Torrey, 1902, p. 71, pi. 9, figs. 84-86. 
Trophosome. Stem 150 mm. long, with short internodes. Hydro- 
cladia alternating, one to an internode; divided into equal internodes by 
faint nodes which may be wanting. Hydrothecae each longer than diame- 
ter of aperture; 9 irregular marginal teeth, median tooth sharp and 
recurved, adjacent teeth longest, smallest teeth next the hydrocladmm. 
Mesial nematophore reaches level of hydrothecal aperture. Septal ridge 

34 University of California Publications. [Zoology 

just below supraealycine nematophores and one just above floor of hydro- 

Gonosome. Corbulae 3 to 4 times as long as broad, formed of 8-10 
pairs of alternating leaflets, 8 nematophores on anterior edge of all but 
first and last. One, rarely two hydrotheeae on anterior edge of all but 
first and last. One, rarely two hydrotheeae between corbnla and stem. 
Gonophores in two rows, about 12 in number. 

Distribution. San Diego Bay, 1-7 fathoms: False Bay. The 
corbulae on the False Bay colonies, collected in January, 1904. 
are longer than that, figured in my previous paper, collected in 
July, 1901; usually with ten Leaflets. 

86 Aglaophenia inconspicua. 

Aglaophenia inconspicua, Torrey, 1902, p. 73, pi. 9, figs. 87-89. 
Trophosome. Stems stout, in clusters, 35-40 mm. high; divided by 
anterb-posteriorly oblique aodes into internodes as broad as long. Hydro- 

eladia bora i same side of stem, alternate, one from each Lnternode, 3-4 

mm. long; divided transversely into equal internodes. A nematophore in 

the axil of each bydrocladium and two at its base in a line parallel with 
its axis. Eydrothecae deep, slightly compressed, free for aol more than 
one quarter their length; 9 marginal teeth, median tooth recurved, the 
next on each side longest. Intrathecal ridge extending obliquely upward 
from near base of theca. Two ridges on each Lnternode. Mesial nemato- 
phore reaching nearly or quite to the mouth of the theca. Supraealycine 
nematophores divergent, not reaching level of mouth of theca. 

Gonosome. Corbulae not more than twice as long as deep, arched, 
slightly compressed; formed of 4 to 6 leaflets, the longest with 10 nemato- 
phores on distal edge and occasionally one or two on proximal edge near 
tip. One thecate Lnternode between corbula and stem. Sporosacs 6-12. 

Distribution. S;m Diego. 5 fathoms: gonosome present, 
July, 1901. 

37. Aglaophenia pluma (Linn.). 

Sertularia pluma, Linnaeus, 1707, p. 1309. 

Aglaophenia pluma, Lamouroux, 1816, p. 170. 

Aglaophenia pluma, Hincks, 1868, p. 286, ,pl. 63, fig. 1. 

Aglaophenia pluma, Torrey, 1902, p. 73, pi. 10, figs. 90-91. 
Trophosome. Stems attaining height of 100 mm. or more, grace- 
fully rising from creeping stolon. Hydrocladia alternate, one to an inter- 
node. Hydrotheeae each with 9 teeth, median tooth not recurved. Mesial 
nematophore not reaching level of aperture. 

Gonosome. Corbulae of about 9 leaflets, arched; a single hydro- 
theca at base of each. 

Distribution. Off Coronado, Cal., on kelp. South Africa. 
Belgium, Mediterranean, Gt. Britain (Hincks). 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 35 

38. Aglaophenia struthionides (Murray). 

Plumularia struthionides, Murray, 1860, p. 251, pi. 12. fig. 2. 

Aglaophenia franciscana, A. Agassiz, 1865, p. 140. 

Aglaophenia struthionides, Clark, 1876«, p. 2G2, pi. 41, fig. 3. 

Aglaophenia struthionides, Torrey, 1902, p. 73. 
Trophosome. Stems long, strong, often attaining height of 150 mm., 
occasionally bearing stem-like branches; divided obliquely into short equal 
internodes each bearing a hydrocladium. Hydrothecae with flaring margin 
armed wih 11 irregular teeth: median tooth long, sharp, recurved; next on 
each side long and directed forward, next bent outward. Mesial nemato- 
phore usually reaches level of aperture. 

Gonosome. Oorbulae each formed of 8-13 pairs of leaflets; with 3, 
occasionally 2, hydrothecae at base. 

Distribution. Puget Sound to San Diego. This is the com- 
monest hydroid on the coast, frequently cast up on California 
beaches. Corbulae present, January, June, July. 

Gen. DiplocheilllS, Allman, 1883. 

Trophosome. All internodes thecate, each internode with an infra- 
calycine mesial nematophore not in contact with the hyclrothcea, and a 
supracalycine median sarcostyle without definite nematophore; each hydro- 
theca with anterior intrathecal ridge. 

Gonosome. Gonangia unprotected. 

Allman founded this genus on the following characters : a 
duplicating of the walls of the hydrothecae "forming an external 
calycine envelope," a shielddike mesial nematophore not adnate 
to the hydrotheca, and the absence of lateral nematophores. 
Bale ('93) has demonstrated that the hydrothecae of the single 
species (D. mirabilis Allman) for which the genus was created 
do not possess the double walls described by Allman, but are 
constructed after the fashion of the hydrothecae of Kirchenpau- 
eria producta Bale, with anterior intrathecal ridges which, from 
certain viewpoints, suggest a duplicature of the walls. Bale has 
also demonstrated the opening of a median sarcostyle above each 
hydrotheca, flanked by webs of perisarc between theca and inter- 
node which form a broad, non-typical nematophore. Allman 's 
definition has been modified to accord with these facts. 

All the trophosomal characters of D. mirabilis which have 
been mentioned are found also in K. producta Bale. The strik- 
ing similarity of the trophosomes of the two species leaves no 
doubt of their generic unity, in spite of the absence of the gono- 

36 University of California Publications. [Zoology 

some in f>. mirabilis. Bale, however, is certainly in error in 
finding in Jickeli's Kirchenpaueria the bond of union. Accord- 
ing to Jickeli's figure ( '83, pi. 28. fig. 27). the hydroid for which 
he erects the genus is an eleutheroplean plumularian — probably 
a Plumularia — with neniatophores broken away. The frequent 
absence of neniatophores ii species which characteristically pos- 
sess them and the absence of any other distinguishing characters 
remove the slender claims to priority over Diplocheilus which 
have been made for this inadequate genus. 

Diplocheilus allmani, n. sp. 

Halicornaria producta, Torrey, 1902, p. 7-". pi. 10, fig. ;».".. 

Trophosome. Colony with simple stem, divided obliquely into inter- 
nodes which var\ in length according to age. Bydrocladia alternate, each 
from a shoulder process projecting from the middle region of each inter 
node Each hydrocladium divided more or less obliquely into equal 
thecatc Lnternodes. Each hydrotheca somewhal compressed below, some- 
what flaring distally, with a broadly oval, smooth orifice; about as deep 
as long; free for one third of its length. Strong anterior intrathecal Sep 
turn about two thirds the length of the hydrotheca from the bottom, 
reaching about one third across it at widest point. Cauline nematophorea 
absent with the exception of single axillary aematophores. Mesial nemato- 
phore short, not reaching the base of hydrotheca. expanding into the 
form of a sickle shaped segmenl of a saucer with a diameter two thirds 
that of hydrotheca and embracing the internode for half its circumference. 
Single median supracalycine saicostyie, flanked by two webs of perisarc 
Stretched between theca and internode. forming a non-typical median 

Gonoscmn absent. 

Distribution. Pt. Loma, Cal., on seaweed and sponges. 

The differences which separate />. mirabilis Allman. D. pro- 
ducta (Bale) and I), allmani are slight. I), allmani, originally 
thought to be identical with /). producta, possesses hydrothecae 
with flaring rims and broadly ovate apertures instead of the 
compressed form and narrowed apertures of D. producta. It 
differs from l>. mi nihil is in the absence of all cauline neniato- 
phores save those in the axils of the bydrocladia, and the cauline 
internodes never bear more than one hydrocladium each. The 
immaturity and paucity of my material make it impossible to 
determine the real value of these differences. For the present, 
then, it seems desirable to distinguish the species. 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 37 

Gen. Plumularia, Lamarck, 1816. 
40. Plumularia alicia. 

Plumularia alicia, Torrey, 1902, p. 75, pi. 10, figs. 96, 97. 

Trophosome. Stems in clusters, slender, loosely branching, 7 to 13 
cm. high: divided transversely by faint nodes into short equal internodes. 
Hydrocladia alternate, one from distal end of each internode, and with 
4 to 7 hydrothecae; thecate and non-thecate internodes alternate, separated 
by nodal septa which are alternately transverse and oblique; thecate inter- 
nodes twice as long as non-thecate. Proximal and distal septal ridge in 
each internode. Hydrothecae free for at least half their length, adcauline 
contours, in profile, somewhat recurved. A single nematophore on each 
internode of stem on side opposite origin of hydrocladium; 2 nematophores 
in each axil ; each hydrocladial non-thecate internode with 1 nematophore ; 
thecate internodes with 1 mesial and 2 supracalycine nematophores. Per- 
isarc of stem thick and brown, of hydrocladia delicate and colorless. 

Gonosome. Male gonangia small, ovate, attached by very short 
peduncles between the nematophores in the axils of the stem or branches. 

Distribution. San Diego, 15 to 25 fathoms; Long Beach, 
Cal., 5 to 13 fathoms. Gonosome present, June and July, 1901. 

41. Plumularia megalocephala Allman. 

Plumularia megalocephala, Allman, 1877, p. 31, pi. 19, figs. 1, 2. 
Plumularia megalocephala, Nutting, 1900, p. 57, pi. 1, fig. 5. 

Trophosome. ' ' Hydrocaulus irregularly branched, not fascicled ; 
pinnae alternate, each borne close to the distal end of an internode of the 
stem, where it is supported on a long stout process of the internode; prox- 
imal internode of pinna short and destitute of hydrotheca; following inter- 
nodes longer, every alternate one carrying a hydrotheca, and slightly 
longer than the others. Hydrothecae small and shallow, each borne near 
the middle of its internode, and supporting a very large hydranth. Beside 
the supracalycine pair of nematophores, each hydrotheca-bearing internode 
carrying a single mesial nematophore at the proximal side of the hydro- 
theca; intervening internode carrying two mesial nematophores, and short 
basal internode carrying one. 

"Gonosome not known" (Allman). 

Distribution. Off San Diego, in 40-75 fathoms. Off Alli- 
gator Reef, 14 fathoms (Allman). Albatross Station 2669, lat. 
N. 31° 9', long. W. 79° 33', 352 fathoms (Nutting). 

"The internode intercalated between the hydrotheca-bearing 
internodes was sometimes present, sometimes absent, and was of 
variable length. The internodes of the stem carry two nemato- 
phores placed laterally and alternately, and one or two pairs on 
it a lateral process" (Allman). 

38 University of California Publications. [Zoology 

Nutting" adds the fact in his description thai the hydrocladia 
"alternate as a rule, but not regularly so in some specimens, 
where they are occasionally opposite toward the distal end of the 

The San Diego material consists of two stems, the longest 
measuring 100 mm., both unbranched. The stem internodes vary 
in Length, due to the obliteration of one or two nodes, and bear 
one, two or three hydrocaldia respectively. The number of their 
lateral uematophores varies with their length, from two to four. 
There is a pair of oematophores on each basal process, also an 
unpaired conical open process from which coenosarc projected 
in one case, and is probably to be reckoned as a nematophore. 
The hydrocladia are slender. In the first formed regions of the 
colony they alternate-, in the younger, distal regions they are 
usually opposite, as Nutting has said. A further variation in the 
distal region consists in an alternation of successive pairs of 
hydrocladia. so iluit the members of r\iT\- other pair lie in a 
plane making an angle somewhal less than 90 with the original 
plane of the colony. The basal internode of each hydrocladium 
is short toward the base of the stem, with a single mesial nemato- 
phore. In the younger distal pari of the colony it is usually 
wanting. Occasionally a non-thecate internode fuses with a 
thecate internode. The thecate internodes frequently hear two 
mesial uematophores. 

One stem had produced a heteromorphic shoot with several 
hydrocladia alternately placed, as is the rule with the basal pari 
of the stem. 

This species appears to be closely allied with P. filicula All- 
man, but its hydrothecae are no1 so deep and its habit is less 

42. Plumularia plumularioides (Clark). 

Halt .rinnii .' i plumularioides, dark. 1876, p, 217, pi. 10, figs. 

16, 17. 
Plumularia plumularioides, Nutting, 1901, p. 62, pi. 4, fig. 3. 
Plumularia plumularoides, Torrey, 1902, p. 78, pi. 11, figs. 103, 
Trophosome. "Hydrocaulus erect, simple, straight, divided by 
transverse joints into internodes of considerable length, regularly branched 
and with a few annulations at the base; branches arranged alternately on 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 39 

opposite sides of the stem, one to each internode, having their origin in 
a small shonlder-like process just below each joint, divided usually into 
regular internodes, though in some cases, short internodes occur between 
the longer ones. Hydrothecae arranged uniserially, usually one to each 
internode, partly adherent to the stem, or entirely free, shallow, tapering 
slightly to the base, with an entire rim." (Clark). 

Gonosome. Gonangia borne on the shoulder processes supporting 
the hydrocladia. Immature ones alone known; widest distally, tapering 
abruptly to base. 

Distribution. San Diego, 15 to 25 fathoms. Cape Etolin, 
Al., 8 to 10 fathoms (Clark). 

43. Plumularia setacea (Ellis). 

Corallina setacea, Ellis, 1755, p. 19. 

Plumularia setacea, Lamarck, 1816, p. 129. 

Plumularia setacea, Hincks, 1868, p. 296, pi. 66, fig. 1. 

Plumularia setacea, Clark, 1876a, p. 261, pi. 41, figs. 1, 2. 

Plumularia setacea, Nutting, 1900, p. 56, pi. 1, figs. 1-4. 

Plumularia palmeri, Nutting, 1900, p. 65, pi. 6, figs. 4, 5; 1901, 
p. 188. 

Plumularia setacea, Torrey, 1902, p. 79, pi. 11, fig. 105. 
TropJwsovie. Stems 5 to 100 mm. long, non-fascicled, divided into 
internodes, each bearing a hydrocladium from a distal process. Hydro- 
cladia alternate ; basal internode short, non-thecate ; thecate and non- 
thecate internodes alternate; there may or may not be septal ridges at 
either end of each internode and associated with the hydrothecae. Latter 
not deeper than broad, broadest at margin. Nematophores polythalamic, 
2 supra and 1 infra-calyeine, 1 on each non-thecate internode except the 
basal internode of each hydrocladium, 2 on each cauline internode, 1 on 
the side opposite the hydrocladium, the other axillary. 

Gonosome. Gonangia borne on the stem near the axils of the hydro- 
cladia, much elongated, female somewhat longer and stouter than the 
male, with a long, narrow neck; small terminal aperture. 

Distribution. Pt. Loma, La Jolla, Catalina I., San Pedro, 
and Monterey, Cal. Victoria, B. C, Santa Barbara and San 
Diego (Nutting). Eastern U. S. (Nutting). Coasts of Europe 
and Gt. Britain (Hincks). Helgoland (Hartlaub). 

40 V a i versify of California Publications. [Zoology 


NOTE.— Only papers directly concerned with Pacific Coast hydroids or 
specifically referred to in the synonymies, are listed below. 

Agassiz, A. 

1865. North American Acalephae. 111. Cat. Harv. Mus. Comp. Zool., No. 2. 

Agassiz, L. 

1860-2. Contributions to the Natural History of the United States. 

Alder, J. 

1856. A Notice of Some New Genera and Species of British Hydroid 

Zoophytes. Ann. & Mag. N. H., (2), XVIII, p. 353. 

1857. A Catalogue of the Zoophytes of Northumberland and Durham. 

Trans. Tynes. Nat. Field Club, III. p. 1. 

Allman, G. J. 

1864. On the Construction and Limitation of Genera among the 

Hydroida. Ann. & Mag. N. H.. (3), XIII. No. 77. p. 345. 
L871. A Monograph of the Gymnoblastic or Tubularia Hydroids. Ray 

1877. Report on the Rydroida collected during the Exploration of the 

Gulf Stream. Mem. linn. Mus. Comp. Zool. V. No. 2. 
1883. Report on the Eydroida Dredged by B.M.S. Challenger. Pt. I. 

1885. Description of Australian. Cape and Other Eydroida, mostly 

new. from tin' Collects I Miss Gatty. .lour. Linn. Soc. 

Lond., XIX. p. 132. 
1888. Reporl on the Hydroida Dredged by H.M.S. Challenger. Pt. II. 

The Tubularinae, Cormorphinae, Campanularinae, Sertul- 

arinae, and Thalamophora. 

Bale, W. M. 

1888. On Some New and Rare Hydroida in the Australian Museum Col- 
lection. Proc. Linn. Soc. N. S. W., (2), III, Pt. I, p. 745. 

1893. Further Notes on Australian Hydroids. with Descriptions of 
Some New Species. Proc. Roy. Soc. Vict., VI, p. 93. 

Calkins, G. N. 

1899. Some Hydroids from Puget Sound. Proc. Bost. Soc. Nat. Hist., 
XXVIII, No. 13, p. 333. 

Clark, S. F. 

1876. Report on the Hydroids Collected on the Coast of Alaska and 
the Aleutian Islands by W. H. Dall. Proc. Ac. Nat. Se. Phil., 
XXVIII, p. 209. 

1876a. The Hydroids of the Pacific Coast of the United States south of 
Vancouver Island. Trans. Conn. Ac, III, p. 249. 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 41 

1879. Report on the Hydroida collected during the Exploration of the 
Gulf Stream and Gulf of Mexico, etc. Bull. Mus. Comp. 
Zool., V, No. 10, p. 239. 

Ehrenberg, C. G. 

1834. Beitrage zur physiologischen Kenntniss der Corallenthiere im 
allgemeinen, und besonders des Rothen Meeres, etc. Abh. 
Berl. Ak. for 1832, p. 225. 

Ellis, J. 

1755. An Essay toward a Natural History of the Corallines and other 
Marine Productions, etc. London. 

Ellis and Solander. 

1786. The Natural History of many curious and uncommon Zoophytes, 
collected from various parts of the globe. London. 

Fewkes, J. W. 

1889. New Invertebrata from the Coast of California. Bull. Essex 
Inst., XXT; also separate, pp. 3-50, 7 pis. 

Fleming, J. 

1828. A History of British Animals. Edinburgh. 

Hartlaub, C. 

1894. Die Coelenteraten Helgolands. Wissensch. Meeresuntersuch., I, 
h. 1, p. 162. 

1900. Revision der Sertularella-Arten. Hamburg. 

1901. Hydroiden aus <lem Stillen Ocean. Zool. Jahrb., Abth. Syst., etc., 

XIV, p. 349. 

Hincks, T. 

1861-2. A Catalogue of the Zoophytes of South Devon and South Corn- 
wall. Ann. & Mag. Nat. Hist., (3), VIII, p. 293. 
1868. A History of the British Hydroid Zoophytes. London. 

Johnston, G. 

1847. A History of the British Zoophytes. 2d. Ed., London. 

Lamarck, J. B. P. A. de. 

1815-22. Histoire Naturelle des Animaux sans vertebres. Paris. (Vol. 
IT, 1816). 

Lamouroux, J. F. V. 

1812. Bulletin Societe Philomatique. 

1816. Histoire des Polypiers Coralligenes flexibles vulgairement nommes 

Zoophytes. Caen. 
1821. Exposition Methodique des Genres de l'Ordre des Polypiers. 


Lesson, R. P. 

1836. Tableau de la Famille des Zoophytes Beroides. Ann. Sc. Nat., 
V, p. 254. 

Linnaeus, C. 

1767. Systema Naturae, 12th ed. 

42 University of California Publications. [Zoolocy 

Marktanner-Turneretscher, G. 

1890. Die Hydroiden des k. k. Naturhist. Hofmus. Ann. Hofmus. 
Wien, V. 

McCrady, J. 

1858. Gymnothaltnata of Charleston Harbor. Proc. Elliott Soc. S. I '.. 
I. p. 103. 

Mereschkowsky, C. 
1878. Studies on the Hydroida. 111. Systematic Pacts. Ann. & Mag. 
N. II. (•",), I. p. 322. 

Murray, A. 

I860. Description of New Sertulariadae from the California Coast. 
Ann. & Mag! X. E., (3), V. p. 250. 

Nutting, C. C. 

1899. Hydroida from Alaska and Puget Sound. Proc. U. S. Nat. Mas., 

XXI, p. 741. 
L900. American Hydroids. Pt. I. The Plumularidae. Spec. Bull. No. 

4, U. S. Nat Mm 
1901. Papers from the Harriman Alaska Expedition. XXI. The 

Hydroids. Proc. Wash. Ac. Sc., Ml. p. 157. 
1901a. The Hydroids of the Woods Hole Region. Bull. U. S. Pish 

Comm., XIX. p. 325. 
L902. American Naturalist, XXXV. p. 789. 
1904. American Hydroids. Pt. II. The Sertularidae. Spec. Hall. No. 

4, I'. s. Nat. Mas. 

Oken, L. 

1816. Lehrbucb der Naturgeschichte. Weimar. 

Pallas, P. 

1766. Elenchus Zoophytorum. Hagae Comitum. 

Peron et Lesueur. 

1S09. Tableau des caracteres generiques et specifiques de toutes les 
especes de Me'duses connues jusqu'a ce jour. Ann. Mns. 
d'Hist. Nat., XIV, p. 325. 

Sars, M. 

ls:;.">. Beskrivelser og Jagttagelser, etc. Bergen, p. 81. 

1862. Bemaerkningor over fire norske Hydroider. Porh. Vidensk. Selsk. 

Christ iania. 

Schneider, K. C. 

1897. Hydropolypen non Rovigno, nebst Uebersicht des System der 
Hydropolypen im Allgemeinen. Zool. Jahrb., Abth. Syst., 
etc.. X. p. 472. 

Thompson, D'A. W. 

1877. The Hydroida of the Vega Expedition. Vega Exp. Vetensk. 
Jakttagelser, IV, p. 387. 

Vol. 2] Torrey. — Hydroids of the San Diego Region. 43 

Torrey, H. B. 

1902. The Hydroida of the Pacific Coast of North America. Un. Cal. 

Publ., Zobl., I, p. 1. 
1902a. American Naturalist, XXXVI, p. 987. 

Trask, J. B. 

1854. On Some New Microscopic Organisms. Proc. Cal. Ac. Sc. 1853, 
1, p. 110. 

Van Beneden, E. 

1844. Eecherches sur l'Embryogenie des Tubulaires, etc. Nouv. Mem 
Ac. Brux., XVII, p. 3. 

Verrill, A. E. 

1871-2. Eeport upon the Invertebrate Animals of Vineyard Sound, etc. 
Eep. U. S. Comm. Fish and Fisheries. 

Wright, T. S. 

1859. Observations on British Hydroid Zoophytes. Edinb. New Phil. 
Jour., n. s., X, p. 105. 



Vol. 2, No. 2, pp. 45-51, PI. 1. December 21, 1904. 










Ctenophorae spherical, cylindrical or compressed, with or without 
winglike aboral processes; two simple or pinnate tentacles usually retractile 
into a sheath; meridional and oesophageal canals end blindly. 

Para Pleurobrachiidae. 

Cydippida without winglike aboral appendages. Body approxi- 
mately round in cross section. Sub-tentacular and sub-oesophageal rows of 
swimming plates equal in length. 

Gen. Euplokamis, Chun, 1880. 
Body elongated; cylindrical or moderately convpressed; rows of 
swimming plates reaching from pole to pole; tentacle sheath present. 

Three ctenophores taken in Prince William Sound, Alaska, by 
Professor Ritter, and now in the collection of the University of 
California, belong undoubtedly to Mertens' Beroe cucumis 
( E i^lokamis cucumis, Chun). All are compressed somewhat, so 
that the transverse diameters are to each other as 6 to 5. Pleuro- 
brackia, a typical example of a spherical ctenophore, may also 
be compressed to the same extent. For these reasons, too much 
stress should not be laid on the circular cross section of Euplo- 
kamis, which is rather to be distinguished from Pleurobrachia by 
its elongation, from Mertensia by its slight degree of compression 
and equal rows of swimming plates. According to recent figures 
by Vanhoffen (:04), cross sections of specimens of Mertensia 
ovum taken in Greenland were three to four times as long as 

46 University of California Publications. [Zoology 

1. Euplokamis calif ornensis, n. sp. 

Body moderately compressed, somewhat flattened at sensory pole, 
narrowed toward mouth. Tentacle sheaths about three fourths the 
length of the body, lying close to and parallel with the oesophageal canals, 
diverging slightly to openings near sensory pole. The four interradial 
canals arise independently from funnel. Distances from funnel to aboral 
and oral poles as 1 to 2. Tentacles yellow brown; inner opening of oesoph- 
agus purple. 

Distribution. San Diego, Cal. Taken at the surface and in 
vertical hauls from various depths to 125 fathoms with non- 
closing nets, during .May. June and duly. None were more than 
25 nun. Long. This species is very closely related to I', oucumis, 
with which it may prove to he identical. It is near, also, to the 
Cydippt elliptica of Eschscholtz, from the tropical Pacific. 

Pleurobrachia, Fleming, 1822. 

Body spherical, interradial canals from two stem canals. 

2. Pleurobrachia bachei A. Ag. 

Pleurobrachni bachei A. A.g., 1.. Agassiz, I860, p. 294. 
\. Agassiz, L865, p. 3 1. 
Oesophagus equal to or less than tunnel tube in length; tentacle 
sheaths distant from funnel, aboul half as long as body, divergent, open- 
ings about one fourth the distance from pole to pole from sense organ; 
stems canals long, all canals slender. 

Remarkably transparent, and colorless with ihe exception of the 
tentacles, which are yellowish red, and the oesophagus, which is blotched 
with deep purple proximally. 

Distribution. San Diego to Pugel Sound. This species 
differs from /'. j>il< us i Kabr.) of the Atlantic, having a shorter 
oesophagus and longer funnel tube, and longer and more slen- 
der stem and interradial canals. The openings of the tentacle 
sacs are somewhat farther from the sensory pole. 


Body compressed, with two lateral lobes. Subtentacular rows of 
swimming plates shorter than others, with four auricular processes at their 
ends. Mouth large. Four interradial canals direct from the funnel. 
Tentacles rudimentary, near oral pole, without sheathes. A Mertensia 
stage in the development. 

Vol. 2] Torrey. — Ctenophores of the San Diego Region. 47 

Pam. Bolinidae. 

Bolina, Merteus, 1833. 

3. Bolina sp. 

There are two recognized species of Lobata on the western 
coast of North America : Bolina septentrionalis Mertens, from 
Behring Str. and B. microptefa A. Ag., from the Gulf of Georgia. 
Agassiz and Mayer have described another, EucJiaris grdndi- 
formis, from the Fiji Islands. It is probable that the very young 
Lobata which have been taken in large numbers off San Diego 
for the past two summers belong to B. microptera, which may 
ultimately prove to be identical with Mertens' circumboreal spe- 
cies. But the development of these immature individuals has 
not proceeded to the appearance of the auricles, and the total 
absence of mature individuals make it obviously impossible for 
the present to determine even the family of the species with 


Ctenoplwrae elongated, conical or ovate, compressed, with 
large mouth and oesophagus. Tentacles and tentacle canals 
wanting. Meridional canals communicate with oesophageal 
canals at the edge of the mouth, and send out numerous branches 
which may form a peripheral network. 

Pam. Beroidae. 

With the characters of the order. 

Beroe, Browne, 1756. 

With the characters of the family. 

4. Beroe forskali M. Edw., Chun. 

fBero'e rufescens Forskal, 1775, p. 111. 

Cydalisia mitraeformis, Lesson, 1843, p. 138, pi. 2, fig. 2. 
ldya penicillata, Mertens, 1833, p. 534, pi. 12. 
Beroe Forskalii, Milne-Edwards, 1841, p. 207, pi. 5. 
Beroe Forskalii, Chun, 1880, p. 309, pi. 14, figs. 3-5; pi. 14a, 
figs. 6-10. 

48 University of California Publications. [Zoology 

Body much compressed, conical, tapering from the very broad mouth 
with full lips to a narrow sensory pole. Fine network of vessels between 
meridional canals, communicating also with oesophageal canals. Gonads 
in lateral follicles of meridional canals. Bows of swimming plates reach 
almost from mouth to tip. 

Distribution. San Diego, Cal. Peru (Lesson). South Pacific 
(Mertens). Mediterranean (Forskal). 

Taken about ten miles off shore, at the surface and in vertical 
hauls from various depths to 125 fathoms with non-closing nets, 
during May. June and July. A single mature specimen was 
taken, with the typical pointed form which is much more pro- 
nounced than in young individuals. The early stages were com- 
monly taken in considerable numbers, and resemble in shape the 
young of B. (roseola) cucumis according to L. Agassiz, and the 
adult of B. cyathina according to A. A.gassiz, the aboral end 
hemispherical and the rows of swimming plates short. The 
transition to the pointed forms is gradual and convincing The 
very young are colorless, the half grown are rosy, with brilliantly 
irridescent rows of swimming plates. 

\ t ol. 2] Torrey. — Ctenophores of the Man Diego Region. 49 


Agassiz, A. 

1865. North American Acalephae. 111. Cat. Harv. Mus. Corap. Zool., No. 2. 
Agassiz and Mayer. 

L899. Acalephs from the Fiji Islands. Bull. Harv. Mus. Comp. Zool., 
XXXII, No. 9. 

Agassiz, L. 

1860. Contributions to the Natural History of the United States, ITI, 
p. 153. 

Brandt, J. 

1838. Ausfuhrliehe Beschreibung, etc. Mem. Ac. St. Petersb., (6), IV, 
p. 237. 

Browne, P. 

1756. Civil and Natural History of Jamaica. 

Chun, C. 

1880. Die Ctenophoren. Fauna iind Flora des Golfes von Neapel. 

1897. Die Beziehungen zwisehen dem Arktischen und Antarktischen 


1898. Die Ctenophoren. Ergebnesse der Plankton Expedition, [I:k:a. 

Eschscholtz, F. 

1829. System der Akalephen. Berlin. 

Fabricius, O. 

1780. Fauna Groenlandica. Hafniae et Lipsiae. 
Fleming, J. 

1822. Phil. Zoology. 

Forskal, P. 

1775. Descriptiones Animalium, etc. Hauniae. 

Lesson, R. P. 

1843. Histoire Naturelle des Zoophytes. Acalephes. Paris. 

Mertens, H. 

1833. Beobachtungen und Untersuchungen iiber die Beroeartigen Aka- 
lephen. Mem. Ac. St. Petersb., (6), II. p. 479. 

Milne-Edwards, H. 

1841. Observations sur la structure et les f'onetions de quelques Zoo- 
phytes, Mollusques et Crustacea des cotes de la France. Ann. 
Sc. Nat. (2), XVI, p. 193. 


1S57. Naturh. Bid. til en Besk. af Gronland. 

Vanhoffen, E. 

1895. Die G-rbnlandischen Ctenophoren. Bibl. Zool., VIII (h. 20), p. 15. 
1904. Nordisches Plankton. XI. Ctenophoren. 


Pig. 1 .— Mertensia ovum. 

Fig. 2.— Beroe forskali. 

Fisr. :>. — Pleurobrachia bachei. 




Vol. 2, No. 3, pp. 51-112, Pis. 2-3. January 31, 1905 











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Wm. e. ritter. 

In accordance with the general plan of the series of faunistic 
papers of which the present is one, the object has been kept con- 
stantly in view not merely of describing the new pelagic tuni- 
cates occurring in the area and of reporting the presence of such 
already familiar ones as have thus far been found; but of fur- 
nishing a ground work, as well designed and securely constructed 
as possible, for future investigations into the general biology of 
this group of animals. 

Diagnoses are given of all the species thus far observed — in 
the Thaliacea covering each of the different generations; and 
as far as possible these have been written from the living animals. 
Furthermore, all the forms actually observed in any numbers 
are figured. Everyone, however, who has had experience with 
these creatures is aware of the impossibility of making drawings 
of them that are at best more than crude outlines ; and that to 
accomplish this the carcasses, always at the best badly out of 
shape, must be resorted to for many points. Lateral views are 
shown in every case, partly because it seemed that on the whole 
these are the most useful as aids to identification, and partly 
because most of the published figures of the same species present 

52 University of California Publications. [Zoology 

dorsal views ; and I have thought the lateral views would supple- 
ment to good advantage the figures of other writers. 

Before the alternation of generations in salpa was known, the 
oozooid and blastozooid generations were described as separate 
species; and, of course, were given different specific names. In 
most instances the describers of the two generations were differ- 
ent persons. Prom this duplication of dames considerable con- 
fusion has arisen, not all writers having adopted the same course 
in the treatment of the names after the true relations between 
the two generations had been recognized. Following the lead 
of Krohn, 184(>, the majority, probably, of authors have retained 
both the specific names, writing them with a hyphen, thus, Salpa 
<l<niocrata-»nicronata. Others, on the contrary, have retained 
lull one of the names, hut unfortunately those who have adopted 
this course have not all retained the same name: thus Lahille, 
1890, has used confoederata, and Brooks, 1893, scutigera, for S. 
confoederata-scutigera. I have followed Krohn, with the slight 
modification that I have uniformly placed that name firsl in the 
couplet that was proposed at the earlier date. This has reversed 
the order in several species, as. for example. 8. fusiformis-runci- 
nata, this being written by other authors runcinata-fusiformis. 
But as fusiformis was proposed by Cuvier in 1804, while runci- 
itala was introduced for the other generation by Chamisso in 
1819, it seems more fitting that fusiformis should lead in the 
couplet, as should also the name Cuvier indicating the author's 
names after the species. 

The species treated in this paper are as follows: 


Cyclosalpa bdkt ri,>. 

Cyclosalpa ajfinis. 

Sul jut fusiformi8-rundnata. 

Salpa tilesii. 

Salpa democratica-mucronata. 

Salpa sonaria-cordiformis. 

Salpa cylindrica. 

Salpa confoederata-scutigera. 


Doliohim tritonis. 
Dol ioht m eh renbergii. 
Doliohim miilleri. 

Vol. 2] Hitter.— The Pelagic Tunicata. 53 


Pyrosoma giganteum. 
Of these, Cyclosalpa bakeri and the trophozooid of Doliolum 
tritonis have not hitherto been described. 

Class UROCHORDA Lankester (Tunicata, Lamarck). 
Order I.— THALIACEA Van der Hoven. 
Pelagic urochorda, with body of zooid more or less cylin- 
drical, the branchial and atrial orifices being situated at oppo- 
site ends, or nearly so, of the body. Mantel musculature limited 
almost entirely to encircling fibres, these being grouped into defi- 
nite bands. Propagation through an alternation of heteromor- 
phic oozooid and blastozooid generations. 

Fam. I. — Salpidae, Forbes, 1853. 
Thaliacea in which the branchial stigmata are reduced to a 
single pair, each very large, between which, extending the entire 
length of the great pharyngeal cavity, is a prominent vascular 
band known as the gill. Muscle bands rarely extending around 
the body without interruption, the break usually being on the 
dorsal and ventral sides. Oozooids and blastozooids presenting 
each one form only. 

Genus 1— Cyclosalpa, DeBlainville, 1827. 

Thalia, Browne, 1756. 

Salpa, Forskahl, Ouvier, Traustedt, and most writers. 

Cyclosalpa, Herdman, Lahille, Apstein. 

Salpidae in which the intestine never forms a compact spher- 
ical mass known as the "nucleus," but is either extended along 
the dorsal side of the "gill" as a straight tube, or disposed in a 
large ring. Blastozooids set free from the proliferating stolon 
of the parent in circular groups, or whorls. 

Genus 2. Salpa, Forskahl, 1775. 

Hololhuria, Linn, 1758. 

Dagysa, Banks and Solander, 1773. 

Biphora, Bruguiere, 1789. 

Tethys, Tilesius, 1802. 

Pegea, Savigny, 1816. 

Jasis, Savigny, 1816. 

Pterolyra, Lesson, 1830. 

Dubrieullia, Lesson, 1830. 

54 University of California Publications. [Zoology 

Salpidae in which the intestine is massed into a compact 
spherical body known as a nucleus. Blastozooids detached from 
the proliferating stolon of the parent either one by one or in 
groups consisting of two parallel series. 

Cyclosalpa bakeri n. sp. 

(a) Solitary (budding) <i< hi ration. — PI. II, all figures. Body 
somewhat cask-shaped, the posterior end a little narrower than 
the anterior: a distinct, though not deep, constriction setting 
off the anterior end. Long axis nearly straight as seen in pre- 
served specimens, 1 PI. II, fig. 1 ; section of body everywhere cir- 
cular. Length of largest zooid seen. 41 mm., this with 
salpa chain well developed. 'Pest exceedingly soft and trans- 
parent and wholly devoid of colored pigment. Five white "longi- 
tudinal organs" on each side, the first pair between second and 
third muscle bands. Upper lip of branchial orifice truncate, 
Lower lip founded and protruding more than upper. Atrial ori- 
fice without lips. Muscle bands very delicate and difficult to 
trace. Body bands ten, though at neither end decisively separ- 
able from orifice muscles. All hands except ninth and tenth. 
and sometimes eighth, interrupted dorsally. and all except first, 
ninth and tenth interrupted vent rally. First trending backward 
on dorsal side, the ends becoming nearly parallel and reaching 
as Ear hack as the third: also connected on dorsal side with the 
posterior lip hand by two parallel Longitudinal muscles. Second 
also trending backward on dorsal side to terminate almost coin- 
cidentally with first and third. Seventh bending abruptly for- 
ward on dorsal side and running parallel close together as far 
as the fifth, sometimes broken into fragments, PI. II, fig. _. 
Eighth also turning forward on dorsum, but less abruptly than 
seventh, sometimes interrupted in median line, sometimes not. 
Second band trending backward on ventral side; likewise third, 
though less than second. Fifth inclined somewhat forward ven- 

1 Though attention should be called here to fig. 4, PI. II, which outlines 
the form of a specimen alive and swimming. From this observation, and 
from the fact that C. aflinis certainly has normally a more sinuate general 
form in life than after preservation, I am inclined to believe that when 
sufficient numbers of living individuals of this species have been examined 
to determine the point, it will be found that fig. 4 represents the normal 
form more nearly than does fig. 1. 

Vol. 2] Bitter.— The Pelagic Tunicata. 55 

trally, and sixth bending abruptly forward to terminate on a level 
with the fifth. Ninth inclining forward on each side to touch 
the eighth tangentially, then bending sharply back to cross the 
tenth, so that on the median ventral line ninth is behind tenth. 

Three bands in dorsal lip of branchial orifice, the first divided 
into a broader and narrower portion, the second trending broadly 
backward on ventral side to become confluent with second body 
muscle; third joining second behind angle of orifice. Second 
muscle of ventral lip trending sharply backward laterally to ter- 
minate at the crossing of the second upper lip and first body 
muscles. A longitudinal band on each side extending forward 
from the second body band to angle of orifice. Endostyle rather 
slender, and with a gentle dorsal curvature near its anterior end, 
extending from midway between the second ventral lip and the 
first body muscle, to the seventh muscle. Gill of usual form and 
extent, posterior termination nearly coincident with that of the 
endostyle. Ganglion considerably behind anterior termination 
of gill, and remote from hypophysis; sense organ broad horse- 
shoe shaped, open end forward. Hypophysis at the anterior 
termination of the gill ; in the form of a twisted horseshoe with 
open end forward and slightly to the right. Intestine straight, 
extending forward above the gill nearly to the ganglion. Stom- 
ach globular, not large. Two strap like lobes of nearly equal 
size and length given off backward from the short bend of the 
intestine. Heart slightly in front of the posterior termination 
of the endostyle, in the interval between the sixth and seventh 
muscle bands. Stolon apparently arising in front of the heart, 
extending forward in the 'mid-ventral line to emerge to the out- 
side through an opening between the second and third muscle 

(b) Aggregate (sexual) generation. PI. Ill, figs. 7 and 8. 
The only zooids seen of this generation were still attached to the 
stolon, and I am, consequently, unable either to describe the full 
grown animal, or to say anything positive about the whorls. 
From the fact, however, that the musculature is essentially the 
same in form and arrangement as that here described, through 
several stages preceding the one on which the description is 
based, it is safe to presume that it will be found to be practically 

56 University of California Publications. [Zoology 

the same in the adult. The hypophyseal-ganglionic complex, and 
especially the digestive tract, are, however, obviously still imma- 
ture. As to the whorls of zooids, it can only be said at present 
that the close similarity of this species to CycJosalpa affinis and 
C. pinnata in the arrangement of the zooids in the chain, makes 
it highly probable that the whorls are likewise much the same in 
the two. The diagnosis is from zooids 4 mm. long, exclusive of 
the intestinal tract. 

Body compressed, cask shaped, the ends imperfectly truncate. 
though adult form probably not yet assumed. Test thick, and 
consistency of animal as ,-i whole much firmer than adults of soli- 
tary generation. Peduncle for attachment to stolen situated 
on ventral side nearer anterior end. relatively Long and narrow, 
containing continuations of the firsl and second body muscles. 
Branchial orifice terminal, lips (a1 iliis stage) scarcely recog- 
nizable. Atrial orifice small, situated at dorso-posterior angle 
(in this stage). Body muscles. 4 on dorsal side, and 4 on ven- 
tral, these branching and anastomosing Laterally in a complicated 
way. and always with a <l</inii< asymmetry, the arrangement on 
either side of a given zooid depending on whether the zooid be 
a right or left one in the salpa chain; or, what amounts to the 
same thing, whether the side is turned toward the anterior or the 
posterior end of the parent of the chain. Arrangement on side 
toward anterior end of parent is as follows: second muscle bifur- 
cates ;i little distance from median dorsal line, the branches unit- 
ing again about opposite the endostyle to extend into the ventral 
peduncle, PL III. tig. 7. Third muscle likewise bifurcates about 
same distance ,-is second from dorsal median line, the anterior 
branch extending down to cross 1 he ventral median line and to be- 
come continuous with its fellow of the opposite side of the zooid, 
and also anastomosing with a ventral longitudinal band running 
into the peduncle. The posterior branch likewise extending 
across ventral line to join fellow of opposite side, but also send- 
ing two delicate branches posteriorward, approximately parallel 
with the endostyle, the more dorsal passing midway between the 
atrial orifice and the esophagus, the other ventrad of the esopha- 
gus, and both extending into the "post abdomen," PL III, fig. 7. 
Arrangement on side turned toward posterior end of parent is 

Vol. 2] Bitter.— The Pelagic Tunicata. 57 

as follows : second muscle not bifurcating ; third bifurcating near 
dorsal line as on opposite side, the anterior branch again bifur- 
cating to send a branch forward which anastomoses with second 
muscle, the single band thus produced passing into the ventral 
peduncle, PL III, fig. 8. In the asymmetry of the muscles this 
species resembles 8. rostrata, 8. punctata, 8. niagallianica of Ap- 
stein. The climax of the phenomenon is reached in S. rostrata. 
First and second muscles united on each side by two longi- 
tudinal bands, PI. Ill, fig. 7. In addition to the dorsal 
lip muscle given off as a branch from first body muscle, 
an angular lip muscle on each side its dorsal limb reaching 
over and becoming continuous with its mate of the other side ; 
another small upper lip muscle near edge of lip. The fourth body 
muscle of dorsal side, relatively very small, bifurcating on each 
side short distance from dorsal median line, the anterior branch 
anastomosing with posterior branch of third muscle, and pos- 
terior branch passing under atrial orifice. The fourth ventral 
muscle is really the posterior branch of third dorsal muscle. 

As already stated, the ganglio-hypophyseal complex and the 
intestinal tract are clearly immature in the largest zooids seen. 
I, consequently, refrain from including a characterization of them 
in the diagnosis of the species. Certain facts about the intestinal 
tract, however, should be mentioned. In the first place, the late 
period in the life of the zooid at which it becomes complete, at 
least as compared with C. affinis, is noteworthy. In the latter 
species, the intestine has assumed its final form and position 
while the bud is still in the chain, and even before the whorls 
are formed. In C. bakeri, on the contrary, the way in which the 
anal end of the intestine projects freely from the posterior end 
of the body as a whole, shows clearly that the organ is not yet 
complete, even in the oldest zooids found. "With little doubt the 
final form is a circle here as in affinis. An apparently wholly 
unique feature in bakeri, however, seems to be the two appendages 
of the intestine shown in the figures. The intestinal tract as a 
whole has the form of a horseshoe, .the plane of the shoe being 
approximately at right angles with the sagittal plane of the 
zooid. The entire bow projects backward beyond the posterior 
end of the endostyle and gill. The mouth of the esophagus is 

58 University of California Publications. [Zoology 

situated a little dorsal and to the right of the end of the endo- 
style ; and the anus enters the atrium to the left of the endostyle. 
The esophagus is marked off from the stomach by being distinctly 
less in diameter than the stomach. At the anterior end of the 
intestine a cecum nearly as large as the intestine itself is given 
off, which curves backward and upward and forms a very con- 
spicuous object in all the stages of development observed, co., figs. 
7 and 8. 

From the posterior extremity of the intestinal bow a great 
finger-like outgrowth of the mantle extends backward and up- 
ward. This appendage is even longer and more conspicuous than 
the stomachal cecum described above. Into it extend prolonga- 
tions of the posterior branches of both dorsal and ventral body 
muscles; and in addition it contains a well defined axial strand, 
the connections and nature of which are doubtful, p. d., fig's. This 
appendage would seem to be comparable with the portion of the 
post-abdomen that extends beyond the intestinal loop in various 
compound ascidians. The axial strand is probably the testis, or 
a portion of it. The ovary is situated on the right side of the 
body at the extreme posterior end. midway between the atrial 
orifice and the esophagus; and the oviduct, which is unusually 
long, extends forward to a level with the third ventral muscle 

C. bakeri appeals to have more in common with C. floridana, 
Apstein, than with any other known species. It is, however, 
very distinct from this latter, as is obvious from the following, 
among several other differences: The largest specimen of C. 
floridana seen by Apstein was 12 mm long. In view of the con- 
siderable number of specimens taken by the Plankton Expedi- 
tion, the great disparity in size thus indicated shows pretty con- 
clusively that C. b(ih( ri is a much larger species than floridana. 
The lateral glandular organs of the solitary floridana are dis- 
tinctly less extensive than in bakeri, and are, according to Ap- 
stein 's statement, continuous on each side as in C. pinnata. The 
intestine of floridana has a single appendage, while that of bakeri 
has two. The closest resemblance between the two is in the mus- 
culatures of the solitary forms, but even here there are well 
marked differences, which, however, need not be dwelt upon, 

Vol. 2] Ritter.—The Pelagic Tunicata. 59 

since a comparison of my figures with that of Apstein will 
make them clear. It is worth noting that this adds another to 
the list of species in which the intestine of the aggregate genera- 
tion is in the form of a circle, these species being C. affinis, C. 
floridana and C. bakeri. 

The specimens of C. bakeri thus far obtained are few, and 
the variations in the muscle bands in these few suggest that fur- 
ther study of more ample material may modify somewhat the 
scheme given in the diagnosis ; I cannot, however, believe that such 
modification can materially alter the results so far as concerns 
the definition of the species. In no species of salpa with which 
I have had experience have I found so much difficulty in tracing 
the muscles. This difficulty is due to the softness of the animal, 
and the extreme delicacy and transparency of the muscles them- 
selves. The separation of the muscles into body muscles on the 
one hand, and orifice muscles on the other, I recognize as being 
a particularly arbitrary matter in this species. For example, 
there would be almost as much reason for considering what I 
have enumerated as the first body muscle, a lip muscle; or, on 
the other hand, for calling what in my scheme is the third upper 
lip muscle, a body muscle. In fact, I have little doubt that what 
Apstein has designated number one in floridana corresponds to 
my third dorsal lip muscle. But the homologizing of the muscle 
bands in the different species of salpa is an exceedingly difficult, 
if indeed possible, thing; though comparison of the develop- 
mental stages would probably help in the matter. 

In all, about fourteen specimens of the species have been 
taken during the last three years, all on the coast of southern 
California, and all excepting one, which was taken in March, 
during the months of June and July. 

Cyclosalpa affinis (Chamisso). 

Salpa affinis Chamisso, 1819, p. 11, pi. figs. 2A-C, solitary genera- 
tion; 2D-E, aggregate generation. 
Cyclosalpa affinis Blainville, 1827. 
Salpa affinis Meyen, 1832, p. 407. 

Salpa affinis Traustedt, 1885, p. 357, PI. T, figs. 6, 7, and 8. 
Cyclosalpa affinis Herdman, 1888, p. 86. 
Cyclosalpa affinis Lahille, 1890, p. 11. 
Salpa affinis Apstein, 1894, p. 4; Cyclosalpa, p. 24. 


University of California Publications. 


(a) Solitary (budding) generation. — Fig. 9. Body distinctly 
larger at anterior end, and tapering nearly uniformly to the 
posterior end ; the anterior end with a pronounced ventral bend, 
the posterior with a nearly equal dorsal bend. Test rather thin 
and soft, and highly transparent, without special thickenings or 

*?■ h 

Fig. 9. — »S. affinis. solitary gcniM-atioii. 

asperities; a pair of lateral and somewhal dorsal appendages at 
the posterior end. No lineaform "glandular" organs. Length 
of full grown zooids, 80 to 100 nun. Branchial orifice directed 
somewhal vrentrad, lips prominent, upper overarching. Atrial 
orifice without lips, directed somewhal dorsad, armed, particu- 
larly in older specimens, with a pair of latero-dorsal appendages. 
Body muscles eight, all excepting lasl interrupted on ventral 
side. First and second interrupted on dorsal side (in older 
specimens only). First trending backward to some extent on 
ventral side. Lip muscles of branchial orifice complicated; two 
sphincters in upper and one in lower: two longitudinal bands on 
each side running forward from the first body band, one to the 
dorsal lip, the other to the ventral: a band on each side extend- 
ing from the angle of the orifice postero-dorsad ; a pair of short, 
strong dorsal longitudinal bands in dorsal lip. Endostyle dis- 
tinctly curved in its anterior third to correspond with the ventral 
bend of the body as a whole: extending from far forward in the 
ventral lip back to the esophagus; both extremities turned 
abruptly up, the anterior more conspicuously so than the poste- 
rior. Ganglion slightly in front of anus. Hypophyseal mouth a 
large, highly convoluted, nearly closed ring, with opening to the 
left (fiff. 11). Gill long and narrow, extending from slightly 
in front of the anus to the esophagus, terminating, consequently, 

Vol. 2] 

Bitter. — The Pelagic Tunicata. 


Fig. 11. — Hypophysis of C. affiitis. 

nearly coincidently with the endostyle. Intestine nearly straight 
and of uniform character, though turned a little dorsad 
and to the left at anal end ; the anus somewhat trumpet shaped. 
Stomach in the sharp curve of the tract, flattened; a large cecum 
extending backward appearing as a direct posterior prolongation 
of the intestine. Entire intestine uniform orange, generally, but 
occasionally devoid of color. Pericardium-heart large, situated 
ventrad of the posterior end of the endostyle. Salpa chain reach- 
ing forward in mid-ventral line under the endostyle and emerg- 
ing to the outside far forward, between the ventral ends of the 
first bodv muscle band. 

■pi a. 

Fig. 10. — C. affinis, aggregate generation. 

(b) Aggregate (sexual) generation. — Fig. 10. Aggregations 
containing from nine to twenty zooids, united radially about a 
common centre, by the large ventral peduncle, remaining intact 
until zooids are fully grown ; in nature, six or eight or more of 

62 University of California Publications. [Zoology 

the whorls united tangentially. Body in general cylindrical, 
though somewhat arched dorsally, and tapering at the atrial end. 
Rather soft and of uniform consistency, there being no thick- 
ened or specially stiffened areas in the test; the connecting 
peduncle and protruding intestinal tract forming very promi- 
nent projections from the ventral side. Full grown zooids from 
60 to 70 mm. long. Body usually quite transparent and without 
pigment, but occasionally traces of pink on surface of test about 
anterior end. Lips of branchial orifice prominent, of approxi- 
mately equal size: atrial orifice without lips. Body muscles five 
on the dorsal side and six on the ventral, ;ill continuous acrdss 
the dorsal side, ami ;ill except sixth interrupted by a narrow- 
interval ventrally; the fifth and sixth ventral muscles joining 
laterally to form the fifth dorsal. The second band giving off a 
longitudinal muscle laterally, which extends forward to the angle 
of the branchial orifice; the fifth giving off a small branch on 
each side, which extends back close under the intestine; the 
sixth, with a branch on each side connecting with the firsl atrial. 
A well developed band in each Lip of the branchial orifice meet- 
ing in a wide angle on each side. Firsl atrial band nearly as 
broad as body bands: the other al rials, about eight in number, 
very delicate. Endostyle extending from slightly in front of the 
first body muscle to behind the fifth muscle, and quite to the 
intestine. Gill extending from somewhat in front of the second 
muscle fully to the intestine, hence terminating near the poste- 
rior end of the endostyle. Ganglion and sense organ under the 
second body muscle, and a little posterior to the anterior end of 
the gill. Hypophyseal mouth close in front of the anterior end 
of the gill, large and much convoluted, becoming rosette shaped, 
the band narrowly open to the right. The intestine projecting 
from the ventral side like a hernia; forming almost a circle of 
relatively large size, in full grown zooids 1.5 cm. in diameter, 
the curve extending downward and forward, so that the anus 
is very near to, and to the left of, the esophageal mouth. Stom- 
ach scarcely larger in diameter than intestine, and not distinctly 
set off from it. Esophagus short and much smaller in diameter. 
Deep 3 r ellow\ nearly uniform throughout. Heart conspicuous, 
immediately in front of intestinal ring on ventral side. Placenta 

Vol. 2] Ritter. — The Pelagic Tunicata. 63 

with embryos on right side between fourth and fifth muscle 
bands. Testis, an elongated white mass in the intestinal ring 
closely applied to the intestine. Vas deferens given off from 
the middle of the testis, passing across the intestinal circle, and 
opening near the esophageal mouth. 

Down to the present time this has been one of the rarer of 
the early known species of salpa. It was described by Chamisso, 
by whom it was taken in the region of the Sandwich Islands. 
It did not occur in the Challenger collections, and has not until 
now been reported again from the Pacific, so far as I am aware. 
Meyer observed it, though not closely, nor in great abundance, 
about the Canary Islands, and we have several other ref- 
erences to its occurrence in the tropical and subtropical 
Atlantic. The Plankton Expedition took, according to Apstein, 
a total of nine specimens at two stations ; one in the Gulf Stream, 
Lat. 41.6, the other in the Sargasso Sea, Lat. 31.5. Voigt, 1854, 
includes it in his list of species of the Mediterranean in the 
vicinity of Nice, but gives no further information about it. Its 
constant abundance on the southwestern coast of North America, 
in a plankton area at least adjacent to, if not in reality part of, 
that from which it was originally described, and its apparent- 
rarity in other parts of the world, would seem to indicate that 
the headquarters of the species is here, though such a suggestion 
relative to the distribution of strictly pelagic organisms can 
have little value until supported by much more data than we 
yet possess. Despite the considerable differences between the 
Salpa here treated as C. afftnis and any of the published descrip- 
tions and figures of the species, I am convinced of the correct- 
ness of the identification. All the discrepancies of any moment 
may be accounted for from the fact that the descriptions and 
figures hitherto published have probably all been made from 
museum specimens. For example, the straight long axis of the 
solitary generation as shown in the figures of Chamisso and 
Traustedt, give a wholly erroneous impression of the general 
form of the species, but the true shape as shown in lateral view, 
fig. 9, can be fully appreciated only by examining the living 
swimming animal. Preserved specimens have more the form of 
the fioures of the authors above mentioned. Again, the two 

64 University of California Publications. [Zoology 

processes at the atrial end are not adequately recognized in any 
of the published figures. This is due in part to the fact that 
they are very short in the younger zooids, and in part to their 
being, in adult preserved specimens, either broken or worn off, 
I have examined specimens of about 1.5 cm. in length from the 
mid-Pacific, practically the region in which Chamisso obtained 
his, and find the processes short though distinctly indicated, 
essentially as they are shown in Traustedt's figs. 7 and 8, PI. I. 
Finally, Traustedt figures an arrangement and anatomosing of 
the seventh and eighth muscle bands of the solitary generation, 
somewhat different from anything I have seen, but the point is 
a trivial one, even if his representations are entirely correct. 

The species has been obtained at Santa Catalina Island and 
off San Diego during the months of March, June, July, and 
August. It was particularly abundant during March. 1!)(>4. in 
the last named locality, and was reproducing actively, both sex- 
ually and asexnally. 

Salpa fusiformis-runcinata Cuvier.-Cham. 

Salpa fusiform is Cuvier, L804, p. 23, fig. 10. 

Salpa nmoinata chainisso, 1819, p. 16, PI. figs. 5A-51. 

Salpa runcinata-fusiformis Crohn, 1846, p. 112. 

ShI/hi runcinata- f fiinn is, Leuckart, 1S54, p. 3 ad seq. PI. I, figs. 

6, 8, 16, 17, 18; and PI. II, figs. 1, 3, 4, 5, 13, 15, and 18. 
Salpa niiicniiild-fiisifnrniis Traustedt. L885, p. 370, PI. 2, figs. 

29, 30, 31. 
Salpa runcmata-f usifi ) mi is Henlinan, IsSS, p. 7(i, PI. (j, figs. 5-12. 
Salpa fusiformis Apstein, 1901, p. 1117, figs. 6a and 6b. 

Fig. 12. — £. fusifoniiis-nuicinata, solitary generation. 

(a) Solitary (budding) generation. — Fig. 12. Nearly cylin- 
drical, somewhat larger at the atrial end, both orifices terminal, 
truncate in general effect at both ends. Length of largest speci- 

Vol. 2] Hitter. — The Pelagic Tunicata. 65 

mens, 70 mm., 76 mm., to 80 mm. Test variable, in some thin 
and soft, particularly anteriorly; in others, thicker and firmer, 
particularly posteriorly ; a number, from eight to twelve, more 
or less regular, longitudinal, serrated ridges running from the 
atrial end forward a variable distance, but most prominent over 
the nucleus. Branchial orifice with two lips, of which the dorsal 
is distinctly the higher. Atrial orifice also with inconspicuous 
dorsal and ventral lips. Body muscles nine, confined to the 
dorsal side and reaching down laterally scarcely half way to the 
endostyle, where they terminate abruptly. First three anterior 
body muscles confluent dorsally ; all the muscle bands distinctly 
broader dorsally. A constrictor muscle in the dorsal lip, and 
one in the ventral lip, both terminating behind the angle of 
branchial orifice, where the ends cross each other and extend a 
short distance beyond the point of crossing. Atrial orifice with 
six or eight muscles of a few fibres each, those of the dorsal and 
ventral lips terminating at the angles, where they cross one 
another. Endostyle slender, straight, extending from the level 
of the angle of the branchial orifice back to the intestine, on a 
level with the eighth muscle band. "Gill" long, narrow, and 
nearly straight, extending from a little in front of the first 
muscle band to the ninth band. Hypophysis horseshoe shaped, 
its plane nearly in the sagittal plane of the animal's body, about 
midway between the muscle band of the upper lip and the first 
body band. Intestinal tract making a compact "nucleus," cor- 
responding to the interval between the eighth and ninth muscle 
bands, and projecting somewhat on ventral side ; the broad short 
end of the rectum projecting dorsad from the nucleus to open 
into the cloaca. Dark red generally, though not universally. 
Heart on ventral side, immediately in front of intestinal mass. 
Chain of buds extending forward along ventral median line from 
near the nucleus for a variable distance, then bending on itself 
and reaching back to emerge to the outside through an orifice 
behind the nucleus. 

(b) Aggregate (sexual) generation. — Fig. 13. Body ellipt- 
ical in outline, with processes at each end, short and broad in 
the young, much longer in full grown zooids, where they become 
as long as the body. At the outset these processes are always 


University of California Publications. 


asymmetrical, the anterior being to the right, the posterior to 
the left, or vice versa (the one figured is anterior process right, 
and posterior left). Length of large specimen, 25 mm.; usually 
smaller. Branchial orifice inclined distinctly upward, though 
not wholly dorsal. Atrial opening nearly directly backward. 
Lips of branchial orifice not prominent, dorsal deeper, but ven- 
tral extending farther forward in correlation with the obliquity 
of the orifice. Atrial orifice scarcely lipped. Body muscles 

Fig. 13.-5. fusiformis-runcinata, aggregate generation. 

seven, never interrupted on dorsum, and never continuous across 
the ventral side. Anterior four, and posterior three, confluent 
on dorsum. Last of anterior group and first of posterior group 
confluent laterally, but the two disconnected ;i1 ends. Posterior 
two, the sixth and seventh, confluent well down the side, the two 
separating, the sixth to pass in front of the nucleus, the seventh 
behind it. The seventh usually confluent with a smaller muscle 
belonging to the atrial orifice. A large muscle band in the dorsal 
lip of the branchial orifice a short distance back from the edge, 
a delicate one at the very edge, and a broad band in the ventral 
lip. An angular muscle band at the angle of the branchial ori- 
fice on each side, its angle directed toward the angle of the orifice, 
and its two limbs directed, the one ventrad, the other dorso- 
posteriad. Endostyle slender, extending far forward under the 
ventral lip to a level with the ventral ends of the sixth muscle, 
some distance in front of the nucleus. Gill rather shorter, rela- 
tively, than in the solitary generation, and making a wider angle 
with the endostyle. Anterior end about middle of the interval 
between the dorsal lip and the anterior group of muscle bands, 
posterior end at the nucleus, hence some distance behind the 
posterior end of the endostyle. "Nucleus" rather small, com- 
pact, egg shaped, situated far back, projecting somewhat from 

Vol. 2] 

Hitter. — The Pelagic Tunicata. 


the general surface of the animal. Color of nucleus orange, 
though not uniform in all parts. Heart between posterior end 
of endostyle and nucleus. 

This is by considerable the most abundant species of salpa 
of the western shores of North America, and probably of the 
whole Pacific Ocean, at any rate north of the equator. It has 
been taken at almost every point on the coast from Alaska to 
Lower California, and at many of these in large numbers. 
On the whole California coast it has been taken in nearly every 
month of the year, though the systematic collecting at San Diego 
thus far indicates it to be considerably more abundant during 
the summer than in the midwinter months. 

Salpa fusiformis-runcinata, form echinata. 


at.o X 



Fig. 14. — S. fusifonnis-runcinata, echinate form. Postero- 
dorsal view, showing serrations of test, and muscle bands. 


p.dp. ' ,' smr' l.r. 

Fig. 15. — S. fusiformis-runoinata, echinate form, ventral view 
of surface. 

g# University of California Publications. [Zoology 

Having worked over a large quantity of material of 8. fusi- 
formis-runcinata with reference to the question of the status of 
echinata, I reach the conclusion that a well marked style, or form 
of the species, which may be called echinata, must be recognized, 
but that it is not a distinct, persistent variety, as Apstein has 
treated it; much less a species as Herdman concluded from his 
study of the Challenger material. 

There are two particulars by which, at their fullest expres- 
sion, the form is distinguished from the typical fusiformis- 
runcinata. One is the ecbination of the test, the other the 
arrangement of the body muscles. Pigs. 14 and 15 are dia- 
grams, though made with special care, from the examination of 
three specimens which agreed almosl perfectly in these partic- 
ulars. Fig. 1-1 is a posterb-dorso-dextral view. It shows two 
double rows of echiualions on the dorsal surface that begin some 
distance behind the anterior end and extend to the posterior end, 
but do not terminate in spines. Laterally from these is a row 
on each side on the edge of a prominent ridge — almost a tin — 
of test, this ridge extending the entire length of the body and 
terminating posteriorly in two prominent processes, r. and i. p. 
d. p. On the ventral side are two sets of submedian rows (Fig. 
15, s. in. r. and s. in. r.' ). an anterior and a posterior, each-set 
open anteriorly but joined posteriorly, the posterior junction of 
the posterior set being in a prominent posterior ventral process 
p. r. p., and laterally from these are again two more rows. As 
to the muscles, the three anterior ones, though converging some- 
what, scarcely touch one another as they do typically in fim- 
formis-runcinata. The eighth and ninth do not even converge; 
they are entirely parallel. Herein is perhaps the most striking 
difference between echinata and the type of the species. 

This description applies exclusively to the solitary genera- 
tion. As to the aggregate generation, one finds an occasional 
lot of zooids that are unusually robust, this being especially 
apparent at the posterior end of the animal, where the posterior 
process of test becomes much thickened and solidified, with the 
serrated edges highly developed. In one lot of this sort observed 
off San Diego in March, 1904, the animals reached a total length 
of 50 mm. or more ; and several zooids in one gathering made 

Vol. 2] Fitter.— The Pelagic Tunicata. 69 

by the Albatross (data as to time and locality lacking, but cer- 
tainly Pacific Ocean material) a total length of 60 mm. was 
reached, the body here, exclusive of the processes, being 40 mm. 
These last were somewhat larger than the largest Challenger 
specimens of echinata. That these robust aggregate zooids belong 
with the echinata form of the solitary generation may be held 
as probable, although nothing less than absolute proof of this 
will warrant associating them positively in classification. 

Now a few more words in support of my opinion that we have 
here a case of extreme, for this group of animals, individual vari- 
ation, or fluctuation, rather than a true variety, or "elementary 
species." In the first place, as to the echination of the test. 
It is doubtful if this is ever wholly absent in S. fusiformis- 
runcinata. Certainly if it is, it is so only exceptionally. So 
far as the evidence goes on this point, it is to the effect that the 
thickening of the test at the posterior end, the prominence of 
the ridges, and the serrations increase with the size, and pre- 
sumably with the age, of the zooids in both generations. But 
more extended and exact information is needed here. There 
are undoubtedly some observations opposed to this supposition. 
For example, I have one specimen of the solitary generation 
taken at Bolinas Bay, California, November 18, 1895, which, 
although scarcely more than half the size of the largest echinata, 
yet possesses the longest, heaviest three posterior processes I have 
seen in any zooids whatever. But here the serrations are almost 
entirely wanting. In this specimen, too, the muscle plan is 
strictly that of fusiformis-runcinata — that is, the anterior three 
and posterior two are fully fused. And here I would say that 
the examination of a large number of specimens with reference 
to the point has failed to discover a single instance of the sepa- 
ration of these muscles in a small zooid. I consequently incline 
to the opinion that the separation of the muscles is an age char- 
acter. But here, too, more positive evidence is needed. My pro- 
visional conclusion is, then, than echinata is an old age form of 

In view of the usually clear delimitation of species in Salpa, 
the question of the status of echinata is especially interesting. 
It well deserves more extensive and critical examination. Ap- 


University of California Publications 


stein's suggestion that we have here a variety that pertains to 
the solitary generation alone is interesting, but can hardly be 
regarded as of much value until established by direct evidence. 
This form has been taken several times at various places on 
the California coast during the last ten or twelve years. It 
would seem to be coincident with the typical fusiformis-ni ad- 
nata in distribution. 

Salpa tilesii-costata (Cuvier-Quoy et Gaim.). 

Salpa tilesii Cuvier, 1804, p. 375, figs. 3-6. 

Salpa costata Quoy et Gaim, 1834, Zool. t. 3, p. 587, PI. 86, 

figs. 1-5. 
Salpa costata-tilesiA Krohn, 1846, p. 114. 
Salpa costata-tilem Traustedt, 1885, p. 379, PI. 1, figs. 10 and 11 : 

and PI. II. figs. 38-41, ami 47. 
Salpa costata-lih svi Eerdman, 1888, p. 60, PI. 4, figs. 1. 4, 8. 
Salpa costata Brooks. 1893, p. 10 (particularly), PI. IV, fig. 4; 

PI. VIII, fig. 4. 
Salpa Tilesii A.pstein, L894, p. 16; 1901, p. Ill 10, figs. 11a, lib. 

Fig. 16. — .S. tilesii-costata, solitary generation. 

(a) Solitary (budding) generation. — Pig. 16. Body much 
Larger anteriorly, tapering gradually back to the region of the 
nucleus, then expanding again rather abruptly to the atrial ter- 
mination. A prominent hump on the ventral side corresponding 
to the nucleus. Anterior end rounded: posterior truncate but 
for slight lateral notches, and armed with a pair of firm com- 
pressed, green edged appendages arising laterally from a little 
in front of the edge of the atrial orifice. Both orifices distinctly 
terminal. Lips of branchial orifice prominent, the ventral some- 
what more so, upper finely serrated on its inner edge. Length 
of longest specimen, 19 cm., exclusive of appendages, which were 
47 mm . Test thick and firm, particularly on the ventral side, 

Vol. 2] 

Fitter. — The Pelagic Tunicata. 


and most of all over the nucleus. Whole surface, except for a 
broad area at the anterior end, beset with low, spine like processes. 
Frequently a girdle of indefinite limitation of dark green in the 
mid body, across the dorsal side and extending well down toward 
the mid-ventral line, the color, which is on the surface of the 
test, being densest laterally. The edges of the posterior append- 
ages densely and uniformly colored with the same green. Body 
muscle bands 20, occasionally 19 or even 18, uniformly spaced 
and mostly all parallel, but the first and second inclined some- 
what backward on the dorsum. All (in old zooids only?) limited 
to the dorsal half of the body, and all interrupted in the mid- 
dorsal line, and typically, the 7th or 8th or 9th interrupted 
laterally. A single broad band in dorsal lip broadly interrupted 
on each side of median line; also a pair of short, longitudinal, 
nearly parallel, widely separated bands in this lip ; two bands 
in ventral lip ; a complicated and variable crossing of short 
bands at angle of orifice. Nine or ten or more delicate, wavy, 
more or less interrupted bands belonging to the atrial siphon. 
Endostyle slender, gently curved to conform to the outline of 
the body, reaching entirely back to the nucleus. Gill relatively 
rather short, extending from a little in front of the first muscle 
band to the nucleus. Hyphysis close to anterior end of gill, but 
short distance in front of ganglion, forming a distinctly pendant 
tubercle, on which the rather large, irregularly triangular mouth 
mouth is situated. 

Fig. 17. — Costata-tilesii, aggregate generaion. 

(b) Aggregate (sexual) generation. — Fig. 17. Body in gen- 
eral cylindrical, but irregular, especially posteriorly from the 

72 University of California Publications. [Zoology 

projection of the intestinal mass and the atrial siphon. Remain- 
ing in the aggregated condition and firmly united at least until 
7 cm. long, and while thus united, body somewhat asymmetrical 
from the mode of aggregation, the branchial and atrial orifices 
being turned respectively to the right or left, depending on 
whether the zooid be right or left in the chain. Largest zooid 
seen, 14 cm. Test rather thick and stiff, particularly on ventral 
side, and most of all over nucleus, where in old zooids it becomes 
opalescent. Surface in some, though not in all, beset with low. 
broad, scattered processes, these on the whole more pronounced 
on dorsum. An irregular area of yellowish green in the test 
over the nucleus, and occasional small patches of this on dorsum. 
Lips of branchial orifice prominent, of nearly equal height, the 
dorsal overarching; ventral projecting forward and below in a 
blunt prow. Atrial siphon narrow, thin walled, elongated, with- 
out lips. Body muscle bands five, limited to the dorsal side, ami 
not extending more than half way down to the mid-ventral line. 
The first three drawing together, bu1 no1 touching on the dorsum, 
and each interrupted by a narrow interval in the mid-dorsal line. 
These three muscles, and frequently the fourth, interrupted on 
the side toward the axis of the chain, hut usually not on the other 
side. Fifth muscle forked on each side. A single strong band 
in the dorsal lip. widely interrupted on both sides of the middle, 
two hands in the ventral lip, the dorsal and ventral lip bands 
crossing and intermingling in a complex hut somewhat variable 
way at the angles of the orifice. Two shorl longitudinal bands 
in dorsal lip. Numerous delicate hands in atrial siphon, all 
confluent with a longitudinal hand on each side. Endostyle 
slender, nearly straight, reaching hack entirely to the nucleus. 
Gill relatively short, scarcely reaching into the anterior third of 
the animal; hypophysis small, hardly recognizable without dis- 
section. Nucleus relatively large, compact, ovate, regular; rectum 
far back, projecting dorsad several millimeters above the gen- 
eral level of the nucleus, greenish brown at its anterior end, 
yellow posteriorly, with an irregular scarlet area on its dorsal 
side and extending somewhat on to the gill. Embryos normally 
four, situated dorsally to the right of the median line, between 
the fourth and fifth bodv muscles. 

Vol. 2] 

Bitter. — lit e Pelagic Tunicata. 


Although this, by far the largest, most magnificent of all 
our species of Salpa, has never been taken in great abundance 
on the California coast, it is by no means rare, since a few speci- 
mens at a time have been frequently collected at numerous points 
during the last fifteen years. The collections show it to have 
been taken in March, May, June, July, August, and November, 
with the largest numbers in March and July. 

Salpa democratica-mucronata Forsk. 

Salpa dcnwcratica Forskahl, 1775, p. 113, PI. 36, fig. G (solitary 

Salpa mucronata Forskahl, 1775, p. 114, PI. 36, fig. D (aggregate 
gener. ) 

Salpa Cabotti Agassiz, 1886, p. 17, figs. 1-5. 

Salpa democratica-mucronata Krohn, 1846, p. 112-113. 

Salpa democratica-mucronata Leuekart, 1854, p. 3, et seq., PI. I, 
figs. 1, 3, and numerous others. 

Salpa democratica-mucronata Traustedt, 1885, p. 365, PI. 2, figs. 
25 and 28. 

Salpa democratica-mucronata Herdman, 1888, PL VIII, figs. 1-10. 

Salpa democratica Brooks, 1893, pp. 6-16, particularly for anat- 
omy. PI. 2, especially. 

Thalia democratica-mucronata Herdman, 1899, p. 748. 

Salpa mucronata Apstein, 1901, p. Ill 5, figs. 5a and 5b. 

Fig. 18. — S. democratica-mucronata, solitary generation. 

(a) Solitary (budding) generation. — Fig. 18. Form ovate, 
the posterior end armed with a pair of long,slender, bilaterally 
placed processes, with a median ventral process, usually bifid, 
the ventral horn much shorter than its mate, which is sometimes 
nearly as long as the laterals; usually with a median dorsal 
process, sometimes of considerable length, but more frequently 


University of California Publications. [Zoology 

short or occasionally wanting. Length of body, exclusive of 
processes, about 8 mm. Test very thick and exceedingly trans- 
parent. A blunt pocket of the mantle reaching into the base of 
the long lateral processes. Branchial orifice inclined somewhat 
upward, with dorsal and ventral lips, the dorsal distinctly higher 
and broadly notched in middle. Atrial orifice nearly terminal, 
but inclined a little upward, without lips. A girdle and half 
another of minute spines on the test encircling the branchial 
orifice, and four bands of similar processes running Lengthwise 
of the animal on the dorsal surface. Body muscle bands six, 
the first interrupted on dorsum; second, third, and fourth in 
contact, though hardly confluent dorsally; fifth and sixth also 
in contact dorsally. Bands all continuous across the ventral 
side, excepting the sixth, which is interrupted. Muscles all deli- 
cate, the sixth particularly so. A single rather broad muscle 
1);iik1 belonging to the branchial orifice situated some distance 
away from the edge of the opening, continuous except for an 
interruption on the ventral side. A few scattered fibres around 
the atrial orifice. Endostyle relatively short, extending back 
only to the third muscle band, consequently leaving a wide inter- 
val between its posterior end and the nucleus, somewhat curved 
and rather thick. Gill extending from a little behind the first 
body muscle band back to the nucleus. Ganglion slightly in 
front of the anterior end of the gill. Hypophysis considerably 

in front of ganglion, and l'r surface view wholly disconnected 

from it: rather small, somewhat triangular in dorsal vieAv. 
Nucleus very small, ovate, situated far back, light yellow. Chain 
of buds encircling the nucleus. Pericardium and heart so deli- 
cate as to be seen with difficulty in preserved specimens. 

Fig. 19.— S. democratica-mucronata, aggregate generation. 

Vol. 2] Bitter.— The Pelagic Tunicata. 75 

(b) Aggregate (sexual) generation. — Fig. 19. Agreeing 
with solitary generation in general form, size, orifices, thickness 
and transparency of test, endostyle, gill, ganglion and hypo- 
physeal organ, and nucleus. The pair of long posterior pro- 
cesses constantly present in the asexual generation, not present 
here, but a few processes into which the mantel extends are found 
on various parts of the body, but wholly irregular as to location. 
Body muscles four, first three confluent dorsally, but each inter- 
rupted ventrally; the fourth bending abruptly forward on the 
ventral side. A band as broad as the body bands at the branchial 
orifice, situated some distance back from the orifice itself, and 
continuous around the animal except for an interruption on the 
ventral side. Laterally on each side a branch given off from this 
band extending postero-dorsad ; also a delicate muscle on each 
side extending from the band toward and around the extreme 
of the orifice; a pair of delicate longitudinal muscles in dorsal 
lip. Posteriorly, a delicate muscle given off from the fourth 
body band on the dorsal side, and extending to a variable extent 
backward and downward in the region of the atrial siphon. A 
number of delicate muscles belonging to atrial orifice. Ovary 
situated on right side, midway between fourth muscle band and 

In treating the muscles of this form I have followed the 
usual custom and enumerated four body muscles only. I strongly 
suspect, however, that in reality we have here six, as in the soli- 
tary form. On this view, the first muscle of the solitary form 
would be represented by the postero-lateral branches, fig. 19, 
of the sexual form ; and the sixth of the solitary form would be 
represented by the delicate band given off from the fourth. 

The species is, next to fusiformis-runcinata, the most abun- 
dant in the area. It has been taken during every month in the 
year in which plankton collecting has been systematically done, 
and in all probability is always present. 

76 University of California Publications. [Zoology 

Salpa zonaria-cordiformis, Pall.-Quoy et Gaim. 

Holotlniriunt zonarium Pallas, 1774, Fasl. X, p. 26, PI. I, figs. 17, 

A, B, and C. 
Salpa polycraiica Forsk., 1775, p. 116, PI. 36, fig. F. 
Salpa sonaria Cham., 1819, p. 12, PI. figs. 1A to G. 
Salpa cordiformis (,)uoy et Gaim., 1327, p. 225, PI. 8A, figs. 3-6. 
Salpa cordiformis-eonaria Krohn, 1846, p. 112. 
Salpa cordiformis-polycratica Vogt, 1854, p. 7. 
Salpa cordiformis-eonaria Traustedt, 1884, p. 382, PI. II, fivs. 

18, 19, 20, and 21. 
Salpa cordiformis-zonaria Herdman, 1888, p. 70, PI. VII, figs. 1-9. 
Salpa cordiformis Brooks, 1893, p. 10 (particularly); PI. Ill, 

figs. 2 and 3; PI. IV, figs. 3, 5, 6; PI. VIII, fig. 5. 
Salpa zonaria Apstein, 1894, p. 19; 1901, p. Ill, 10, figs. 10a, 10b. 

Fig. 20. — S. cordiformis-zonaria, solitary generation. 

(a) Solitary (budding) <i< m ration.- Fig. 20. Body unusu- 
ally firm and hard, probably more so than in any other species; 
form regular, of nearly uniform diameter throughout, though 
slightly larger at posterior end: somewhal quadrilateral, though 
this perhaps due to shrinkage, and may he eharacteristic of pre- 
served animals only. Long axis almosl perfectly straight. Wholly 
colorless except for light lemon yellow of nucleus. Maximum 
Length of zooids so far obtained in our area, 3-1 mm., though a 
length of 60 mm. is recorded by other observers. 

Anterior end strikingly truncate, the orifice being exactly 
terminal, the usual salpa lips scarcely suggested, and the lateral 
angles clear cut and almost exactly right angles. The somewhat 
Larger posterior end produced into three ridged, sharp pointed 
processes, two of which are shorter and dorso-lateraL in position, 
and tlie third, considerably longer, situated in the median ven- 
tral plane and directed somewhat ventrad as well as backward. 

Body muscle hands six in number, exceedingly broad, all 
quite paralLeL with one anotlier in the course around the body; 
all interrupted both dorsally and ventrally, and by about the 

Vol. 2] 

Bitter. — The Pelagic Tunicata. 


same interval, excepting that the interruption of the sixth on the 
ventral side is much greater than that of any of the others. Two 
muscles, the one dorsal, the other ventral, probably representing 
lip muscles of other species, but rather remote from the branchial 
orifice, meeting in an acute backwardly directed angle on each 
side. A short, semi-lunar scrap of muscle on each side, close 
within the antero-lateral angles. The posterior orifice small, 
terminal in position, with rather distinct dorsal and ventral lips, 
each containing a delicate muscle. 

Endostyle straight and slender, extending nearly the entire 
length of the body, but terminating posteriorly somewhat short 
of the nucleus. Gill likewise peculiarly long and slender, extend- 
ing back of the nucleus, hence beyond the posterior end of the 
endostyle. Hypophysis simple, narrowly elliptical, projecting 
but slightly into the pharyngeal cavity, situated exactly in the 
sagittal plane. Ganglion rather small and eye spot scarcely pig- 
mented. Nucleus small, situated very far back in the base of 
the posterior ventro-median process of the body. A pointed 
outpocketing of the mantel projecting behind the nucleus into 
the process of test. The stolon beginning far back and extend- 
ing somewhat backward to wind around the nucleus from right 
to left. 

Fig. 21. — S. sonaria-cordiformis, aggregate generation. 

(b) Aggregate (sexual) generation. — Fig. 21. Body firm, 
from the hardness of the comparatively thin test ; irregularly 
cigar shaped in general outline. Branchial orifice fully on the 
dorsal side and looking directly upward, its two lips prominent 
and equal in size. Posterior end produced into a broad process 
of varying length, in some zooids it equaling half the length of 
the body, while in others it is much shorter, and in still others 
almost wholly wanting. In all the specimens that have come 

78 University of California Publications. [Zoology 

under my observation the posterior process, when present, is 
directed somewhat to the right, with the atrial orifice turned cor- 
respondingly to the left; but this is quite surely correlated with 
the position occupied by the zooid in the salpa chain, and in half 
the individuals the process is probably situated to the left, with 
the atrial orifice to the right. Atrial orifice opening somewhat 
upward, but less so than the branchial. Lips rather prominent 
here also. 

Body muscle bands six. very broad and heavy. First and 
second interrupted dorsally, the first widely so, and all inter- 
rupted ventrally. First and sixth inclined strongly toward the 
middle of the body on both dorsal and ventral sides. A single 
muscle in the anterior dorsal lip. but none in the ventral. A 
few fibres entirely encircling the atrial orifice. First band 
bifurcates on each side. Sixth band bifurcates on righl side (not 
shown in Fig. 21 ), but not on left, this being probably correlated 
with the fact that the post-body process is to the righl side. (The 
asymmetry of this muscle in this species may be compared with 
that of the muscles of Cyelosalpa bakt ri.) 

Endostyle relatively short, rather stout, somewhat curved to 
correspond with the gentle convexity of the ventral surface of 
the zooid: extending from within the prominent extension of the 
body in front of the branchial orifice to the posterior edge of the 
fourth muscle band. Cill likewise rather short, the anterior end 
being in the interval between the second and third muscle bands, 
and the posterior end at the nucleus on a level with the anterior 
edge of the sixth band. Hypophysis a considerable distance in 
front of the anterior end of the gill, narrowly elliptical, project- 
ing but slightly into the pharyngeal cavity. Ganglion situated 
at the immediate posterior end of the hypophysis. 

Nucleus small, ovoid, far back, not projecting from the gen- 
eral surface of the body. Embryos four, situated dorso-laterally 
in the interval between the fifth and sixth muscle bands. 

This is, from its rigidity and angularity, one of the best 
defined species of Salpa in our fauna. Though not abundant, it 
cannot be said to be very rare, since it has been taken at least 
eight times during the last three years, in no instance, however, 
in any quantity. 

Vol. 2] 

Bitter. — The Pelagic Tunicata. 


Salpa cylindrica Cuvier. 

Sal pa cylindrica Cuvier, 1817, p. 22, figs. 8 and 9. (Solitary 

Salpa cylindrica Saw 1816, p. 124, PI. XXIV, figs. 2 1} 2 2 , 2 3 . 

(Iasis cylindrica in description of figures.) 
Salpa cylindrica Traustedt, 1884, p. 377, PI. II, figs. 35, 36, 37. 
Salpa cylindrica Herdman, 1888, p. 72, PI. VII, fig. 10. 
Salpa cylindrica Brooks, 1893, PL III, figs. 5, 6, 7 ; Pl.VIII, fig. 2. 
Salpa cylindrica Apstein, 1894, p. 16. 

Fig. 22. — <S'. cylindrica, solitary generation. 

(a) Solitary (budding) generation. — Fig. 22. Body nearly 
uniform in diameter from end to end, the anterior end slightly 
larger; long axis quite straight; both orifices terminal. Test for 
the most part thin and soft, though in some zooids a pair of 
rather prominent flanges with finely serrated edges situated 
dorso-laterally at the posterior end. Largest zooid seen, 23 mm. 
long. Lips of anterior orifice of equal size. 

Body muscle hands nine, all interrupted ventrally and unin- 
terrupted dorsally, first four confluent dorsally. All the bands 
rather broad and heavy. Two narrower bands in dorsal anterior 
lip and two in ventral ; a segment of a circular band behind the 
angle of the anterior orifice, and a pair of short, longitudinal- 
oblique bands in the dorsal lip. Several delicate bands in each 
lip of the posterior orifice, the dorsal and ventral groups coming 
together on each side in an acute angle. Endostyle nearly 
straight, extending the entire length of the body; rather slender. 
Gill likewise unusually long and slender. Hypophysis long, 
elliptical, situated at the immediate anterior end of the gill. 
Ganglion with the specially conspicuous eye spot situated a little 
nearer the hypophysis than the level of the first body muscle 
band. Nucleus small, spherical, situated a little short of the 

80 University of California Publications. [Zoology 

posterior end of the body, at the angle formed by the junction 
of endostyle and gill. 

Stolon with salpa chain when well developed reaching along 
the entire ventral side of the zooid, parallel with the endostyle. 
(In the specimen from which Fig. 22 was drawn the salpa chain 
was but slightly developed, and hence did not yet present the 
condition described.) 

Fig. 22'. — S. cylindrica, aggregate generation (after Brooks). 

(b) Aggregati (sexual) generation- Fig. 22', copied from 
Brooks. Having seen do specimen of this generation, I give a 
translation of Traustedt's Latin diagnosis: "Body ovate: ex- 
tremities terminating in conical, scarcely elongate appendages. 

Apertures dorsal. Body muscles five, all interrupted on ventral 
side; three anterior and two posterior confluent in median dorsal 
line; all interrupted ventrally." 

This is one of the rarer forms of salpa of the California 
coast. 1 find but four lots of it in all the collections that have 
been made during the last fifteen years, by both the Albatross 
and the University. Two of these were by the Albatross, one 
in January. 1889, the other in April, 1904. The other lots, col- 
lected by the San Diego station, were taken in July, one of 1903, 
the other of 1904. All were from the coast of southern Cali- 
fornia. Curiously enough, all the specimens so far seen, with 
one doubtful exception, were of the solitary generation. 

Salpa confoederata-scutigera Forsk.-Cuvier. 

Salpa confocderata Forsk., 1775, p. 115, PL 36, fig. A. 
Salpa scutigt ra Cuvier, 1817, p. 18, figs. 4 and 5. 
Salpa octofora (?) 2 Cuvier, 1817, p. 20, fig. 7. 

1 While 1 follow Traustedt in regarding S. octofora of Cuvier to belong 
to the present species, the figure and positive statement of Cuvier as to the 
elliptical shape of octofora make it not improbable that a rather distinct. 
octofora variety does exist. 

Vol. 2] Hitter.— The Pelagic Tunicata. 81 

Salpa (Pegea) octofora, Sav., 1816, p. 124, PI. 24, fig. 1.1 and 1.2. 

Salpa ferruginea, Cham., 1819, p. 23, figs. 10A-D. 

Salpa scutigera-confoederata Vogt, 1854, p. 6. 

Salpa scutigera-confoederata Traustedt, 1885, p. 362, PI. II, 

figs. 23, 24. 
Salpa scutigera-confoederata Herdman, 1888, p. 84, PI. IX, fig. 9. 
Pegea confoederata Lahille, 1890, p. 12, text, figs. 1, 2, 3A, 3B. 
Salpa scutigera Brooks, 1893, pp. 6-16, anatomy; PI. IV, figs. 1-7. 
Salpa confoederata Apstein, 1894, p. 12, PI. II, fig. 16. 

(a) Solitary (asexual) generation.-— As I have seen but a 
single small specimen of this generation, I neither figure it nor 
give a diagnosis of it. But these may be the less disadvantage- 
ously omitted from the the fact that the two generations are so 
very similar. The chief differences between them are that the 
general body form is relatively shorter and more rotund in the 
solitary generation, and that the atrial siphon projects farther 
backward from the level of the nucleus in the solitary than in 
the aggregate generation. 

Fig. 23.— S. confoederata-scutigera, aggregate generation. 

(b) Aggregate (sexual) generation. — Fig. 23. Aggrega- 
tions remaining intact at least until zooids are 25 mm. long, but 
at this stage falling apart with great ease. Body cylindrical, 
straight, and regular, the regularity broken only by a slight con- 
striction behind the branchial orifice, by a pair of low, lateral 
prominences in this constriction; and a pair of low, broader, 
blunter lateral test tubercles at the posterior end, into which a 
double outpocketing of the mantle on each side projects. Test 
rather delicate, without specially thickened areas, transparent, 
the iron rust colored pigment characteristic of the species being 
mostly confined to the mantle. 

Branchial orifice terminal, lips not very prominent; atrial 
orifice but little smaller than branchial, directed slightly upward, 
with well defined lips. Largest zooids, 70 mm. long. A brick 

82 University of California Publications. [Zoology 

red or iron rusty pigment widely present in the mantle, but most 
pronounced at two ends, and more abundant posteriorly than 

Body muscles four, all confined to the dorsal half, the first 
two bent toward each other and nearly touching in the mid-dorsal 
line; the last two likewise similarly inclined, sometimes connected 
by a short ridge in the mid-dorsal line, and sometimes not in 
contact. A single band in the dorsal lip and two in the ventral. 
A pair of angular Viands on the dorsal side behind the branchial 
orifice, the apices directed backward, and the dorsal limb of each 
leaching well into the dorsal lip to serve as a retractor of 
it. (Dorsal limb unfortunately omitted in Fig. 23.) A 
short oblique band on each side, near tangential with the apex 
of these angles. A single band in each of the posterior lips, 
nearly as strong as the body muscles. Endostyle slender, 
straighl excepl for a distinct ventral curvature jusl short of the 
posterior end. the posterior termination separated from the 
nucleus by the Length of the pericardium. ('ill relatively rather 
short, beginning but little in front of* the lirst body muscle, which 
is unusually far back, and not reaching to the nucleus poste- 
riorly. Hypophysis large, irregularly triangular, placed vertic- 
ally, the opening anterior, suspended as a prominent tubercle 
immediately at the anterior end of the gill, and ventrad of the 
ganglion, the hypophysis and ganglion being consequently in 
unusually close relation. Nucleus subglobular, small, slightly 
compressed dorso- ventral ly, projecting but slightly, if at all, from 
the general ventral surface of the animal, but extending back- 
ward fully even with the posterior extremity of the atrial siphon, 
its color nearly uniform seal brown. 

The aggregate generation is not rare in the urea, but not a 
single zooid of the solitary generation has thus far been observed. 
The species has been taken on the California coast from Monterey 
Bay southward, and in the months of January. February, March, 
May. dune. duly, and August. 

Fam. II. — Doliolidae Bronn, 1862. 

Thaliacea in which the body form is typically and perfectly 
that of a barrel; the body muscle bands are complete rings; and 

Vol. 2] Hitter.— The Pelagic Tunicata. 83 

the branchial apparatus consists of a membranous partition be- 
tween the pharyngeal and atrial ends of the animal, pierced 
by pairs, never very numerous, of stigmata, Both branchial and 
atrial orifices terminal and at opposite ends; both encircled by 
lobes; those of the branchial orifice always more prominent. 
Branchial tentacles and dorsal lamina wanting. Intestinal tract 
small but never nueleaform; situated on ventral side of body, 
immediately behind the branchial membrane. Hermaphroditic, 
the gonads of both sexes always simple ; ovary never producing 
a large number of ova. 

In addition to the sexual method of reproduction, in which a 
larval stage of the typical tunicate tadpole is passed through, a 
process of gemmation prevails in which three polymorphic gen- 
erations occur. 

Genus Doliolum Quoy et Gaimard, 1835. 

With characters of the family. (The genera Anchinia and 
Dolcliinia not occurring, so far as known, in our area, are not 
taken into account in the diagnoses. ) 

Three species, namely, D. tritonis, D. ehrenbergii and D. mul- 
leri, are here recognized as belonging to our fauna. Whether 
certain forms of doubtful status, not now treated, belong to one 
or another of these species, more extended observation in the 
future must determine. 

Owing to the fact that zooids of the generations produced by 
budding, far advanced in development, rarely reach the hands 
of the student still attached either to their parent or to one 
another, the difficulty of arriving at unquestionable conclusions 
as to how the various forms of the different species go together 
is great. As to the old "nurses," i.e., oozooids in which the 
internal organs have wholly disappeared through degeneration, 
and the muscle bands have become greatly broadened, we are not 
yet in a position to specify with absolute certainty the specific 
characters of this form in a single species of the genus. My 
diagnoses of this generation in each species relates almost wholly 
to the young zooids before the degradation of the organs lias 

g4 University of California Publications. [Zoology 

The relations of the several generations in the life cycle, as 

now understood, may be indicated by the following scheme, made 

without regard to theoretical views as to the exact nature of the 

relationships : 


Embryo = Beginning of cycle 






Embryo = Beginning of cycle 

A greal variety of nomenclature has been employed by dif- 
ferent writers to designate the several generations of Doliolnm. 
That here used, though not without objections, appears to me 
to be the best of any ye1 proposed. It is in the main from Herd- 
man, L888 and 1904. 

The chief synonyms found in the Literature of the subjecl 
are as follows : 

Gonozooid (Herdman) ; sexual animal (Krohn, Grobben, 
Uljanin) ; sexual generation A (Keferstein and Ehlers). 

Ooozooid (Herdman): asexual animal (Krohn, Gegenbaur) ; 
generation B (Keferstein und Ehlers); first nurse generation 
(Grobben) ; nurse (Uljanin) ; blastozooid (Herdman, '88). 

Trophozooid I Herdman) ; lateral buds, generation C 1 (Kefer- 
stein und Ehlers) ; lateral buds (Grobben) ; nutritive animals 
(Fol. Uljanin). 

Phorozooids (Herdman); median buds (Krohn, Gegenbaur, 
Grobben) ; generation C m (Keferstein und Ehlers) ; foster-ani- 
mals (Uljanin). 

Vol. 2] 

Bitter. — The Pelagic Tunicata. 


Doliolum tritonis Herdman. 

Doliolum denticulatum Herdman, 1883, p. 101, Pis. 18, 19, an. I 20. 
Doliolum tritonis Herdman, 1888, p. 47, PL 3, fig. 3. 
Doliolum tritonis Traustedt, 1893, p. 4, PL 1, fig. 10. 
Doliolum tritonis Borgert, 1894, p. 19, PL 5, figs. 17 and 18. 
Doliolum tritonis Borgert, 1901, p. Ill 3, fig. 3. 

Fig. 24. — Doliolum tritonis, gonozookl. 

(a) Gonozooicl. — Fig. 24. Length, 15 mm., 12 mm., 12 mm.. 
13 mm., 14 mm.; form very regular; test always very thin, and 
general consistency delicate. Lobes of branchial orifice prominent, 
rounded, somewhat broader than high, apparently always twelve : 
those of atrial orifice much less pronounced, usually two. Muscle 
bands eight, first and eighth considerably smaller, all excepting 
these two with a narrow line running lengthwise along the 
middle. Atrial orifice with ten very low, broad lobes, also 
rounded. Endostyle rather slender, extending from the middle 
of the second intermuscular space to a little beyond the middle 
of the fourth space; anterior end pinkish yellow in many speci- 
mens. Ganglion in the third intermuscular space, beyond the 
third muscle band about the width of the band. Hypophyseal 
duct long and slender, the mouth, which is rather small, situated 
a little in front of the middle of the second intermuscular space. 
Peripharyngeal band trending forward from the endostyle on 
each side in a broad curve which reaches in front of the second 
muscle band, then running backward again on the dorsal side 
of the right limb, passing into a broad open spiral behind the 
hypophyseal mouth. Dorsal limb of the branchial series begin- 
ning on a level with the ganglion a little behind the third muscle 

86 University of California Publications. [Zoology 

band, and extending back to a little behind the sixth band. Ven- 
tral limb beginning slightly behind the fourth *band. About 
seventy long stigmata on each side in the dorsal limb in the 
largest individuals, and about fifty in the ventral limb, but the 
number apparently varying with size and age. Esophageal 
mouth much nearer the ventral than the dorsal side of the poste- 
rior pharyngeal wall, rather large, broad elliptical, its anterior 
edirc .m ;i level with the fifth muscle band. Stomach large, irreg- 
ularly globose, in the fifth intermuscular space, its posterior end 
on a level with the sixth muscle band. Intestine compressed 
dorso-ventrally, in the form of a broad spiral of a little more 
than one turn, the axis of the spiral directed dorso-ventrad. Anus 
in the posterior half of the sixth intermuscular space. Stomach 
and the pyloric enlargement of the intestine brick red in many 
specimens, but apparently not in all. Ovary latero-ventral, 
immediately in front of seventh muscle band. Testis long, irregu- 
larly cylindrical, situated on the left side; posterior end near 
the ovary, anterior end on left side, usually in fully developed 
state slightly behind the second muscle band, but the termina- 
tion variable in position. Anterior end frequently, though not 
always, with pronounced hook. 

(b) Ooznoid.— ^ee Figs. L'T. L'S. and 29 of D. elm nbergii and 
iniilhri. this generation, which have the essential features of the 
present species.) Body of the usual doliolum form; muscle hands 
nine, first and ninth relatively very narrow, seventh hand alone 
interrupted and extending into dorsal process. Lobes of branchial 
orifice prominent, usually ten in number, occasionally eleven 
or twelve: atrial lobes very low or wholly wanting. Stigmata 
eight? i surely not more), in four pairs. Endostyle, peri- 
pharyngeal band and hypophysis as in gonozooid and phorozooid. 
Ganglion situated in fourth intermuscular space: otocyst large. 
on left side in third intermuscular space at anterior edge of third 
muscle band. 

Proliferous stolon on ventral side in fifth intermuscular 
space: dorsal process varying from short conical in small zooids 
to long and slender in large ones. Intestinal tract similar to 
that of gonozooid, excepting that the intestine proper is, as com- 
pared with the other parts of the tract, much shorter and forms 

V0L - 2 J Bitter.— The Pelagic Tunicata. 87 

less of a spiral, the length of the loop of the intestine being 
scarcely greater than that of the esophagus. Degeneration of 
the interna] organs, with the simultaneous increase in width of 
the muscle bands apparently beginning considerably earlier in 
this than in either of the other species — D. mulleri and D. ehren- 
/x rgii, with which it occurs in this region. Old individuals (of 
this species, probably) reaching a length of 25 mm., with thick- 
ness of 7 mm., while length of individuals just before beginning 
of degeneration of internal organs, 1.38 mm. 

While the evidence that the oozooid here described belongs 
to tritonis is not complete, it is sufficient to admit of little doubt ; 
indeed, there is with this, as with other species here treated, an 
element of uncertainty about the assignment of the oozooids to 
their proper species, since in no one of them have I seen the ori- 
gin either of the embryo from the egg, or the production of the 
phoro- and gonozooids by the oozooid. My chief reliance for 
the identification has been the intestinal tract. Those oozooids 
with a tract entirely similar to that of the gonozooicl of D. ehren- 
bergii I assume to belong to this latter species. Similarly, those 
in which the organ is the same as that in D. mulleri, it is assumed 
belong to that species. Now it is true, as indicated in the diag- 
nosis, that although the intestinal tract of the animal at present 
under notice differs somewhat from that in the gono- and phoro- 
zooid of tritonis, nevertheless its resemblance to the tract of 
tritonis being closer than it is to that of either ehrenbergii or 
mulleri, the only other species thus far found in this area, the 
conclusion that the animals belongs to tritonis seems justified. 

Another point that, on the whole, speaks for the same con- 
clusion is the character of the lobes of the atrial orifice. These 
are so low in both the gono- and 'phoro- zooids of tritonis as to 
render them scarcely recognizable. This is likewise true of the 
oozooid now being considered. In fact, only in an occasional 
specimen have I been able to see anything at all like lobes. True, 
the atrial lobes are considerably less easily seen in all the species 
with which I am familiar than are the branchial ; but in the 
present zooid their conspicuousness is less than in the corre- 
sponding zooids of any other species. Another consideration 
that I believe points in the same direction is the great size of 

University of California Publications. 


an oozooid constantly oeeurring in our area, which is probably 
the senescent stage of this generation of tritonis. Many of these 
specimens reach a length of 125 mm., or even more, and a thick- 
ness of 7 linn. This is near the maximum size recorded for any 
Doliolum. When it is recalled that the gono- and phoro- zooids 
of tritonis are likewise the largest for these generations, of any 
species, the reasonableness of the supposition that this Large 
nurse belongs 1<> tritonis is apparent. It must be said, however, 
that this line of consideration really has less weight than it might 
seem to have, since we are entirely without information as to 
the size that may he attained by the oozooids of any of the species, 
n.janin, '84, has recorded the occurrence in the Mediterranean 
of a nurse measuring 30 mm. in Length, and this he assumes 
belongs to />. ehrenbergii. The identification is, however, by no 
means certain. As a matter of fact, in the presenl state of knowl- 
edge, l do not believe it possible to assign with certainty any 
oozooid. after the defeneration of its organs is complete, to its 
species: and since the degeneration is complete at an early time 
in all the species, so far as known, the difficulty of disposing of 
t hese old nurses is obvious. 

Vig. 25. — D. tritonis, Phorozooid. 


(c) Phor&zooid. — Pig 25. This differs in no particular from 
the sexual generation except in the absence of the sexual glands 
and the presence of the ventral process. This latter is short and 
thick, is situated in the median ventral line in the sixth inter- 
muscular space a little in front of the seventh muscle band. The 
following lengths of the zooids are typical: 12 mm., 11 mm., 
11 mm.. 11.5 mm.. 12.5 mm.. 13 mm. The gonozooids and phoro- 

Vol. 2] Bitter.— The Pelagic Tunicata. 89 

zooids have been taken together in great numbers off San Diego, 
particularly in May and June. The following table shows the 
number of each generation in lots taken at random from collec- 
tions made in May, June, and July, 1904. 































In view of the constant great preponderance in individuals 
of the sexual generation here, and the entire absence of any other 
than the sexual generation in the vast quantities of specimens 
taken by the "Triton" in the north Atlantic, it would seem 
that this generation is normally more abundant in individuals 
than is the phorozooid generation. Borgert, 1894, reports both 
generations from the Plankton Expedition, but gives no informa- 
tion as to the relative numbers of each. 

Fig. 26. — D. tritonis, Trophozooid. 

(d) Trophozooid. — Fig. 26. General form, the peduncle dis- 
regarded, that of a truncated cone with an oblique base ; intest- 
inal tract extending prominently behind the level of the endo- 
style. A maximum, so far as known, of 25 pairs of branchial 
stigmata. Peduncle very broad at distal end and narrowing 
toward the body of the zooid; length of largest specimens seen, 

90 University of California Publications. [Zoology 

3 mm. : 2 mm. broad in broadest part. Endostyle forming 
considerably less than a right angle with the main axis of the 
1>ody. A single short, blunt test process on the anterior half of 
the dorsal edge of the body. 

The above are the characters which seem to distinguish this 
from any hitherto described trophozooid. I am unable to find 
lobes about the branchial orifice, but, owing to the slight develop- 
ment of these in other species, and to the fact that I have had 
only preserved specimens to examine, and but few of these, I 
assume that they are present but escape observation. 

Although no mantle muscles are present that are not found 
in some, at any rate, of the other described species, they all would 
appear to be particularly well developed here. This is espe- 
cially true of the anterior, a.m., and posterior, p.m. prebranchial 
bands. The sharply cut-off ending of these muscles is deserving 
of notice. The dorsal muscle, dm., extends back to about the 
middle of the length of the body and terminates, not in a sharp 
cut-off, but in a point by a gradual running out. The stalk, or 
peduncle muscle, is also well developed. The ganglion, g., and 
hypophysis. ////., need no special comment for the present pur- 
pose. The peripharyngeal band,, is very close to the 
branchial orifice, and, from the great size of the orifice, is very 
long. At the great bend of the intestine there occurs a patch 
on the intestinal wall, made up of round bodies, probably large 
cells, gl.b. These are presumably glandular, though they may 
be excretoiy. Two other well defined patches, of different struc- 
ture, however, from the one just described, occur on the rectal 
portion of the intestine, p. and p' . The nature of these is un- 
known. Many points in the structure, and particularly the 
physiology, of this generation of Doliohtm are in urgent need of 
further study. 

The only specimens of fully grown trophozooids detached from 
their nurse thus far taken in our area were the ones here de- 
scribed. They were secured by Professor H. B. Torrey at San 
Clemente on June 9. 1896. A piece of dorsal process 8 or 10 cm. 
long, bearing the trophozooids, was secured. The evidence, there- 
fore, that it belongs to D. tritonis is not direct. I infer that it 
does from the following considerations : It clearly does not belong 

Vol. 2] Bitter.— The Pelagic Tunicata. 91 

to ehrenbergii or mulleri, as comparison with published descrip- 
tions of the trophozooids of these species shows. D. tritonis being 
the only other species known to occur in the area, and at the 
same time the most abundant of all the species, would be the 
one to which it would seem most likely to belong. The large 
size of the zooids and the length of the dorsal process, as inti- 
mated by the fragment secured, tends to support the same con- 

D. tritonis is the most abundant, as it is the largest, spe- 
cies in the area. The gono- and phoro- zooids were taken in 
great abundance at San Diego in May and June of 1904. The 
giant oozooids, which I have supposed to belong to this species, 
occurred in particularly large numbers at Avalon in July, 1901. 
The gonozooids and oozooids have also been taken, though in 
less quantity, in March, October, and November. 

Doliolum Ehrenbergii Uljauin (notKrohn). 

Doliolum Gegenbauer, 1856, PI. XVI, figs. 12 (?) and 13 (?), 
and fig. 15. 

Doliolum Gen. 2B and 4B, Keferstein und Ehlers, 1861, p. 68, PI. 
IX, figs. 5 and 7 ; and PI. X, fig. 4. 

Doliolum denticulatum Grobben, 1882, pp. 23-41, Pl.l, figs. 3, 4, 
and 5; PI. 2, figs. 7 and 8. 

Doliolum ehrenbergii Uljanin, 1884, pp. 132-133, PI. 5, figs. 1 and 
3; PI. 12, fig. 8. (Numerous other figures are assigned to 
, ehrenbergii, but I here refer to only those that undoubt- 
edly represent the species as here understood.) 

Doliolum ehrenbergii Herdman, 1886, p. 46, PI. Ill, figs. 5 and 7. 

Doliolum ehrenbergii Lahille, 1890, p. 65, figs. 47, 48, 49, arid 50. 

(a) Gonozooid. Unknown. 

Fig. 27.— D. ehrenbergii, oozooid, before loss of internal organs. 

(b) Oozooid. — Fig. 27. Length from .925 mm. to 2.77 mm., 
between disappearance of larval characters and beginning of 


University of California Publications. [Zoology 

degeneration of intestinal tract. Form variable, from strongly 
bulged in middle region to relatively narrow here. Muscle bands 
nine, first and ninth so much smaller than the others, both as to 
size of muscle and diameter of ring, and so near the adjacent 
muscle as to be easily overlooked. Branchial orifice with from 
eight to ten moderate lobes of unequal size: atrial with usually 
twelve lobes. Endostyle rather thick, extending from near the 
second muscle to the fifth. Stigmata eight, large, in two sets 
of two pairs each, one set dorsad of esophageal mouth, the other 
ventrad; dorsal stigmata between sixth and seventh bands, ven- 
tral opposite fifth. Esophageal mouth large, with thick lips; 
situated near center of posterior end of the pharynx, in the 
space between the fifth and sixth muscle bands. Stomach glob- 
ular, situated under and extending in front of the sixtb muscle 
band. Intestine rather long, extending hack nearly to eighth 
muscle; slightly curved, the convex side turned ventrad. Gan- 
glion large, between fourth and fifth muscles; hypophyseal duct 
running forward, its mouth ltd ween third and fourth muscles. 
Otolith sac on left dorsal side in third intermuscular space. 
Dorsal process before beginning of degeneration of internal or- 
gans generally short, with a constriction which sets off a top- 
shaped terminal portion. Proliferous stolon, without distinctive 
features so far as examined. 

Fig. 28. — V. ehreribergii, oozooid, after complete defeneration 
of internal organs. 

The degeneration of the viscera appears to be inaugurated 
relatively considerably later in this oozooid than in that of tri- 
tonis and miilleri, and this accounts for the fact that while tri- 
tonis oozooids are much more abundant than those of the present 
species, one finds fewer by considerable of them with the organs 
still intact than he does of the gegenbaurii zooids in the same 

Vol. 2] Bitter.— The Pelagic Tunieata, ' 93 

condition. The following table shows the length of the animal 

and the width of the muscle bands, and the state of the viscera, 

in seven specimens observed at the San Diego laboratory in June, 


No. Lengb of Zooid. Width of Muscle. Condition of lntestirfe. 

1 2.22 mm. .20 mm. organs intact 

2 2.59 mm. .18 mm. organs far degenerated 

3 2.60 mm. .129 mm. organs intact 

4 2.77 mm. .148 mm. organs intact 

5 3.33 mm. .27 mm. organs nearly gone 

6 4.25 mm. .37 mm. organs wholly gone 

7 5.00 nun. .55 mm. organs wholly gone 

While, owing to the difficulty in getting accurate measure- 
ments, and of expressing in precise terms the stage of degenera- 
tion of the organs, and probably, more than all, to individual 
variation, such data as this are not very significant, they show 
in a general way what is undoubtedly true; viz., that the degen- 
eration begins relatively late here, and then, that the increase in 
size of the zooid and width of the muscle bands go on pari passu 
with the rather gradual degeneration and resorption of the 
internal parts. 

A few remarks must be made concerning my position with 
reference to the status of D. ehrenbergii and the representatives of 
the four generations assigned to it. First, in regard to the name. 
I agree with the proposal of Borgert, '94, and the practice of 
Lahille, '90, that if such a species as ehrenbergii is to be recog- 
nized at all. it should be Uljanin 's, and not Krohn's. Krohn, 
'56, proposed this specific name for what he regarded as Quoy 
et Gainard's D. denticulatum, on the wholly arbitrary and un- 
permissable ground that Quoy et Gainard's name was "unpas- 
send, " since other species as well as this are denticulated about 
the branchial orifice. Since no author, so far as I am aware, 
between Krohn and Uljanin applied the name gegenbaurii either 
to D. denticulatum or to any form supposed to belong to this 
species, the real cjuestion is, Do all the forms assumed by Uljanin 
to belong to D. denticulatum, and hence called by him gegen- 
baurii, actually belong to one species, or was he in reality dealing 
with generations representing two species, one of which was 
denticulatum and the other an undescribed, or at least an un- 

94 University of California Publications. [Zoolog? 

named species? I agree with Borgert that the latter is the case. 
Since, consequently, D. gegenbaurii Uljanin is only in part a 
synonym, the canon of nomenclature, "once a synonym, always 
a synonym,'' does not apply, and the name may stand for the 
unnamed forms with which Uljanin was dealing. That this 
unnamed oozooid which he had was the same as the "Doliolum 
sp." of Gebenbaur. '56, and shown in his PI. XVI, fig. 15, I have 
little doubt. I am strongly of the opinion, too, that "Gen. 2P> and 
4P>" of Keferstein und Ehlers, and shown in their PL IX, figs. 
5 and 7, and PI. X. fig. 4. likewise belong to the same species, as 
do also Grobben's D. denticulatum, shown in his PI. T, figs. 3 
and 4, particularly, and pretty certainly also in fig. .">. As to his 
fig. 7, PI. II, I am in considerable doubt. 

The species is by no means uncommon off San Diego during 
the summer months, though it has not been taken in "swarms." 
as lias />. tritonis. So far il has not been taken in the fall and 
winter months, unless some of the lew old OOZOOids at present in 
doubt as to species belong here. 

(c) Phorozooid. Probably similar to the gonozooid, with the 
exception of the absence of sexual organs and presence of the 
ventral process, but it is doubtful if fully developed specimens 
of this generation are known. 

(d) Trophozooid. Not known with certainty in the fully 
grown state, but probably shown by Grobben, '82, as he himself 
believed, in his PI. II. fig. S. As I have not seen specimens of this 
generation except as very young buds still attached to the dorsal 
process of the oozooid. I do not give a diagnosis and figure of 
it, but refer to the above mentioned figure by Grobben, assigned 
by him to D. denticulatum. The form described and figured by 
this author apparently differs from the trophozooid assigned by 
me to I), tritonis, fig 26, in the folloAving particulars: It is some- 
what broader in proportion to its length, particularly at the 
anterior end; it has a somewhat less number of branchial stig- 
mata, the maximum reported for it being eighteen (Gegenbauer, 
'56), while the tritonis zooid has at least twenty-three; it has 
test processes at the posterior end as well as at the anterior, 
where alone, so far as we know, one is present in tritonis; and 
these processes are more filiform than in tritonis. 

Vou 2 1 Bitter. — The Pelagic Tunicata. 95 

Doliolum miilleri Krohn. 

D. miilleri Krohn, 1852, p. 58, PI. II, fig. 4. 

D. vordmanni Krohn, 1852, p. 59, PI. II, fig. 6. 

Doliolum sp. Gegenbauer, 1856, PI. XV, fig. 8. 

Doliolum Gen. 3B, Keferstein unci Ehlers, 1861, p. 68, PI. X, fig. 3. 

Doliolum miilleri Grobben, 1882, pp. 55-65, PI. II, figs. 9 and 10; 
PI. Ill, figs. 14, 15, 16, 17, and 18; PL IV, figs. 21 and 22. 

Doliolum miilleri Uljanin, 1884, pp. 127-130, many figures of ana- 
tomical and developmental detail, and in addition the fol- 
lowing of special importance for identification: PI. 4, figs. 
1, 3, 4, and 5, larvae; PI. 7, fig. 11; PI. 8, fig. 10; PI. 9, 
fig. 6; PI. 11, fig. 9; PI. 12, figs. 2, 3, and 4. 

This is the least common of the three species of Doliolum thus 
far observed in our area. Hardly more than a dozen specimens 
all told have been taken. These have all been oozooids, and as 
only a portion of them were still in possession of their internal 
organs, my material for study has been scant. Since, however, 
the other generations are sure to turn up some time, as collecting 
goes on, 1 give the diagnosis of all the generations, relying on 
Uljanin. the most recent writer, chiefly, for all the generations 
except the oozooid, this being made mainly from my own obser- 

(a) Gonozooid. — Length about 3.5 mm., sometimes reaching 
4 mm. ; mantel soft and sticky, in consequence of which surface 
is always covered with foreign particles ; muscle bands extremely 
small ; gill membrane extending from above opposite the fifth 
muscle band downward and forward into fourth intermuscular 
space; pierced by from ten to twelve branchial stigmata. Endo- 
style extending from slightly in front of third muscle band to 
a little in front of the fifth band. Intestinal tract U-shaped, 
esophageal opening near the center of the branchial membrane, 
the entire loop being situated in the fifth intermuscular space. 
Testes pear-shaped, in fully developed state thrusting out the 
body wall into a hillock; situated alongside the intestinal tract. 
Ovary close behind the testes, containing but very few ova; ova 
maturing earlier than the sperm. 

Color markings: Intestine violet or rose; orange red pigment 
spots on the edge of both branchial and atrial orifices. 


University of California Publications. 


Pig. 29.— D. nuilliii. oozooid, before degeneration of internal 


(b) Oozooid. Fig. 29. Maximum Length before beginning of 
degeneration of intestine something less than - mm.; test rela- 
tively thick ;iml soft ; muscle bands nine, without distinctive 
characters as compared with same generation of other species. 
Branchial orifice with normally ten lobes: atrial with twelve, 
the former distinctly more prominent. Branchial membrane 
with its stigmata, endostyle, peripharyngeal band, ganglion, 
hypophysis, and otocysl differing in no easily recognizable way 
from the corresponding parts in />. ehrenbergii. Intestinal tract 
U-shaped, esophageal mouth very near the center of the branchial 
membrane, whole trad in fifth intermuscular space. 

Dorsal process shorl and thick, with but few buds, sometimes 
having a pronounced kink near the base, hut never, so far as 
observed, with the top-shaped terminal piece characteristic of 
D. ch)-< ii Ik rgii. 

In this, as in the other species, little of positiveness is known 
about the old "nurses." Several times catches containing 
oo/.ooids with internal organs intact have also contained several 
scenescent specimens of medium size and having the form of the 
old-style, bell-mouthed cannon shown by Keferstein und Ehlers, 
"Gen. IB," PI. X. fig. 1. These I suspect belong to the present 
species. I do not imagine this peculiar form to be wholly dis- 
tinctive. It is probably due to contraction ; but it certainly does 
not occur frequently in old nurses of what I suppose to be tri- 
tonis and ( Jin nbergii. 

Vol. 2] Bitter.— The Pelagic Tunicata. 97 

(c) Phorozooid. — Entirely similar to gonozooid except for 
absence of gonads and presence of ventral process which is rela- 
tively long and cylindrical. 

(d) Troplwzooid — Small, but long in proportion to breadth, 
and with a relatively long peduncle. 

As already said, D. mulleri is the least abundant of our spe- - 
cies. Thus far it has been taken in mid-summer only, off San 

Order II.— ASCIDIACEA de Blainville, 1827. 

Pelagic or sedentary urochorda, with proportionally very 
large branchial sac having many stigmata. Branchial and atrial 
openings not at opposite ends of body, except in Pyrosoma. 
Test in most cases large in quantity and forming a common 
matrix in which the zooids are embedded in most colonial forms. 
A free swimming larval or "tadpole" stage in the life career of 
nearly all species, this undergoing a profound metamorphosis 
to give rise to the adult. An asexual reproduction by budding 
in many, but no true alteration of generations or polymorphism, 
as in the Thaliacea. 

Pam. Pyrosomidae T. Rupert Jones, 1848. 

Pelagic colonial Ascidiacea, with the colony in the form of a 
hollow cylinder closed at one end. Zooids embedded in the thick 
test constituting the greater part of the wall of the cylinder, and 
so arranged that the branchial orifice opens on the external sur- 
face of the cylinder, while the atrial orifice opens into its inte- 
rior; branchial and atrial orifices consequently at opposite ends 
of the zooid. Branchial sac very large, the stigmata placed per- 
pendicularly to the endostyle, each extending from the endostyle 
to near the mid-dorsal line. No peribranchial chamber present, 
the atrial orifices opening directly into the great common cloaca, 
which constitutes the hollow of the cylinder of the colony, as 
above indicated. Embryo arising from the egg, known as the 
" cyathozooid, " giving origin, by a peculiar process of trans- 
verse fission, to the first four blastozooids, from which as the 
starting point the remainder of the colony arises by typical ascid- 
ian budding. Species all, so far as known, highly phosphorescent. 


University of California Publications. 


Genus Pyrosoma Peron. 

But one genus in the family, hence generic characters same 
as those of the family. 

Pyrosoma Peron, 1804, p. 437, PI. 72, and all succeeding writers. 

Pryosoma giganteum Lesueur. Fig. 30. 

P. giganteum Lesueur, 1815, p. 70, PI. I, figs. 1-15. 

P. giganteum Savigny, 1816, p. 207, PI. IV, fig. 7. and Pis. XXII 

and XXIII, many figures. 
/'. giganti inn Keferstein and Killers, 1861, pp. 72-77, PI. XII, figs. 

4, 5, 7, and 8. 
/'. giganteum Panceri, 1872, pp. 1-25, Pis. I and II. 
/•. giganteum Eexdman, 1888, pp. 26-29, PI. I, figs. 4-21. 
P. giganteum Seeliger, 1895, pp. 61-62, Pis. I and II; PI. IV, 

figs. 3 and 4. Fig. 30. 

fig. 30.— Pyrosoma giganteum colony, 4.5 cm. Long. 

Colony cylindrical, or narrowing slightly toward the closed 
end: quite rigid from the firmness of the test; surface bearing 
prominent test processes, these varying much in number, size and 
form, but on the whole inclined somewhat toward the open end 
of the colony, the atria] orifices of the zooids corresponding to 
the processes being on the more convex side of the processes. 
The processes. ;is ;i rule, with an oblique, more or less distinct 
plane, with finely serrate edges ;it their summits. A velum or 
diaphragm of test shutting off to a variable extent the opening 
of the colony. .Measurements of three Largesl preserved colonies: 
first, length 25 cm., greatest thickness near open end L-'..~> cm.: 
second, Length 25 cm., greatest thickness 3.5 cm.; third, length 
19 em., greatest thickness about middle of length, 3 cm.; largest 
colonies observed, measured in life, 60 cm. long. 40 cm. Long, 
and 35 cm. long. Thickness of wall of largest colonies about 
o' mm. Color varying from an entire absence of pigment and ex- 
treme transparency to livid pink, due to pigment in the oozooids. 

Vol. 2 J 

Bitter. — The Pelagic Tunicata. 


Fig. 31.— Single zooirt of P. giganteum, with the test 
process on which the branchial orifice opens. 

Ascidiozooids. — Fig. 31. Varying in length, depending on 
the length of the branchial siphon, but branchial sac in full 
grown zooids quite constantly about 3 mm.; length, including 
both siphons, 5 mm. to 6 mm. The large branchial sac much com- 
pressed, varying in form from approximately square with some- 
what rounded angles, to pronouncedly elongate. Branchial ori- 
fice in some zooids but slightly above the general level of the test 
surface; while in others it is high up on the above described 
oblique planes of the test processes. In these latter zooids the 
branchial siphon is long and narrow. Branchial tentacles 
close within the branchial orifice, however long this may be; 
tentacles few and short, a single one on the ventral side distinctly 

Largest number of branchial stigmata seen, thirty-two, on 
each side of sac ; maximum number of internal branchial vessels, 
fifteen. Dorsal languets, six to eight. Intestinal tract project- 
ing but little behind branchial sac; esophageal mouth at the 
dorso-posterior angle of the branchial sac, esophagus nearly as 
long as the loop of the intestine proper ; stomach globular or egg- 

100 University of California Publications. [Zoology 

shaped; intestinal loop narrow, the amis situated on left side of 
the stomach. 

Testis rosette shaped, with about ten lobes; when fully ripe 
projecting prominently from the surface of the body on the 
ventral side, a short distance behind the proliferous stolon, which 
is between it and the endostyle; in younger zooids the testis not 
projecting thus from the body surface. The ovary, with its one 
large ovum, closely associated with the testes. 

Although I have decided, alter much perplexity, to call our 
one species of Pyrosoma, I', giganteum, I must confess that the 
decision as between giganteum and atlanticum lias little more 
value to my mind than it would have bad it been made by throw- 
ing dice. Having no examples of atlanticum at hand for com- 
parison, I have been obliged to depend upon published descrip- 
tions of this species, and at almost all points at which authors 
make specific differences between atlanticum and giganticum 
I find, among the large number of specimens at my disposal, 
agreements with both, and complete gradations from one to the 
other; in the ease of the zooids, often within the same colony, in 
fact. I am compelled to question the actual existence of both 
atlanticum and giganteum as distinct species. For example, 
Savigny, 18l6, was first to emphasize difference in form of the 
colony as being distinctive of the two species, he stating the 
atlanticum colony to be conical, and that of giganteum cylin- 
drical. This difference appears to have been chiefly relied upon 
by llerdman. '88, '92, for distinguishing the two species. I find 
colonies that would certainly have to be described as cylindrical, 
and others that would as surely be regarded as conical. But 
there are numerous others, again, that the narrowing toward the 
closed end is so exceedingly gradual and slight that to say they 
are conical would be no more apt than to describe the trunk Of 
one of our tallest silver fir trees as conical. Again, as to the 
structure and arrangement of the test processes, I find, even in 
the same colony, essential agreement with those said by Seeliger, 
'95, to be characteristic for P. atlanticum var. tuberculosum; and 
at the same time with those described and figured by various 
writers for P. giganteum. 

VoL - 2 1 Bitter. — The Pelagic Tunicata. 101 

Turning to the ascidizooids, I have not had much better luck 
than with the colony as a whole. Thus the branchial siphons are 
said by Seeliger ("Schlundrohr" of this writer) to reach a much 
greater length in old zooids of giganteum than they ever do in 
atlanticum. In the same colony I find old zooids with long 
siphons, but others again, certainly equally old, as judged by 
position in the colony and development of the gonads, with the 
siphons decidedly short and wide. Of course it may not be Seeli- 
ger 's meaning that the siphon becomes elongate with age in all 
zooids. Indeed, this has been one of the considerations that has 
influenced my decision to call this giganteum. The other point 
that has had weight with me concerns the arrangement of the 
zooids in the younger colonies. Seeliger states that in colonies 
of giganteum 8-9 mm. long' three whorls of zooids, regularly 
placed above one another, are present; while in colonies of 
atlanticum of the same size the zooids are more numerous and 
smaller, and are not disposed with the same regularity. My 
young colonies agree entirely with Seeliger 's account of the 
young gigantt inn. 

So far as concerns the branchial sac, the testes, and the mus- 
culature, upon which some reliance is placed by various writers 
for separating the two species, I am of the opinion that indi- 
vidual variation is so great here that the value of differences can 
be determined only by extensive quantitative studies, careful 
regard being had to the age of the zooids. 

The color variation is also great, this ranging from deep pink 
to an entire absence of the color. No intimation of blue has 
been observed. On the whole, it seems that the older colonies 
are the more deeply colored. In fact, I have not seen any highly 
colored colonies less than 8 or 10 cm. long. It is, however, true 
that one sees colonies of say 20 mm. length some of which arc 
highly colored, while others of the same size are almost if not 
wholly without pigment. 

The species occurs in abundance throughout our area, at least 
from March on through the summer and autumn months. I 
have records for December also, but a few only. It must be 
remembered, however, that we have thus far done but little 
winter collecting. 

102 University of California Publications. [Zoology 


The more important literature of the Pelagic Tunicata. 
Agassiz, A. 

1866. Description of Salpa Cabotti Desor. Proc. Boston Soc. Nat. 
Hist., Vol. XI, pp. 17-23. Good figures of Salpa democrat ica 

Apstein, Carl. 

1894. Die Thaliacea der Plankton-Expedition. B. Vertheilung der 
Ergebnisse der Plankton-Expedition der Humboldt-Stiftung. 

Bd. II, E. a. B. 
New: 8. floridiana, S. verrucosa, S. magalhanica, S. fusiform is 
var. ecMnata proles gregata. S. mania redescribed. Sptvinl 
attention to distribution, geograj>hical. vertical, seasonal. 
1901. Salpen, in Xordisches Plankton, p. LIT 5-III 10. 
Banks and Solander. 

1773. Eawkesworth 's Voyages. Vol. II. pp. 2-3. The description of 
Dagysa by Eawkesworth, but attributing the observations 
and name to Banks and Solander, admits of no question 
thai the animals described were salps. 

Blainville, de H. 

L827. Dictionnaire des sciences, aaturelles, Tunic 17, pp. ( .M-1i'.".. Ge- 
erie name Cyclosalpa proposed. 

Borgert, A. 

1894. Die Thaliacea der Plankton-Expedition C. Vertheilung der 
Ergebnisse der Plankton- Kxpodition. P>d. II, K. a. 

1901. Dolioliden, in Nordsches Plankton, p. Ill l-III 4. 
Brooks, W. K. 

1875. Embryology of Salpa. Proc. Boston Soc. Nat. Hist., Vol. 18, 
p. 193. Author's view that solitary salpa is the female, and 
chain salpa the male, hence nut a true alternation of gen- 
erations, first set forth. 

1893. The Genus Salpa. Memoirs from the Biological Laboratory of 
the Johns Hopkins University, II. Johns Hopkins University 
Press, Baltimore, pp. l-.'!i>:i. Most comprehensive study of 
the genus yet published. Part. I. — "A general account of 
the life history of Salpa. Part II. — The systematic affinity 
of Salpa iu its relation to the conditions of primitive pelagic 
life; the phylogeny of the tunicata; and the ancestry of 
the Chordata. Part III. — A critical discussion, of my own 
observations and those of other writers, on the sexual and 
asexual development of Salpa. ' ' Excellent figures of the 
adults of several species. 

Vol. 2] Bitter.— The Pelagic Tunicata. 103 

Browne, P. 

1756. The Civil and Natural History of Jamaica, p. 384, Tab. 43, 
figs. 3 and 4. First recognizable description of a Salpa. 
From the description of the "crest," the intestine, and the 
size of the zooid, there can be little doubt that the author 
was dealing with a Cyclosalpa. 

Bronn, H. G. 

1862. Mantelthiere: Tunicata Lmk. Die Klassen u. Ordnungen des 
Thierreichs. Bd. Ill, Abth. 1, pp. 103-223. Family Dolio- 
lidae in tabular arrangement, p. 216. 

Chamesso, Adelbert de. 

1819. De animalibus quibusdam e classe vermium Linnaeana in cir- 
cumnavigatione terrae auspicante Comite U. Eomanzoff duce 
Ottone de Kotzebue, Annis 1815, 1816, 1817, 1818 peracta. 
Fasciculus primus. De Salpa. Berolini. pp. 1-24. 
New species Salpa afflnis (both generations); S. runcinata (soli- 
tary generation; S. pinnata, well figured, and several other 
forms described and figured, most of which cannot be identi- 
fied with certainty. Alternation of generations in the genus 
first recognized here. "Species Salparum sub duplici con- 
spieiunter forma, prole per totum vitae cursum parenti dis- 
simili, stirpem autem huic similem generante, ita ut quaeli- 
bet Salpa matri aeque ac filiabus dispar, aviae, neptibus 
et sororibus par sit." 

Cuvier, G. 

1804. Memoire sur les Thalides et sur les Biphores. Annales du Mu- 
seum d'Histoire naturelle, T. IV, p. 360. Fullest account 
of the anatomy up to this time; relationship to ascidians 
recognized; six species described and well figured: S. Tilesii, 
S. scutigera, S. cylindrica, S. fusiformis, S. cristata and S. 
octofora, of which the first four still remain as good species. 
1817. Memoires pour servir a l'histoire et l'anatomie des Mollusques. 
Memoire sur les Thalides (Thalia. Brown), et sur les Biphores. 
(Salpa. Forskaohl), 22 pp., 1 pi. A reprint of the 1804 

Delage, Yves et Edgard Herouard. 

1898. Trataite de Zoologie Concrete. T. VIII, les Procordes. Thalia- 
cea, pp. 174-227. Many good figures of adult and develop- 
ing structure. 

Fal, H. 

1876. Ueber die Schleimdriise oder den Endostyl der Tunicaten. 
Morph. Jahrb., Bd. I, pp. 222-242. The nutritive function 
of the lateral buds of Doliolum first pointed out. 

104 University of California Publications. [Zoology 

Forskal, Petrus. 

1775. Descriptions Animalium Avium, Amphibiorum, Piseium, Insec- 
torum, Vermium; quae in itinere orientali observavit. 
Salpa, p. 112. Genus instituted and defined as belonging to 
the mollusca: 8. maxima, S. pinnata, S. democratica, S. mu 
cronata, S. confoederata, S. africana, S. punctata, S. fasciata 
S. sipho, S. solitaria, S. polycratica. 

Gegenbaur, C. 

1 s">4. Ueber die Hntwickelung von Doliolum, der Scheibenquallen und 
von Sagitta. Briefl. Niettheilungen an A. Kolliker, Zeitschr. 
f. wiss. Zool., Bd. V, p. 13-16. 
1856. Ueber den Entwickelungseyclus von Doliolum aebst Bemerkun- 
gen i'dier die Larven dieser Thiere. Zeitschr. f. wiss. Zool., 
Bd. VII. p. -JS3-314. 
hVi-ognizod the difference between the median and lateral 
buds on the dorsal process, without, however, finding how 
these are related t" one another and to tin 1 ''nurse**; and 
nt her important new tacts. 

Grobben, Carl. 

1882. Doliolum und sein Generationswechsel, etc. Wien, Issl'. One 
of the most importanl contributions in the knowledge id' 
the genus. 

Herdman, W. A. 

1882. Reporl on the Tunicata collected during the ('raise of II. M.S. 
Triton in t lie Summer el' 1882. Trans. Boy. Soc. Edinburgh, 
Vol. XXXTT, pp. '.t:;-H7. 

1888. Keport on the Tunicata collected during the Voyage of H.M.S. 
Challenger, Pari III. Pyrosomidae, Thaliacea, Larvacea. 
Challenger Reports, Zoology, Vol. XXVII, pp. 1-163, Pis. 
I II. Xew salpidae: S. musculosa, S. mollis, S. nitida, S. 
quadrata, S. echinata. Xew Doliolidae: l>. affine, I>. challen~ 
geri, ]). Jcrohni, I>. tritonis. New Pyrosomidae: /'. spinoswm. 
All new and numerous old species fully described and fig- 
ured, and relationships discussed. 

1891. A Revised Classification of the Tunicata, etc Linnean Society's 
Journal, Zoology, Vol. XXIII. Sanctions Lahille's proposal to 
divide the old genus Salpa into the additional genera, viz.: 
Thalia, Blumenbach, 1810 (not Browne, 1756; Pegea, Savigny, 
1816, and lasis, Savigny, 1816. 

1904. Ascidians and Amphioxus. The Cambridge Natural History, 
Vol. VII. Macmillan and < !o. 

Huxley, T. H. 

1851. Observations upon the Anatomy and Physiology of Salpa and 
Pyrosoma. Philos. Trans. Roy. Soc. London, part IL, pp. 
567-593. In addition to the important discussion of the 

VoL - 2 J Bitter.— The Pelagic Tunicata. 105 

structure and affinities, an excellent review of the litera- 
ture. Author's interpretation of reproduction in Salpa: 
' ' The parent and offspring are not dissimilar, but the indi- 
vidual is composed of two zooids. ' ' 

1851. Eemarks upon Appendicularia and Doliolum, two genera of 

the Tunicata. Ibid., pp. 595-605. 

Jones, T. Rupert. 

1818. Tunicata. Cyclopedia of Anatomy and Physiology. Vol. IV, 
Part II, pp. 1185-1243. 

Keferstein, W., und Ernst Ehlers. 

1861. Zoologische Betrage gesammelt in Winter 1859-60, in Neapel 
und Messina. Ill Ueber die Anatomie und Entwicklung 
von Doliolum, pp. 56-71, Pis. IX, X, XL Important. IV 
Bemerkungen iiber die Anatomie von Pyrosoma, pp. 72-77, 

Krohn, A. 

1846. Observations sur le generation et le developpement des Bi- 
phores (Salpa). Ann. d. sci. Nat., ser. 3 (Zool.), t. VI, pp. 
110-131. First important confirmation and extension of 
Chamisso's discovery of "Alternation of generations"; 
sexual elements in proles gregata, and budding in proles soli- 
taria observed; idenification of the two generations belong- 
ing to the same species first made for several species. 

1852. Ueber die Gattung Doliolum und ihre Arten. Archiv. fiir Natur- 

geschichte, pp. 53-65, PI. II. The tailed larva observed; 
alternation of generation in this genus first recognized, and 
this compared with the similar reproductive process in 
Salpa. The simple alternation of a sexual with an asexual 
generation, as in Salpa, assumed. D. mulleri, D. nordmanni 
and D. troschelii described as new. 

Lahille, F. 

1890. Contributions a 1 'etude anatomique et taxonomique des Tuni- 
ciers. Theses, Tolouse. 

Lesson, R. P. 

1830. Voyage autour du monde, execute per ordre du roi, sur la cor- 
vette de S.M. la Coquille, 1822-25. Zoologie, t. II, Paris, 
1830, pp. 256-279. 


1813. Memoire sur quelques nouvelles especes d'animaux mollusques 

et radiaires receuillis dans la Mediterranee pres de Nice. 

Bull. d. Scienc de la Societe Philom., Paris. T. 3, p. 281, 

PI. V, fig. 2. 
1815. Memoire sur 1 'organisation des Pyrosomes, et sur la place qu 'ils 

semblent devoir occuper dans une classification naturelle. 

Bull. d. Scienc. de la Societe Philom., Paris, 1815, p. 70, PI. 

I, figs. 1-15. Also, Journ. de Phys. de Chim., et d'hist. nat., 
T. LXXX, p. 413. 

106 University of California Publications. [Zoology 

Metcalf, M. M. 

1893. On the Eyes and Subneural Gland of Salpa. Supplement to The 
Genus Salpa, by Brooks. In addition to the most detailed 
study of the organs treated yet published, bearings of the 
results on question of relationships of the species considered. 

1900. Notes on the Morphology of the Tunicata. Zool. Jahrb., Bd. 
XIII, pp. 495-602. Numerous anatomical observations, par- 
ticularly on the brain, subneural gland, and associated parts 
in many species of the class, including several Salpae; struc- 
ture and affinities of OctacTiemits patagomensis. 

Panceri, Paolo. 

1872. Gli organj Luminosi e la luce dei Pirosomi e <lelle Foladi. Atti 
della R. Acad, della Scienze fisiche e mathematiche de 
Napoli, Vol. V. No. 13, p.51, Pis. 3. In addition to first 
recognition of the function of the light producing organs, 
various observations on anatomy and embryology. Most 
of the observations on P. gigantt um. 

Peron, M. 

1804. Memoire sur le nouveau genre Pyrost i. Annals du Museum 

d'hist. aatar, T. I V, p. 4: , .7. PI. 72. Pyrosoma atlanticum 
Perrier, Edmond. 

1899. Traite de Zoologie. Tasc. V, A.mphioxus-Tuniciers. 
Quoy et Gaimard. 

1834. Voyage de decouvertes de I'Astrolabe. Zoologie, t. III. pp. 559- 
602, Pis. 86-89. Many species of Salpa and Doliolum elab- 
orately, bul in many instances nol accurately, illustrated; 
many of the figures not identifiable. 

Salensky, W. 

Beitriige zur Embryonalentwicklung der Pyrosomen. Zool. 
Jahrb. (Abth. f. Anat. u. Entw.), Bd. V, Hft. 1, pp. 1-98. 
This, along with Salensky's numerous other papers on the 
development not only of Pyrosoma, but also of Salpa and 
compound ascidians, indispensable to students of the devel- 
opment of these groups. 

Savigny, Jules-Cesar. 

1816. Memoires sur les Animaux sans Vertebres. 2d partie, 1st fasc, 
p. 124, PI. XXIV. S. octofora and S. cylindrica examined 
and figured. Affinity of Salpa to ascidians confirmed. 

Seeliger, O. 

1889. Ueber den Generationswechsel der Salpen. Jena Zeitschr. f. 
Naturwiss, Bd. 22, Hft. 3, pp. 399-414. In this and various 
other important papers by this author, affinities of all the 
kinds of tunicates discussed, particularly from the develop- 
mental point of view. 

Vol. 2] Ritter. — The Pelagic Tunicata. 107 

1895. Die Pyrosomen der Plankton-Expedition. Ergebnisse der 
Plank.-Exped. der Humboldt-Stiftung, Bd. II, E. b. Fullest 
most resent treatise on the structure and classification and 
distribution of the group. New species: P. ahemiosum, P. 
minimum, and varieties levatvm and tuberculosum of P. at- 

Todaro, F. 

1902. Sopra gli organi escretori delle Salpidi. Kend. Acead. Lincei 
(5), Vol. 11, Seiu. 1, p. 405-417. In addition to description 
of renal organ in Salpa, proposes genus Helicosalpa. 

Traustedt, M. P. A. 

1S85. Spolia Atlantica. Bidrag til Kundskab om Salperne. Vidensk. 
Selsk. Skr., 6 Eaekke, naturvidenskabelig og niathematisk. 
Afd. II, 8 p. 339-400. Most useful paper on the classifica- 
tion of Salpa yet published. 

1893. Die Thaliacea der Plankton-Expedition A. Systematise!^ Bear- 
beitung. Ergebuisse der Plankton Expedition, Bd. II, E. a. A. 

Uljanin, Basilius. 

1884. Die Arten der Gattung Doliolum im Golfe von Neapel, etc. 
Fauna und Flora des Golfes von Neapel, Monographic N. 
The most extensive treatment of the genus yet published. 

Vogt, C. 

1854. Eecherches sur les Animaux inferieurs de la Mediterranee. 
Second memoire, Sur les Tuniciers nageans de la Mer de 
Nice. Mem. de l'lnstitut national genevois, II. Complete 
list of species of Salpa described up to date of the memoir, 
with association of the two generations of each species; 
budding in Salpa, pp. 5-61. Genus Anchinia, pp. 62-73. 
Genus Appendicularia, pp. 74-86. Genus Pyrosoma, pp. 
87-96. Six plates, illustrating all the genera. 

These abbreviations are used 
of 1 be plates. 

an. anus. 

til. o. al rial orifice 

br. — branchae. 

br. o. branchial orifice. 

coe. — Cecum of intestine of sexual 

I /. eleoblast. 
i ml. endostyle. 
emb. — embryos. 
es. — esophagus. 

d. p. — dorsal process of nurse. 
gn. — ganglion. 
lit. — heart. 
hy. — hypophysis. 

in the text figures as well as in those 
i. c. - 1 ntesiin.-il appendages. 

int.— intestine. 

i a.- " nucleus. 
oc— otocyst. 

OV. - ovary. 

/'. a. |misi abdomen. 

y. hr. h. peripharyngeal band. 

pd. — peduncle t'nr attachmenl of 

sexual zooid t<> stolon. 
/'<(/. peduncle. 
/'in. placenta. 
8. — stomach. 
sto. — stolon. 

1 1 x. — testis. 

V. /'. — ventral process of phorozooid. 



All figures "I' Plates II ami III are <>t' Cyclosalpa bdkeri. 

Pigs I. 2, '■''. Lateral, dorsal, ami ventral \irus, respectively, of the 
solitary generation, all drawn mainly from preserved specimens. 

Pig. l. Outline of same, lateral view, made from a Living, actively 
swimming specimen. 

Pig. 5.- Intestinal tract, "gill," endostyle, peripharyngeal band, hypo- 
physis, ami ganglion with its sense organ, seen from lefl side. 

Pig. ii. Anterior end of "gill," with hypophysis. 


:1 PlATE H. 



Figs. 7 and 8. Lefl and right side views, respectively, of zooids of 
sexual generation, takes from the stolon. 


[Ritter] Plate III. 

FIG. 8 

3 gi- 



Vol. 2, No. 4, pp. 113-233, Figs. 1-62. October 14, 1905 










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The present paper is the result of a study of the pelagic Cope- 
poda collected in the San Diego region during one month of the 
summer of 1903, nearly two months of 1904, and during Decem- 
ber and January, 1903. Likewise, there have also been examined 
a few collections taken during the fall of 1904. Such time as 
has been spent on the subject serves to convince the writer that 
a good deal of further study is necessary, and that, taking the 
field as a whole, this report is incomplete. Such forms as are 
described here seem well established. Without doubt there are 
others which are new, but for lack of proper material they are 
not now dealt with. 

The work was commenced at the San Diego Marine Labora- 
tory connected with the University of California, and completed 
at Harvard University. It is a pleasure to express my apprecia- 
tion of the helpful suggestions and advice of Professor W. E. 
Kitter of the University of California, and Professor E. L. Mark 
of Harvard University. 

In the body of a free-swimming Copepod two regions are 
readily distinguishable, a cephalothorax and an abdomen. In 
the first suborder, Gymnoplea, the cephalothorax consists of the 
head and five thoracic segments, though the head is often fused 
with the first thoracic segment, and the fourth with the fifth. 
In the second suborder, Podoplea, the last thoracic segment is 
drawn into the abdominal portion; this, then, loosely speaking, 
contains one thoracic segment. 

114 University of California Publications. [Zoology 

In typical groups the abdomen proper consists of five seg- 
ments in the male, and always less than five in the female: fu- 
sions often reduce the number of segments to as few as one or 
two. The first segment of the abdomen in each sex bears the 
orifices of the sexual organs, and is called the genital segment; 
the last segment contains the opening of the alimentary canal, 
and is called the anal segment. The furca is a paired appendage, 
borne on the anal segment and carrying usually a fringe of six 
bristles of various lengths. The anterior portion of the head 
segment is known as the front, and terminates ventrally in the 
rostrum. The latter may consist of one or two strong, pointed 
projections, or of the same number of long, slender filaments. 

The appendages of the cephalic segments in order are. from 
front to rear: (1) The anterior antennae; (2) The posterior 
antennae; (3) The mandibles; (4) The maxillae; (5) The ante- 
rior maxillipeds (second maxillae); (6) The posterior maxilli- 
peds ( maxillipeds). The thoracic segments hear the swimming 
feel i four pairs), and also a fifth pair of feel which usually are 
modified in both sexes, and at times are absenl in the female. 
The cephalic appendages from :* to 6, inclusive, are the mouth 
parts. All the appendages except the anterior antennae are 
typically biramous: each consists of a two-jointed basal part 
(basipodite) , which hears the inner and outer rami (respectively 
endopoditi and exopoditt (. Neither ramus is more than three- 
jointed, and in many forms the number of joints is reduced by 

The anterior antennae in the Gymnoplea consist of 25 joints,, 
but this number may be reduced by fusion. The joints 
carry bristles and sense organs, or a< st Ik tasks. The joints in 
an appendage, or a part of one, are numbered from the base 
to the tip. Among the (iymnoplea one of the anterior antennae 
of the male may be modified to form a grasping organ; this is 
usually on the right side, but both appendages may be so modi- 
fied. The grasping organ may be recognized by its want of 
symmetry. The joints on either side of the geniculation are 
often provided with teeth. In many of the Gymnoplea the fifth 
feet in the male form grasping organs ; the fifth feet are always 
asymmetrical in the male, whether they form grasping organs. 

Vol. 2] Estcrly. — Copepoda of the San Diego Region. 115 

or not. In the Podoplea some of the males have anterior grasp- 
ing antennae ; in others this function is performed by the poste- 
rior antennae or posterior maxillipeds. 

The main axis of the appendages (excepting the anterior 
antennae) may be considered as parallel to the dorso-ventral 
diameter of the animal ; accordingly anterior and posterior faces 
are distinguished in an appendage as well as proximal and distal 
portions, and inner or outer margins of the various joints. 
This nomenclature is useful, especially since most appendages 
are flattened. The form, arrangement and number of the bristles 
on the appendages are used a great deal in identification, and 
for that reason the appendages must usually be dissected off. 

It is of importance to have only adult animals for study. 
The adult male in most genera has noticeable sexual peculiarities 
in the structure of the fifth feet or in the grasping antenna. 
The females are certainly adult if carrying eggs or spermato- 
phores. In large animals it is a comparatively easy task to dis- 
sect off the appendages, but in the majority of the Podoplea 
and the smaller Gymnopiea it requires a good deal of patience 
and a steady hand. A dissecting microscope is indispensable, 
and it is a good plan to use fine needles, which may be ground 
down to an edge. Farrant's fluid makes a good mounting me- 
dium for permanent preparations. If it is spread thin over the 
slide the appendages may be placed in order in it and the cover 
glass put on without disturbing the arrangement. For the deter- 
mination of the genus of an individual the feet especially must 
be removed and examined. 

Most of the drawings in this paper were made with the aid 
of the Abbe camera. The keys, and descriptions of all but new 
species, are translated from Giesbrecht's works of 1892 and 1898. 
In the general key for the Gymnopiea the plan adopted in the 
Tierreich (1898) has been followed, but only those genera are 
included which from their distribution might be expected to 
occur in the San Diego region. Species keys are given in some 
cases, but include only the species actually found; however, if 
one sex of a form occurs, and the other has not been obtained, a 
description of the latter is given in most cases. 

116 University of California Publications. [Zoology 

It may be remarked that little defense is needed for translat- 
ing bodily from Giesbreeht, in view of the fact that whatever 
could be written concerning any form known to him has keen 
so well stated that one could not improve upon it, 

Following is a list of the species treated in this paper: 


1. Acartia tonsa Dana '204 

2. A( tideus armatus Brady 145 

3. Arictclliis .si Ins us (Weslinvht 189 

4. Augaptilus longicaudatus Giesbreeht L88 

5. Cain mis fiiiiiiiiirliu-iis dinner 125 

6. Calanus gracilis Dana 128 

7. Calanus minor Giesbreeht 126 

8. Calanus robustior Giesbreeht 129 

9. Calanus tenuicornis Dana 1'_'7 

10. Candacia bipinnata Giesbreeht 195 

11. Ciiniliiciii curia Dana 196 

12. Candacia a< thiopica Dana l'-'ii 

L3. Candacia /" ctmata Brady 193 

14. Ci ill injiinh s bradyi Wliorlor 17"_' 

If). Clausocalanus arcuicornis Giesbrechl 142 

16. Eucalanus attenuatus Dana 133 

17. Eucalanus crassus < riesbreehl 134 

is. Hunila mis iloiu/dtus Dana 131 

19. Eucalanus subti wwis Giesbreeht 135 

20. Euchaeta acuta Giesbreeht 157 

21. Euchaeta media Giesbreeht 160 

22. Euchaeta spinosa Giesbrechl 159 

23. Kuril i n lid amoena Giesbreeht 155 

•J4. Euchirella galeata Giesbreeht 155 

25. Euchirella m< ssint nsis Giesbreeht 151 

26. Eurliiit Ha pulchra Lubbock 153 

27. Euchirella rostrata clans 152 

28. Gaidius pungi ns Giesbreeht 146 

29. Heterorhabdus clausi Giesbrechl 185 

30. //- U rorhabdus longicornis Giesbreeht 186 

31. lh U rorhabdus papilliger < llaus 184 

32. //' U rorhabdus spinifrons Claus 182 

33. Labidocera trispinosa, new species 200 

34. Lucicutia flavicornis clans 180 

35. Mecynoa ra clausi Thompson 137 

36. Mrl ml in luci ns Boecfc 177 

37. Metridia boecki Giesbreeht 178 

38. Faracalanus parvus Glaus 140 

39. Phyllopus bidentatus Brady 191 

40. Pleuromamma abdominalis Lubbock 174 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 117 


41. Pleuromamma gracilis Claus 175 

42. Pleuromamma xiphias Giesbrecht 176 

43. Rhincalanus nasutus Giesbrecht 136 

44. Scolecithrix bradyi Giesbrecht 165 

45. Scolecithrix danae Lubbock 164 

46. Scolecithrix pacifica, new species 168 

47. Scolecithrix persecans Giesbrecht 166 

48. Scolecithrix similis Scott 170 

49. Scolecithrix subdentata, new species 167 

50. Undeuchaeta major Giesbrecht 148 

51. Undeuchaeta minor Giesbrecht 149 


52. Clytemnestra rostrata Giesbrecht 214 

53. Corycaeus carinatus Giesbrecht 226 

54. Corycaeus venustus Dana 225 

55. Euterpe acutifrons Giesbrecht 212 

56. Microsetella rosea Giesbrecht 211 

57. Oithona nana Giesbrecht 209 

58. Oithona plumifera Baird 207 

59. Oncaea conifera Giesbrecht 216 

60. Oncaea minuta Giesbrecht 217 

61. Sapphirina angusta Dana 221 

62. Sapphirina iris Dana 219 

63. Sapphirina lomae, new species 223 

64. Sapphirina scarlata Giesbrecht 222 

I Sub-order— GYMNOPLEA. 

Gymnoplea (sub-order) Giesbrecht, 1892, p. 41. 
Gymnoplea (tribe) Giesbrecht, 1898, p. 7. 

The genital orifices lie in the first segment of the posterior 
division of the body ; they are ventral and paired in the female, 
unpaired and lateral in the male. The fifth pair of feet in the 
female are like the preceding pairs, retrograded or lacking; in 
the male a pairing organ always present. The first segment of 
the posterior division of the body (abdomen) never bears 
appendages. The abdomen of the male is 5 segmented (fig. 3a), 
and the female seldom carries the eggs in sacks hanging from 
the genital orifice. The anterior antennae in the male may be 
symmetrical, or one may form a grasping organ ; in the female 
the antennae are symmetrical. 

118 University of California Publications. [Zoology 


(The genera marked with an asterisk have been found in the 
San Diego region.) 

1. Inner ramus of third and fourth feet 3-jointed, figs. 12a, 19b 2 

1. Inner ramus of third and fourth feet 2- jointed, fig. 44>l 19 

2. Inner ramus of first foot 3-jointed 3 

2. Inner ramus of first foot 2-jointed 32 

2. Inner ramus of first foot 1-jointed, fig. lod 38 

3. A black or brown knob on the first segment of the cephalothorax 

in the antero-lateral angle, on the right or loft side; figs. 33a, 

34a *Pleuromamma 

.".. This knob absent 4 

4. First joint of inner ramus of second fool with proximally curved 

hooks on the inner margin; fig. 35b *Metridia 

4. This joint, like the rest, bearing a bristle 5 

5. Tormina] joinl of outer ramus of third and fourth feet with two 

spines or thorns on outer margin, and one terminal bristle; fig. Id (i 

5. Terminal joinl as above, but with three apines on outer border; 

figs. 17/,. 196 6a 

6. Terminal bristle of outer ramus of third and fourth feet with 

broad, smool h border; ti«, r . \d *Calanus 

6«. Terminal bristle bearing teeth or spines <>n outer border; figs. 

35a, 39a 7 

7. One bristle of left ramus of furea much Longer and thicker than 

the ot her furea] bristles 8 

7. Purcal bristles symmetrical !i 

8. Mandibular blade with three or four teeth, the ventral one 

hooked and separated from the others by a wide spare; fig. 
38/" *Heterorhabdus 

8. Mandibular blade with at least 8 teeth Disseta 

it. Anterior antennae symmetrical 10 

9. Anterior antennae asymmetrical 14 

10. Kami of fifth feet 3-jointed; fig 32c 11 

10. Rami of fifth feet 2-jointed *Augaptilus$ 

10. Outer ramus 3-jointed, inner ramus 2-jointed IsochaetaJ 

Id. Outer ramus 3-jointed, inner ramus L-jointed IsiasO 

10. Outer ramus 3-jointed, inner ramus lacking *Phyllopus9 

10. Outer ramus 1 jointed, inner ramus rudimentary; fig. 42b *Arietellus9 

11. Middle joint of outer ramus of fifth foot with a thorn-like pro- 

cess which is fused with the joint ; fig 32c *Centropages$ 

11. This joint with an awl-shaped or rudimentary bristle on the 

inner border; figs. 36b, 38e 12 

12. Terminal joint of inner ramus of fifth foot with five bristles; 

fig. 366 *Lucicutia$ 

12. This joint with at least six bristles 13 

13. Abdomen with 4 segments HaloptilusP 

13. Abdomen with 3 segments *Augaptilus9 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 119 

14. Grasping antenna on right side 15 

14. Grasping antenna on left side 17 

15. Inner rami of both feet of fifth pair 3-jointed, with plumose 

bristles ; fig. 40a 16 

15. Inner rami rudimentary, without plumose bristles IsiasJ 

16. Inner rami of feet of fifth pair unlike, the right foot with a 

forceps *CentropagesJ 

16. Inner rami alike; figs 41c, 41d *Augaptilus$ 

17. Both rami of each foot of fifth pair 3-jointed 18 

17. Both outer rami 3-jointed, the inner rudimentary *ArietellusJ 

17. Eami of left fifth foot 3-jointed, of the right 2-jointed. . *Lucicutiat 

18. Inner ramus of maxilla present, distal bristles of anterior max- 

illiped naked or set with spines HaloptilusJ 

18. Inner ramus of maxilla lacking; distal bristles of anterior max- 

illiped with mushfoom-shaped appendages (cf. fig. 41&) 

19. Inner ramus of first foot 3-jointed 20 

19. Inner ramus of first foot 2-jointed 25 

19. Inner ramus of first foot 1-jointed Eurytemora 

20. Head without dorsal cuticular lenses or lateral hooks 21 

20. Head with one or two pairs of cuticular lenses and hooks on each 

side 23 

21. Rami of posterior antennae about equal in length Parapontella 

21. Inner ramus much the shorter 22 

22. Abdomen with asymmetrical outgrowths Pontellopsis 

22. Abdomen symmetrical Pontellina 

23. Head with two pairs of eye lenses Anomalocera 

23. Head with but one pair of lenses 24 

24. Last two thoracic segments distinct; anterior antennae of female 

24-jointed; terminal portion of grasping antennae of male 
4-jointed Pontella 

24. Last two thoracic segments fused; anterior antennae of female 

22-jointed; terminal portion of grasping antennae of male 
4-jointed Ivellopsis 

25. Head with one pair of cuticular lenses; fig. 48a *Labidocera 

25. Head without dorsal cuticular lenses 26 

26. Inner rami of fifth feet jointed *Centropages 

26. Inner rami of fifth feet 1-jointed or absent 27 

27. Second maxilliped longer than the first 28 

27. Second maxilliped shorter than the first 29 

28. Furca long and narrow, at least six times as long as broad. . . .Temora 

29. First joint of maxillipeds with few short bristles; fig. 47/i. . *Candacia 

29. This joint bearing long bristles set with spines 30 

30. Outer ramus of posterior antennae less than half as long as the 

inner ramus Acartia 

30. Outer ramus more than half as long as the inner 31 

31. Posterior maxilliped 6- 7- jointed Calanopia 

31. Posterior maxilliped 3-jointed Tortanus 

32. Both feet of fifth pair with inner^rami, which are 2- or 3- jointed, 

bristles plumose " *Lucicutia 

120 University of California Publications. | Zoology 

32. Inner rami without plumose bristles, or lacking; the entire foot 

may be absent on one or both sides 33 

33. Furca long and narrow, at least six times as long as broad. . . .Temora 

33. Furca at most three times as long as broad 34 

34. Middle joint of outer ramus of third and fourth feet with two 

i ristlcs; terminal joint with seven; fig. 12a 35 

34. M Lddle joint with one, terminal with five 37 

35. Oiiier border of outer ramus of Bwimming feet not denticulate.. 


35. Outer border on rear pairs denticulate; fig. 12a 36 

36. Fifth foot lacking in female, or knob-like; in the male only the 

I't'i fifth foot present Acrocalanu;> 

36. Fifth fool in female 2 jointed (fig. L2c); in the male the right is 

2-jointed, the left .".jointed (fig. L2e) *Paracalanus 

:'.7. uiiier ramus of firsl toot 3-jointed; fifth fool absenl in female... 


37. Outer ramus of first foot 2-jointed; fifth foot present in female 

(fig. 106) Rhincalanus 

38. Inner minis of second foot 3-jointed *Mecynocera 

.'is. Inner ramus of second fool 2-jointcd (fig. 286) 39 

38. Inner ramus of second foot 1- jointed (fig. 25§) 51 

39. Terminal joint of outer rami of second to fourth feet with five 

lnist les on inner margin Spinocalanus 

.".9. Terminal joinl with four bristles on inner margin (fig. 14b) 40 

40. Surfaces of rami of second to fourth feet \vilhont very large 

spines; appendages of anterior maxillipeds have the form of 

bristles or hooks -11 

10. Surfaces of outer rami and of the two terminal joints of the 

inner rami of the third and fourth feet with Larger spines (as 

in fig. 286); part of the appendages of the anterior maxilliped 
vermiform I fig. 30d) or pencillate 49 

11. Basals and outer rami in second and third feet broader than in 

the fourth pair, the second basal irregularly toothed on the 
distal border (fig. 136) *Clausocalanus 

41. Second and third feet not differing as above from fourth foot... 42 

42. Outer marginal thorns of terminal joint of outer ramus of third 

and fourth feet comb-like and placed in deep indentations in 
the margin Ctenocalanus 

42. These of the usual shape 43 

43. Fit't h pair of feel svmniet rical or lacking 44 

43. fifth pair of feet asymmetrical or lacking 47 

44. Anterior part of head with a spine in the dorsal median line.... 


44. Head without this spine 45 

45. Fifth foot lacking Pseudocalanus? 

45. Fifth foot 2-jointed, with a thick, curved bristle at the end 46 

46. Terminal bristle of fifth foot much longer than the basal joint 


46. Terminal bristles not as long as, or but little longer, than basal 

joint Stephus$ 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 121 

47. Fifth feet, especially the left, with several apical appendages... 


47. Fifth feet slender, stylet-like, with a few short needles or with 

but one needle or hook-like appendage 48 

48. Fifth feet slender, stylet-shaped, about as long as the abdomen. . . 


48. These shorter than the abdomen, the right foot with a terminal 

hook Drepanopus*, 

49. Cephalothorax broad to globular Phaenna 

49. Cephalothorax elongate, ellipsoidal (figs. 26a, 29) 50 

50. Head distinct from thorax Xanthocalanus 

50. Head fused with thorax (figs. 26a, 27) *Scolecithrix 

51. Fifth foot lacking 52 

51. Fifth foot present 59 

52. Last thoracic segment prolonged laterally into a long, pointed 

process (figs. 14a, 15a) 53 

52. Lateral angles of last thoracic segment rounded or slightly pointed 

(figs. 16a, 19a) 56 

53. Bristles of the sixth joint from the last in the anterior antennae 

thick and transversely ringed BradyidiusJ 

53. These bristles of the usual form 54 

54. Eostrum with two heavy teeth (fig. 14«) *Aetideus$ 

54. Eostrum with one point (fig. 15o) or lacking 55 

55. Outer ramus of first foot 3-jointed ChiridiusJ 

55. Outer ramus of first foot 2-jointed (fig. 15d) "Gaidius^ 

56. Eami of posterior antennae about equal in length *Euchaeta2 

56. Outer ramus at least iy 2 times as long as the inner ramus 

(fig. 20c) 57 

57. Inner border of first basal of fourth foot naked or feathered 58 

57. Inner border with teeth or spines (figs. 19&, 20d) *Euchirella$ 

58. Head with or without crest (figs. 16a, 17a) ; last thoracic segment 

not produced into spines or blunt processes *Undeuchaeta$ 

58. Head with median crest, last thoracic segment produced into 

blunt processes (not spines) Chirundina? 

59. Last thoracic segment on each side with a strong point (fig. 

14a) *AetidiusJ 

59. Last thoracic segment rounded (fig. 20a) 60 

60. Eight fifth foot with forceps (figs. 20&, 216) EuchirellaJ 

60. Eight fifth foot ends in a stylet without forceps (fig. 23a) .... 61 

61. Inner rami of posterior antennae at most equal in length to the 

outer EuchaetaJ 

61. Inner rami of posterior antennae over half the length of 

the outer *Undeuchaeta£ 

122 University of California Publications. [Zoology 


Dana (subfamily), 1852. 

Claus, 1863, p. 166. 

Giesbrecht, 1892, p. 41; 1898, p. 12. 

First antennae of male symmetrical or nearly so, not genicu- 
late; aesthetasks more numerous than in female. Fifth feet of 
female either like the preceding ones or in various stages of 
degeneration, often resulting in complete loss. The males vary 
from the females in the segmentation and form of abdomen, in 
structure, number of joints, and appendages of the anterior 
antennae, and in the form of the fifth foot. Head usually dis- 
tinct from thorax; the two last thoracic segments usually fused. 
Rostrum with one or two joints, sometimes lacking. Abdomen 
of female usually with 4 (fig. la), seldom with 3 or 2 segments; 
thai ill' the male with 5, often with very short anal segment. 
Anterior antennae of female, l<i- to 25-jointed. Outer ramus of 
second antennae one-half in four limes as Long as the inner, 5- to 
8-jointed, middle joints short, end ones usually elongate. Six 
to eight teeth on blade of mandible, outer ramus ;">-. inner ramus 
2-jointed. Maxilla with at least two lobes on inner margin and 
one on outer: outer ramus always present, inner 1- to 3-jointed, 
seldom fused with second basal. Proximal and distal curved 
bristles of first maxilliped usually equal in length, the former 
sometimes modified into delicate saclike appendages (fig. 306). 
Second maxilliped elongate, terminal portion (inner ramus) 5- 
jointed, from one-third to one and one-half times the Length of the 
second basal. Outer rami of first to fourth feet 3-jointed (that 
of first foot occasionally 2-jointed) ; inner ramus of first and 
second pairs 1- to 3-jointed, of third and fourth 3-jointed; term- 
inal bristle of outer rami at times with smooth border (fig. Ic7), 
not serrate. Inner rami of fifth pair of male rarely 3-jointed, 
mostly rudimentary or lacking ; the outer ramus forms hooks 
or shears ; occasionally the entire appendage of one side may be 

Sub-fam. Calaninae. 
Calanina Giesbrecht, 1S92, p. 44. 

Fifth pair of feet in the female in all respects like the pre- 
ceding pairs ; in both sexes all five pairs are provided with 3- 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 12;'. 

jointed inner and outer rami, and the number of outer marginal 
bristles on the outer ramus is the same in all the feet. The 
number of bristles on the inner ramus of the first pair is: one 
on the first joint, two on the second, six on the third (one on 
outer margin) ; on the terminal joint of the second and third 
pairs there are eight (two outer marginals). The fifth pair of 
feet in the male is always modified to form an accessory sexual 
organ : the right foot has a 2-jointed basal and is biramous, each 
ramus with three joints ; the left foot has also a 2-jointed basal, 
and a 3-jointed outer ramus, while the inner ramus is reduced 
and may be entirely absent. 

1. Genus Calanus Leach . 

Monoculus Gunner, 1765. 

Calanus Leach, 1819, p. 539. 

Undina Dana, 1852, p. 1047. 

Cetochilus Claus, 1863, p. 169. 

Calanus Brady, 1883, p. 30. 

Calanoides Brady, 1883, p. 74. 

Undina Brady, 1883, p. 52. 

Calanus Uiesbrecht, 1892, pp. 45, 88, 725. 

Calanus Dahl, 1894b, p. 61. 

Calanus Dahl, 1898, p. 13. 

Calanus Wheeler, 1899, p. 164. 

$ Head free, or fused with thorax ; fourth and fifth thoracic 
segments not fused. Abdomen with 4 segments, genital segment 
symmetrical, furca occasionally asymmetrical. Anterior antennae 
25-jointed, terminal joints with long, plumose bristles ; the anten- 
nae vary in length, in some cases not reaching the posterior end 
of the body, in others extending beyond the furca (fig. 5a). 
Rami of posterior antennae of about equal lengths, outer ramus 
7-jointed. Inner ramus of maxilla 3-jointed. Anterior maxil- 
liped with long, curved bristles on inner border, outer border 
with a plumose bristle. Inner ramus of posterior maxilliped 
long, 5-jointed, bristles long and stiff, usually not plumose. 
Outer and inner rami of first to fourth pairs of feet 3-jointed, 
first, second and third joints of outer rami of all with 1, 1, 2 
marginal spines, respectively, terminal bristle scalpelliform, its 
margin smooth (fig. Icl). Inner ramus of first foot with 1, 2, 6 
bristles on the first, second and third joints in order; terminal 

124 University of California Publications. [Zoology 

joint of inner ramus of second and third feet with eight bristles. 
Fifth foot like the others. 

J Abdomen with 5 segments, genital orifice on left side of 
genital segment. Number of joints in the anterior antennae 
reduced, at least by fusion of the first and second joints. Mouth 
pails often retrograded; swimming feet usually as in the female. 
Fifth feet in some cases similar to the swimming feet, in others 
asymmetrical pairing organs (fig. 1/'. c), 2 basals on each side. 
right foot (fig. 16) with 3-jointed outer and inner rami, the left 
(fig. Iri with 3-jointed outer ramus without plumose bristles, 
inner ramus reduced or lacking. 


$1. Anterior antennae extend beyond end of cephalothorax for :it 

Leasl half its Length ( (i «_r. 5a) 2 

1. Anterior antennae 'I" nut extend beyond end of cephalothorax, 

nr only for a few joints at most -J 

l'. r.iistlcs of furca symmetrical C. tenuicomis 

1'. Furca with an elongated bristle on left side .'! 

3. Ventral surface of genital segmenl very strongly convex (fig. 

C. robustior 

3. Ventral surface of genital Begmenl rather Blightly convex (fig. 

C. gracilis 

4. Head net fused with thorax: cephalothorax with six segments 

[fig ho C. nnmarchicus 

4. Head fused with thorax: cephalothorax with five segments (fig. 

2) C. minor 

Jl. Anterior antennae longer than body by at least six joints 

C. tenuicornis 

1. Anterior antennae nol Longer than body, or bul slightly so 2 

2. Outer margin of terminal joint of outer ramus of second to 

fourth feet denticulate 3 

2. ( hiter margin of same smooth or feathered 4 

3. Inner rami of fifth feet similar in structure C. gracilis 

:'.. Inner ramus of left foot of fifth pair shortened ami without bristles 

C. robustior 

4. Outer ramus of right fifth foot with plumose bristles on inner 

margin C. minor 

4. Outer ramus of right fifth foot without bristles on inner margin 
(fig. lb); outer ramus of left foot (fig. lc) less than twice as 
long as that of the right C. finmarchicus 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 125 

1. Calanus finmarchicus Gunner. 

Nonocuhis finmarchicus Gunner, 1765, p. 175, figs. 20-30. 
Calanus perspicax Dana, 1852, p. 1071; 1855, pi. 74, figs. la-c. 
Cetochilus helgolavdicus Claus, 1863, p. 171, pi. 26, figs. 2-9. 
Calanus finmarchicus Brady, 1883, p. 32, pi. 1, figs. 1-10. 
Calanus finmarchicus Giesbrecht, 1892, pp. 89, 218, pi. 6, fig. 19; 
pi. 7, figs. 32, 33; pi. 8, figs. 3, 15, 21, 31, 33; 1898, p. 14. 
Calanus finmarchicus Wheeler, 1899, p. 164, fig. 1. 

Fig. 1. — Calanus finmarchicus. (a) Female, lateral, X18. (b) Outer 
ramus of right fifth foot of male X195. (c) Left fifth foot 
of male X140. (d) Outer ramus of third foot of male 
X140. St., terminal bristle. Se., outer marginal bristle. 
(e) Basals of fifth foot of female X195. 

$ Head not fused with thorax, front and lateral portions of 
fifth thoracic segment rounded, furcal bristles symmetrical. An- 
terior antennae extending' about to end of abdomen, aesthetasks 
not doubled on any joint. Distal margin of second basal of 
second to fourth feet with a tooth ; the proximal spine of the 
outer margin of terminal joint of outer ramus divides the margin 
in these pairs (respectively) into portions as 2:1, 2:1, 3:1. 

126 University of California Publications. [Zoology 

First basal of fifth pair with concave dentate inner border 


J Head not fused with thorax, anterior antennae straight ; 
outer ramus of right fifth foot without bristles on inner margin 
(fi-. 1&), terminal bristles sometimes dentate, thorn-like. Basals 
and proximal joint of outer ramus of left foot ( 6g. lc) elongate, 
terminal joint of outer ramus shortened. Outer ramus of right 
foot (excluding terminal bristle) readies at most to the distal end 
of the second joint of the outer ramus of the left foot. Inner 
rami of both feet similar in structure. 

Coloration: Rather transparent, with variably distributed 
red pigment. In some eases this is found only in one of the 
anterior antennae, in others in the thorax and appendages, whih 
the entire body of some animals is brightly colored. 

Length: Both sexes. 2.6-3.1 mm. 

Occurrence: Probably the commonesl species in the San 
Diego region, occurring abundantly in nearly all collections with 
the larger nets; in some ciscs ('. finmarchicus is almost the only 
species, and is very often predominant. 

2. Calanus minor Glaus. 

Cetochilus minor Claus, 1863, p. 172, pi. 26, figs. 1-8. 

Calanus valyv.t Brady, 1SS3, ]>. 33, pi. 3, figs. 1-7. 

Calanus minor Giesbre^ht, 1892, p. 90, pi. 6, figs. 3, 1(>, 22; pi. 7, 

figs. 6-22; pi. 8, figs. I, 9, L9, 30; L898, p. L5. 
Calanus minor Wheeler, lSJlit, |>. Ki5, fig. L'. 

Fig. 2. — Calanus minor. Female, lateral, X31. 

5 Head fused with thorax, forehead and lateral edges of last 
thoracic segment rounded. Anterior antennae not as long as the 
body. Distal margin of second basal in second to fourth pairs 
of feet with a tooth; the proximal outer marginal spine of the 
terminal joint of the outer ramus, in the second to fourth pairs, 
respectively, divides the margin into portions as 5 : 4, 10 : 7, 2 : 1. 
Inner margin of first basal of fifth feet straight, more coarsely 
dentate than in C. finmarchicus. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 127 

J Anterior antennae bent into S-shape ; joints 3, 4 and 5, and 
24 and 25 fused. Right fifth foot with 2 bristles on inner margin 
of third joint of outer ramus, terminal bristle short; terminal 
joint of left foot with three small bristles. 

Coloration : About as in C. finmarchicus. 

Length : Female, 1.8-2 mm. ; male slightly smaller. 

Occurrence : Not at all abundant, but coming in most catches 
with C. finmarchicus. 

3. Calanus tenuicornis Dana. 

Calanus tenuicornis Dana, 1849, p. 278; 1852, p. 1069; 1855, pi. 

73, figs. 10a, 10b. 
Calanus tenuicornis Giesbrecht, 1892, pp. 90, 129, pi. 6, figs. 12 r 

13; pi. 7, figs. 5, 16, 23; pi. 8, figs. 18, 27; 1898, p. 18. 

Fig. 3.— Calanus tenuicornis. (a) Male, lateral, X31. (&) Left fifth 
foot of male X83. 

$ Head not fused with thorax; forehead and angles of last 
thoracic segment rounded; bristle of furca asymmetrical, outer 
marginal minute. Anterior antennae at least l 1 /^ times as long- 
as the body. Proximal outer marginal spine of terminal joint 
of outer ramus in second to fourth feet, respectively, divides the 
margin into portions as 5 :4, 10 :7, 7 :4. 

J Anterior antennae as in 5> except for fusion of joints 1 
and 2, 3 to 5, 7 and 8, 9 and 10, 24 and 25. Mouth parts re- 
duced ; no bristle on inner margin of outer ramus of either of 
the fifth feet ; terminal bristle of right thorn-shaped, that of the 
left slender ; inner rami of both feet similar. Basal portion and 
two proximal joints of outer ramus of right elongate, terminal 
joint shortened. 

Coloration : A variable amount of red or orange in antennae- 
and mouth parts and oil drops of the same color in body. 

128 University of California Publications. [Zoology 

Length: Female, 1.8-2 mm.; male, 1.5-1.8 mm. 
Occurrence : Fairly abundant, both in summer and winter 

4. Calanus gracilis Dana. 

Calanus gracilis Dana, 1849, p. 278; 1852, p. 1078; 1855, pi. 

74, fig. 10. 
Cetochilus longiremis Claus, 1SG3, p. 171, pi. 26, fig. l. 
Calanus gracilis Brady, 1883, p. 35, pi. 5, figs. 1-6; pi. 6, fig. 10. 
Calanus gracilis Giesbrecht, 1892, pp. 90, 128; pi. 6, fig. 1; pi. 7, 

fig. 26; pi. 8, figs. 2, 4, 6-8, 12, 16, 26; 1898, p. 17. 

Fig, l.- Calanus gracilis, (a) Genital Begmenl of female, Lateral, X83. 
(fo) Outer margin >>i first basal of anterior mazilliped of 
female ■ 1 10. 

2 Head fused with thorax; forehead and sides of lasl thoracic 
segmenl rounded. Left side of furca with one elongated bristle. 
Anterior antennae at Leasl 1 ' L . times as Long as the body. There 
is a process a1 the base of the inner marginal bristle of the second 
basal of thesecond fool (cf. fig. bd . The proximal outer marginal 
spine of terminal joint of outer ramus in second to fourth feet, 
respeel ively, divides the margin into porl ions as 1: 1, 4:3, 4: 3 in 
length. First basal of lii'ih pair with feathered Inner margin. 

J Head separate from thorax, anterior antennae straight, 
joints 1 and 2, 24 and 25 fused. Mouth pails greatly reduced. 
Outer border of third joint of outer ramus denticulate in second 
t<> fourth feet. Right foot of fifth pair and inner ramus of* the 
left as in the preceding pairs: basals and first two joints of left 
foot elongate, third joint shortened, no bristle on inner border of 
outer ramus. 

Coloration: Transparent, with little or no pigment in body. 

Length : Female, 2.4 mm. 

Occurrence : San Diego, July 14, 1903, one female ; December 
23, 1903, 14 females. 

Vol. 2] Estcrhj.—Copepoda of the San Diego Region. 129 
5. Calanus robustior Giesbrecht. 

Calanus robustior Giesbrecht, 1888, p. 332; 1892, pp. 91, 129; 

pi. 7, figs. 15, 19, 25, 30; pi. 8, fig. 34; 1898, p. 18. 
Calanus comptus Scott, T., 1893, p. 26, pi. 5, figs. 46-50; pi. 6, 

figs. 1-5. 

Fig. 5.— Calanus robustior. (a) Female, lateral, X36. (b) Outer mar- 
gin of first basal of anterior maxilliped of female X140. 
(e) Genital segment of female, lateral, X83. (d) Inner 
ramus of second foot of female X185. B.2, second basal 
of foot. Ri.l, first joint of inner ramus. Si., inner mar- 
ginal bristle. 

Allied to C. gracilis, but in the female the ventral surface of 
the genital segment is much more convex (cf. figs. 4a and 5c), 
and the first basal of the anterior maxilliped has a bulging 
protrusion on the outer border (cf. figs. 46 and 56). 

J Bristles on anterior maxilliped longer than in C. gracilis, 
inner ramus of left fifth foot stylet-like, jointed and without 
bristles, outer ramus much elongated. 

Coloration : As in C. gracilis. 

Length: Female, 3.17 mm. 

Occurrence: San Diego, July 14, 1903, one female; December 
21, 1904, December 29, 1904, one female each day. 

130 University of California Publications. [Zoology 

Sub-fam. Eucalaxixae. 
Eucalanina Giesbreeht, 1892, p. 45. 

5 Body elongate, head for the most part much lengthened 
(figs. 6a, b) and seldom distinct from the first thoracic segment. 
Rostral filaments slender, abdomen usually with three segments, 
seldom with four; furca often fused with the anal segment. First 
and second and eighth and ninth joints of anterior antennae 
fused. The swimming feet, and especially the rami, are short in 
comparison with the Length of the body ; inner ramus of first pair 
1- or 2- jointed, 3-jointed in the following pairs. Terminal bristle 

of miler rami with sn 1h edge, that of the first pair as in the 

succeeding three pairs; fifth pairabsenl or nniramous ; if present, 
with from three to five joints. 

J Body, especially the head, shortened (fig. 6c); anterior 

antennae without reduction in number of joints: furca as in the 

female. The mouth parts may be stunted. Fifth pair of feet 
not well developed, left foot uni- or biramous, right aniramous 
or lacking. 

1. Genus Eucalanus Dana. 

Calanus Dana, 1848, p. 11 ; 1849, p. 278. 

Eucalanus Dana, IS~>_. \>. 1047. 

Eucalanus (in part) Lubbock, L856, p. 13; 1860, p. 160. 

Calanella Claus, L863, p. 174; not Eucalanus Claus, 1881, \>. 325. 

Eucalanus Brady, l s v>. p. 37. 

Eucalanus Giesbreeht, L888, p. 333 : 1892, pp. 46, 131, 739; 1895, 

p. 246; L898, p. 19. 
Eucalanus Wheeler, 1899, p. 166. 

Anal segment and furca fused, hitler asymmetrical; head 
triangular, often elongate, fused with thorax: abdomen short, 
that of female with three or four segments, of the male with 5. 
Anterior antennae longer than body, 23-jointed in female, term- 
inal bristles plumose and colored. Outer ramus of mandible 7- 
or S- jointed and shorter than inner. Mandible of female longer 
than maxilla: second basal of mandible makes with the outer 
ramus a cylindrical body on which the inner ramus articulates 
proxinially to the outer ramus (fig. 7c). Inner ramus of posterior 
maxilliped with long bristles. Swimming feet short; outer rami 
3-jointed, inner ramus of first pair 2-jointed, of second to fourth 
pairs 3-jointed. Fifth pair absent in female; in male (fig. 6d) 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 131 

both are imiramous. The left 4-jointed, the right 1- to 4- jointed 

or lacking. Head appendages of male retrograded and modified, 

body shortened. 


Abdomen with 3 or 4 segments ? 

Abdomen with 5 segments <$ 

$1. Two segments between anal and genital segments (fig. 6a) 

E. elongatus 

1. One segment between genital and anal segments (fig. 76) 2 

2. Inner border of second basal of mandible divided into two 

approximately equal portions by the insertion of the inner 
ramus (fig. 7c) E. attenuatus 

2. Proximal portion much longer than distal 3 

3. Two terminal bristles of left side of furca longer but hardly 

thicker than on the right side; genital segment (fig. 8a*) much 

broader than long, onion-shaped E. crassus 

3. Two terminal bristles of left side of furca longer and much 
thicker than on the right side. Second basal of maxilla with 
four inner marginal bristles; forehead (fig. 9a) triangular, 

rounded in front E. subtenuis 

Jl. Both feet of fifth pair present 2 

1. Eight foot of fifth pair absent 3 

2. Outer ramus of posterior antenna does not extend by far to the 

distal border of the first joint of the outer ramus E. elongatus 

2. Outer ramus reaches almost to the distal border of first joint of 

i nner E. attenuatus 

3. Terminal joint of fifth foot at least as long as the apical bristle; 

outer border of second joint of inner ramus of second to 

fourth (as in fig. 8a) feet without tooth E. crassus 

3. Terminal joint of fifth foot shorter than apical bristle. . .E. subtenuis 

1. Eucalanus elongatus Dana. 

Calanus elongatus Dana, 1848, p. 18; 1849, p. 278; 1852, p. 1079; 

1855, pi. 75, figs, la, b. 
Eucalanus elongatus Giesbrecht, 1892, pp. 131, 149, pi. 11, figs. 

2, 7, 12, 20, 25, 32, 36; 1895, p. 246; 1898, p. 20. 

5 Two free segments between genital and anal, former 
longer than broad ( fig. 6a, 6). Forehead of regular triangular 
shape. First and second joints of outer ramus of posterior anten- 
nae not fused, first joint of inner ramus little longer than the 
second and over three times as long as broad. Inner margin of 
second basal of mandible with three bristles. End of inner ramus 
does not reach distal end of second basal by about the length of 
the ramus ; its first joint with 2, the second with 5 bristles. Sec- 


University of California Publications. [Zoology 

ond lobe of inner margin of maxilla present, third lobe with four, 
second basal with 5 bristles. First joint of inner ramus of pos- 
terior maxilliped with 3 bristles, second with 4. 

Fig. 6.— Eucalamis elongatus. (a) Female, dorsal, X15. (b) Female, 
lateral, X15. (<•) Bead of male, dorsal, X37. (c?) Fifth 
foot of male, X40; left foot at left of figure 

J Pronounced secondary sexual characters; right fifth foot 
present, left as long as the fourth foot exclusive of terminal 

Coloration: Very transparent, with a small though varying 

amount of red in the body, usually limited to a single oil-drop 
near the posterior cud of the thorax. (See note also.) 

Length: Female, 4.4-7.5 mm.; male. 4 mm. 

Occurrence : A very common species, present in practically 

all hauls with the 000 net. both in winter and summer. 

The most numerous specimens belong to a variety of E. elongatus as in 
Giesbrecht's monograph (1892), in which the last thoracic segment is 
rounded instead of pointed. But Giesbrecht, 1895, p. 246, calls attention to 
this difference. The typical form with pointed thoracic segment occurs in 
the San Diego region, and so far one specimen has been taken, a female, 
length iy-2 mm. The bristles on the posterior maxillipeds and the maxilla 
are faintly orange, those on the feet a rich orange, as far as seen, the feet 
being badly broken. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 133 
2. Eucalanus attenuatus Dana. 

Eucalanus attenuatus Dana, 1848, p. 18; 1849, p. 278; 1852, p. 

1080; 1855, pi. 75, figs. 2a-e. 
Calanus mirdbilis Lubbock, 1856, p. 16, pi. 5, figs. 1-6. 
Calanella mediterranea Claus, 1863, p. 176, pi. 28, figs. 6-11. 
Eucalanus attenuatus Giesbrecht, 1892, pp. 131, 150, pi. 3, fig. 1; 

pi. 11, figs. 1, 11, 13, 16, 18, 24, 40; pi. 35, figs. 3, 6, 17, 

25, 34, 37; 1898, p. 20. 


f.M.'seg. r\. 
Fig. 7.— Eucalanus attenuatus. (a) Head of female, dorsal, X20. (&) 
Abdomen of female, X31. Gen. seg., genital segment. F.+a. 
seg., furca and anal segment, (c) Mandibular rami, female, 
X31. Hi., inner ramus. 

2 Forehead (fig. 7a) triangular, indented on each side, much 
tapering. Genital segment longer than broad, between it and 
anal segment but one free segment (fig. 76). First joint of inner 
ramus of posterior antenna 4 times as long as broad and 1% 
times as long as the second ; two inner marginal bristles on second 
basal of mandible (fig. 7c), end of inner ramus distant from 
distal end of second basal more than the length of the ramus. 
Maxilla as in E. elongatus; first joint of inner ramus of posterior 
maxilliped with 3 bristles, second with four. 

J Pronounced secondary sexual characters; right fifth foot 
present, left considerably shorter than the fourth foot. 

Coloration : Similar to that of E. elongatus; I have never seen 
animals with the plumes at the ends of the antennae entire; in 
Wheeler's specimens they were colorless; in Giesbrecht 's at 
times orange and iridescent. 

Length: Female, from 4 to less than 5 mm.; male, under 3.5 

Occurrence : A few come in the hauls with elongatus, but are 
not nearly so common. They were especially abundant in June 
and July, 1903. 


University of California Publications. 


3. Eucalanus crassus Giesbrecht. 

Eucalanus crassus Giesbrecht, 1888, p. 333 ; 1892, pp. 132, 151 ; 
pi. 11, figs. 8, 10, 17, 21, 22, 38; pi. 35, figs. 4, 20, 26-28; 
1898, p. 22. 

Pig. 8. — Eucalanus crassus. (a) Fourth fool of female X<83. (b) Head 
of female, lateral, X 18. (<■') Head of female, dorsal, X 18. 

('/) Abdomen of female, ventral, X83. 

9 Genital segmenl (fig. s '/ much broader than long, onion- 
shaped : between it and ana] segmenl but one free segment. Fore- 
Ih.kI (fig. 8c) flatly rounded, furca and second terminal bristle 
slightly asymmetrical. First two joints of outer ramus of poste- 
rior antennae fused, first joint of inner ramus shorter than 
second ami about twice as long as broad. Inner ramus of man- 
dible reaches the distal margin of second basal ; first joint of 
inner ramus with two bristles, second joint with four. Second 
lobe on inner margin of maxilla absent, third lobe with three, 
second basal with 4 bristles; first and second joints of inner 
ramus of posterior maxilliped with 3 bristles. 

£ Secondary sexual characters not pronounced; right foot of 
fifth pair absent. 

Coloration : Transparent ; there was no pigment in my speci- 

Length: Female, 3 mm. 

Occurrence: San Diego, June 16, 1904, one female. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 135 
4. Eucalanus subtenuis Giesbrecht. 

Eucalanus subtenuis Giesbrecht, 1888, p. 333; 1892, pp. 132, 150, 
pi. 11, figs. 4, 23, 42; pi. 35, figs. 9-11, 18, 29, 30; 1898, 
p. 21. 

Fig. 9. — Eucalanus subtenuis. (a) Head of female, dorsal, X20. (b) 
Head of female, lateral, X20. 

$ Genital segment somewhat longer than broad, between it 
and anal segment one free segment; forehead (fig. 9a) as in E. 
attenuatus, but less prolonged and not indented on sides. First 
joint of outer ramus of posterior antennae fused with second, 
first joint of inner ramus 3 times as long as broad, and as long 
as second joint ; second basal of mandible with 2 bristles on inner 
border, first joint of inner ramus with 2 bristles, second with 4 ; 
second inner lobe of maxilla absent, third with 4 bristles, second 
basal with 4; same number on first joint of inner ramus of pos- 
terior maxilliped. 

J Secondary sexual characters not pronounced; right fifth 
foot absent. 

Coloration : Transparent, without pigment. 

Length : Female, 2.7 mm. 

Occurrence : San Diego, June 16, 1904, one female. 

2. Genus Rhincalanus Dana. 

Calanus Dana, 1848, p. 11; 1849, p. 278. 

Bhincalanus Dana, 1852, p. 1082; 1855, pi. 76, figs. 2a-d. 

Rhincalanus Brady, 1883, p. 40. 

Bhincalanus Giesbrecht, 1888, p. 334; 1892, pp. 47, 152, 7G1 ; 

1898, p. 22. 
Bhincalanus Scott, T., 1893, p. 30. 

5 Five segments in cepalothorax (head and thorax fused), 
fourth and fifth thoracic segments distinct. Head similar to 
Eucalanus attenuatus, but produced into a snout-like process 
(fig. 10a). Abdominal and thoracic segments with spines; abdo- 
men with 3 segments, furca fused with last segment, and asym- 


University of California Publications. 


metrical. Anterior antennae much longer than body, 23-jointed 
(joints 1 and 2, 8 and 9 fused) ; rami of posterior antennae equal 
in length: mandible not longer than the maxilla. Swimming 
feet short, rami of first pair 2-jointed, of second to fourth pairs 
3- jointed. Fifth foot (fig. 106) uniramous, present on both sides, 
each 3-jointed; second joint with one plumose bristle, third with 
two; a thicker bristle a1 end of third joint, plumose on inner 

J Anterior antennae shortened; fifth foot on the left side 

Lef1 fifth loot biramous J 

Both feel of fifth pair uniramous 5 

1. Rhincalanus nasutus Giesbrecht. 

Rhincalanus nasutus Giesbrecht, L888, p. 334; L892, pp. 152, 160; 
pi. 3, fig. 6; pi. 9, figs. 6, li; pi. L2, figB. 9-12, 1-1. 16, 17; 
pi. 35, figs. 40. 17. 4!>; ISSIS. p. -2'2. 

Fig. 10.— Rhincalanus nasutus. (a) Female, dorsal, X18. (ft) Fifth foot 
of female X260. 

$ Front of head elongated; indented on the sides (fig. 10a); 
rostral filaments ventral, not visible from above. Fifth foot with 
one bristle on second joint, three on third (fig. 106.) 

J Eight fifth foot with strongly curved bristle at end; left 
with long outer ramus reaching almost to end of inner ramus. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 137 

Coloration: Transparent; small amount of red on sides of 
body and faint yellowish tinge to whole, distinct from the glass- 
like clearness of Eucalanus. 

Length: Female, 3 mm.; male, 2.7 mm. 

Occurrence: A female was first seen. San Diego, June 10, 
1904, but two or three occur in most catches when Eucalanus is 

3. Geims Mecynocera [. C. Thompson. 

Leptocalanas Giesbreeht, 1888, p. 334. 
Mecynocera Thompson, I. C, 1888rt, p. 150. 
Mecynocera, Giesbreeht, 1892, p. 160; 1898, p. 23. 
Mecynocera Wheeler, 1899, p. 167. 

Pure a symmetrical, articulating with anal segment ; mandible 
shorter than maxilla and less than half as long as the fourth pair 
of feet, similar in structure to that of Calanus, but inner ramus 
is nearly as Long as second basal and twice as long as outer 
ramus; inner ramus of posterior maxillipeds at least as long as 
first or second basal. Firsl pair of feet with outer ramus of three 
joints, inner of one joint; fifth pair present, with five joints on 
either side. J Unknown. 

5 Head distinct from thorax: rostral threads delicate; abdo- 
men short, with three segments; genital segment and furca sym- 
metrical. Anterior antennae of unequal length, more than twice 
as long as the body, with 23 joints, bristles few and very long. 
Inner ramus of posterior antennae nearly twice as long as outer 
ramus. The succeeding appendages, similar to those of Calanus: 
feet short, outer rami with three joints, inner ramus of first pair 
with one joint, of second to fourth with three; fifth pair with 
basals. outer ramus with three joints, inner ramus lacking. 

1. Mecynocera clausi I. C. Thompson. 

Mecynocera clausii Thompson, I. ('., 1888a, p. 150, pi. 11, figs. 1-4. 

LeptOCalanus filiform:* (iiesbvotht, isss, 1'. 334. 

Mecynocera clausii Giesbreeht, L892, p. 1(50, pi. 5, fig. 1; pi. 11, 

ags. 13, 4.1; pi. 35, figs. 21, 22; .1/. clausi, 1898, p. 23. 
Mecynocera clausii Wheeler, 1S99, p. 107, fig. 5. 

The only species of the genus. 

Coloration: Exceedingly transparent, without pigment in my 

138 University of California Publications. [Zoology 

Length: Female, 0.9-1 nun. 

Occurrence : The only specimens I have were collected Decem- 
ber 30, 1903, on the "Banks" off Point Loma. 

Fig. 11.- Mecynoeera clausi. Female, dorsal, ■ r>. 

Sub-fam. Paracalaninae. 
Paracalanina Giesbrivlit. lsi'i'. \>. 4s. 

$ Cephalothorax with four segments, abdomen with from two 
to four-, rostrum ends in two soft filaments. Anterior antennae 
25-jointed, with long terminal joints, but the division between the 
first and second and eighth and ninth joints may not be clear. 
Outer ramus of posterior antennae at most as long as the inner 
ramus; the mouth parts like those of Calanus. Terminal bristle 
of outer rami of feet with smooth border; basals and rami set 
with spines; fifth foot rudimentary (2- 4- jointed) or lacking. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 139 

J Characters as in the male of Calanus; the number of joints 
in the anterior antennae more reduced, the end-joint always 
shortened, and sometimes fused with the preceding' one; fifth pair 
of feet weakly developed, the left 5-jointed, the right 4- or 2- 
jointed, or lacking. 

1. Genus Paracalanus Boeck. 

Calanus Glaus, 1863, p. 172. 

Paracalanus Boeck, 1864, p. 8. 

Paracalanus Glaus, 1881, p. 326. 

Paracalanus Bourne, 1889, p. 145. 

Paracalanus Giesbrecht, 1892, pp. 48, 164, 757; 1898, p. 23. 

Paracalanus Dahl, 1893, p. 21. 

Paracalanus Wheeler, 1899, p. 168. 

Second basal of first pair of feet with an inner marginal 
bristle ; proximal division of outer border of third joint of outer 
ramus of fourth pair (fig. 12a) over twice as long as the distal; 
outer border of the second joint not dentate ; proximal division 
of the outer border of third joint of outer ramus in the third and 
fourth feet dentate ; scalpelliform terminal bristle of the outer 
ramus in the third pair longer than the end joint; second joint 
of inner ramus of first pair with 5, third of same in second pair 
with 7 bristles. The abdomen of the female (fig. 12b) with 4 seg- 
ments; the last joint of anterior antennae less than iy 2 times as 
long as the next to the last. Fifth foot of female short, 2-jointed 
(fig. 12c) ; right foot of male with 2 joints, left with 5 (fig. 12c). 

5 Head fused with first thoracic segment, and fourth thoracic 
segment with fifth. Rostrum produced into two thin filaments. 
Genital segment and furca symmetrical, latter without bristle on 
outer margin. Anterior antennae with 25 joints. Outer ramus 
of posterior antennae shorter than inner; mandible with broad 
blade, the sack-like appendage on the first joint of the inner 
ramus small. Maxilla with obscure segmentation of inner ramus, 
without bristle on the second lobe of outer border, and with but 
one on the first inner marginal lobe. Anterior maxilliped with 
outer marginal bristle. Inner ramus of the first swimming foot 
with 2 joints, of the second to fourth foot with 3 joints. 

J Abdomen with 5 segments. Number of joints of anterior 
antennae reduced through fusion of joints 1 to 6 and 7 to 8, end 
joint shortened but free. Aesthetasks enlarged and numerous. 


University of California Publications. [Zoology 

Mandibular blade, appendages on inner border of maxilla and 
anterior maxilliped stunted, those of posterior maxilliped less so, 
its outer marginal bristles long and richly plumose. The swim- 
ming feet show slight peculiarities. 

1. Paracalanus parvus Glaus. 

Calanvs parvus Claus, 1863, p. ]7:i, pi. 26, figs. 10-14; pi. 27, figs. 

Paracalanus parvus Claus, 1881, p. 327, pL 3, figs. 1-16. 
Paracalanus parvus Bourne, 1889, p. 145, pi. 11, figs. 1-3. 
Paracalanus parvus Giesbrecht, 1892, pp. 164, 170; pi. 1, fig. 5; 

pi. 6, figs. 28-30; pi. 9, figs. 511. 25, 27. 31, 52 ; !S!«s. p. 24. 

Fig. 12. — Paracalanus parvus, (a) Fourth foot of female X195. Ee.3, 
third joint of outer ramus, (b) Female, dorsal, X83. (c) 
Fifth foot of female X410. (d) Second joint of inner 
ramus of second foot of female X195. (c) Fifth foot of 
male. Ps., left foot. Pel., right foot. 

5 Inner bristle of furca barely longer than the furca. Ante- 
rior antennae reach, when brought to the sides of the body, per- 
haps to the posterior border of the third abdominal segment. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 141 

First joint of inner ramus of maxilla with two bristles on ante- 
rior face. Third lobe of second basal of posterior maxilliped with 
two bristles. Inner margin of first basal of the fourth pair of 
feet ends in one or two points (fig. 12a) ; anterior and posterior 
faces of first basal of second to fourth pairs set with hairs and 
spines ; surfaces of first and second joints of outer ramus of the 
third pair and of second joint of the fourth, naked. Fifth foot 
rudimentary, symmetrical. 

J Fifth foot asymmetrical (fig. 12e) ; compare also generic 

Coloration : Rather transparent, with red pigment in varying 
amounts and distribution, never very abundant. 

Length : Both sexes within 0.8-1.2 mm. 

Occurrence: Fairly common in hauls with smaller nets, both 
sexes being present summer and winter. 

Sub-fam. Clausocalaninae. 
Clausocalanina Giesbrecht, 1892, p. 49. 

5 Head usually fused with the first thoracic segment, fourth 
thoracic always fused with the fifth ; rostrum ends in two short, 
soft filaments or is lacking; abdomen with four segments, furca 
symmetrical. Eighth and ninth joints of anterior antennae 
fused ; terminal joint short, seldom fused with the preceding one. 
Outer ramus of posterior antennae 6-jointed and always longer 
than the inner ramus. The other appendages of the head for the 
most part as in Calanus. Inner ramus of the first pair of feet 1- 
jointed, of the second pair 2-jointed, of the third and fourth 3- 
jointed; terminal bristle of the outer rami with dentate border; 
third joint of outer ramus in second to fourth pairs with three 
bristles on outer border. Fifth pair rudimentary on each side, 
3-jointed or lacking. 

J Unknown in Spinocalanus and Ctenocalanus. Abdomen 
with shortened anal segment; anterior antennae and head 
appendages in some cases like those of the female, in others as in 
the Paracalaninae. Fifth pair of feet: the right, 1- to 5-, the left 
5- jointed. 

14l } University of California Publications. [Zoology 

1. Genus Clausoc alarms Giesbrecht. 

Calanus Dana, 1849, p. 278; 1852, p. 1047. 
Calamus Claus, 1863, p. 172. 
Eucalanus Claus, 1881, p. 325. 
Vrepanopus (in part) Brady, 1883, p. 76. 

Clausocalanus Giesbrecht, 1888, p. 334; 1892, pp. 50, 185, 733; 
1898, p. 27. 

Rostrum with two points: second basal of second and third 
swimming feet with toothed distal margin and broad outer ramus. 
Mouth parts and number of segments of anterior antennae 

2 Head fused with thorax and the fourth with the fifth 
thoracic segment. Abdomen with four segments, genital segmenl 
and I'nrea symmetrical. Anterior antennae extend beyond the 
thorax. 23-jointed. Outer ramus of posterior antennae 1 ' . , times 
as Long as the inner, the former 6-jointed, with short bristles on 
the proximal joints. First joim of inner ramus of mandible 
with a vi-iy small, sack like appendage. .Maxilla and maxillipeds 
as in Calanus, miter marginal bristle lacking on anterior maxilli- 
peds. Outer rami of swimming feel with 3 joints: inner ramus 
of lirst foot 1-jointed, of second 2-jointed, of third and fourth 
3-jointed. End joint of outer ramus with finely dentate terminal 
bristle, and four bristles on inner border in second to fourth 
pairs; fifth pair nniramous, 3-jointed. 

J Head fused with lirst thoracic segment, and elongated at 
expense of free thorax rings. Rosl rum suppressed ; abdomen with 
live segments, anal very short. Anterior antennae with joints 
8-10,13-16,20-21, L'4-25 fused. Outer ramus of posterior antennae 
twice as Long as inner. Blade of mandible, appendage of inner 
border of maxilla, and anterior maxilliped suppressed; less so the 
posterior maxilliped. the outer marginal bristle of which is not 
enlarged. Swimming feet elongated. Left fifth foot (fig. 13c) 
Long, nniramous and with 5 joints, right short. 1- to 3-jointed. 

1. Clausocalanus arcuicornis Dana. 

Calami* arcuicornis Dana, 1S49, p. 278; 1852, p. 1056; 1855, 

pi. 72, fig. 9a-b. 
Calanus mastigophorus Claus, 1863, p. 173, pi. 27, figs. 5-8. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 148 

Clausocalanus arcuicornis Giesbrecht, 1888, p. 334; 1892, pp. 186, 
193; pi. 1, fig. 14; pi. 2, fig. 7; pi. 10, figs. 3-8, 14, 16, 17^ 
19; pi. 36, figs. 29-31, 34; 1898, p. 27. 

Clausocalanus arcuicornis Wheeler, 1899, p. 171, fig. 9. 

Fig. 13.— Clausocalanus arcuicornis. (a) Male, lateral, X45. (b) Second 
basal of second foot to show toothed distal margin, X410. 
(c) Fifth foot of male X83. 

$ Genital segment longer than the two following. Furca 
about as long as broad. No aesthetask on fourth, sixth, eighth, 
eighteenth or twenty-second joints of the anterior antennae. 

J Second segment of abdomen at least as long as the third 
and fourth together (fig. 13a) ; right foot of fifth pair with three 
joints (fig. 13c). 

Coloration : Not very transparent, with red pigment in vari- 
ous places on the posterior part of the body and on genital seg- 

Occurrence: San Diego, June 25, 1904, one male. 

Sub-fam. Aetidiinae. 

Aetidiina Giesbrecht, 1892, p. 52. 
Aetidiinae Wolfenden, 1903, p. 263. 

9 Head sometimes distinct from first thoracic segment ; other- 
wise the cephalothorax always has four segments, as has the 

144 University of California Publications. [Zoology 

abdomen invariably. Rostrum strongly ebitinized, usually with 
one point, seldom with two or lacking-. Genital segment and 
furca usually symmetrical. In the anterior antennae the eighth 
and ninth and twenty-fourth and twenty-fifth joints are fused. 
Outer ramus of posterior antennae at least fully as long as the 
inner, and usually longer; the second and third joints of the 
outer ramus are distinct. Mandible as in Calanus, with strong 
blade, and occasionally shortened inner ramus. Maxilla with 
well developed lobes on inner margin and usually with hooked 
bristles even on the second basal and the inner ramus; outer 
ramus relatively small. Bristles of anterior maxilliped short but 
strong, those of the inner ramus relatively slender and sparsely 
plumose; the articulation of the inner ramus is rather on the 
posterior surface of the second basal than at the end. Inner 
ramus of posterior maxillipeds a1 most % as long as the second 
basal. Inner ramus of firsl swimming feet always 1-jointed; that 
(if the second almosl always I- join ted, while in the second and 
third the inner ramus is 3-jointed; the form of the swimming 
feet as in the Clausocalan inae : inner marginal bristle of first 
basal long and plumose. 

J Known in Aetideus, Euchirella and Undeuchaeta, Charac- 
ters like those of Clausocalanus; occasionally the twentieth 
and twenty-firsl joints of one of the anterior antennae arc fused. 
Left foot of fifth pair 5-jointed (if the righ.1 is lacking, or stylet- 
like, in which ease the right is claw-like). 

1. Genus Aetideus Brady. 

Aetidius Brady, 1883, p. 75. 

dius Thompson, ls.ssb. p. Ml'. 
Aetidius Giesbrecht, 1S92, pp. 53, 213. 
Aetideus Wolfenden, 1903, p. 266; 1904, p. 116. 
Aetideus Giesbrecht, 1898, p. 31. 

$ Cephalothorax and abdomen with four segments, symmet- 
rical; rostrum large, prolonged into two thick chitinous prongs; 
last thoracic segment produced into a spine on each side. Ante- 
rior antennae 23-jointed, reaching about to the end of body. Rami 
of posterior antennae about equal in length, outer ramus 7- 
jointed. Outer rami of all swimming feet 3-jointed, inner ramus 
of first and second pairs 1-jointed, of third and fourth 3-jointed. 
Fifth pair of feet absent. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 145 

J Anal segment very short, abdomen with five segments. 
Anterior antennae 20-jointed, joints 8-10, 12 and 13, 20 and 21, 
24 and 25 fused. Blade of mandible, appendages of inner border 
of maxilla and anterior maxilliped stunted. Left fifth foot uni- 
ramous, 5-jointed ; right lacking ; swimming feet as in female. 

1. Aetideus armatus Brady. 

Aetidius armatus Brady, 1883, p. 75, pi. 10, figs. 5-16. 

Aetidius armatus Giesbrecht, 1892, p. 213, pi. 2, fig. 6; pi. 14, 

figs. 1-13; pi. 36, figs. 6-9. 
Aetideus armatus Giesbrecht, 1898, p. 31. 
Aetideus armatus Wolfenden, 1903, p. 266. 

Pig, 1A.— Aetideus armatus. Female, (a) Animal from side X20. (6) 
Second foot X83. St., terminal bristle of outer ramus. 

With the characters of the genus. 

Coloration : Rather transparent ; there was no pigment in my 
specimens, but Giesbrecht says that red may occur in the body. 

Length : Female, 3 mm. 

Occurrence: San Diego, June 9, 1904, one female; June 14, 

two females. 

2. Genus Gaidius Giesbrecht. 

Gaidius Giesbrecht, 1895, p. 249 ; 1898, p. 32. 

Gaidius Wolfenden, 1902, p. 365; 1903, p. 266; 1904, p. 114, pi. 
9, figs. 7, 8. 

Rostrum short, one point (fig. 15b), sides of last thoracic seg- 
ments produced into a sharp spine (fig. 15a). Inner ramus of 
posterior antennae three-fourths as long as outer. Outer ramus 
of first foot 2-jointed, of the second to fourth 3-jointed, inner 
ramus of first and second feet 1 -jointed, of the third and fourth 


University of California Publications. [Zoology 

May be distinguished from Aetideus by the form of the ros- 
trum, relatively shorter inner ramus of the posterior antennae. 
and by the fusion of the proximal joints of the outer ramus of 
the first foot (fig. 15c/). 

1. Gaidius pungens Giesbrecht. 

Gaidim pungens Giesbrecht, 1895, p. 249, pi. 1, figs. 1-4; 1898, 

p. 32. 
Gaidius pungt ns Wblfenden, 1903, p. 266. 

Fig. 15.— Gaidius pungens. Female. («) Dorsal, X18. (b) Head, lateral, 

X45. (r) Tube-like processes on inner distal portion of 

second basal of fourth foot X195. (d) First foot X195. 
/,'(.. inner ramus. 

5 Anterior antennae reaching at least to posterior border of 
thorax. The processes on the inner border of the first basals of 
the fourth feet are heavier and stiffer than in the preceding- 
pairs, being almost tube-like (fig. 15c). 

J Unknown. 

Coloration: Transparent, with little or no pigment. 

Length : Female, 3 to 3.5 mm. 

Vol. 2] Esterhj. — Copcpoda of the San Diego Region. 147 

Occurrence: San Diego, May 31, 1904, eight females; two 
males which seem to be of this species were taken also at this 
time, but they are distinctly immature. 

3. Genus Undeuchaeta Giesbrecht. 

Euchaeta (in part) Brady, 1883, p. 57. 

Undeuchaeta Giesbrecht, 1888, p. 335; 1892, pp. 54, 227, 766; 

1898, p. 33. 
Undeuchaeta Sars, 1900, p. 58, pis. 15, 16. 
Undeuchaeta Wolfenden, 1903, p. 267. 

5 Abdomen with four segments, the first with the genital open- 
ing on the convex ventral surface, at least as long as the second 
and longer than the last segment, Lateral angles of last thoracic 
segment rounded, or at least not produced into spines. Anterior 
antennae 23-jointed, outer ramus of first foot 2-jointed, inner 
ramus 1-jointed. Outer ramus of posterior antennae at least 1V 2 
times as long as the inner; outer ramus of maxilla (fig. 16e) 
small, middle bristles shorter than the distal and proximal ones, 
outer marginal lobe with much elongated middle bristles. 

J Anterior antennae 21-jointed, cephalo-thorax with four 
segments, abdomen with five, anal segment very short. Head with 
rather high crest (fig. 16d), last thoracic segment prolonged into 
angles, but not pointed. Inner ramus of posterior antennae % 
as long as the outer. Mandible, maxilla and maxillipeds much 
reduced. Outer ramus of first foot indistinctly 3-jointed. 
Left foot of fifth pair uniramous (inner ramus reduced 
to a very small, rod-like projection), outer ramus (fig. 16/) end- 
ing in a short style (terminal joint of ramus). Right foot bira- 
mous. Terminal joint of outer ramus produced into a long 
stylet, inner ramus as in Euchaeta (cf. fig. 23a) ; outer ramus of 
each foot 3-jointed. The second joint of the outer ramus of the 
left foot (fig. 16 f. Be. 2) bears a toothed process (fused with the 
joint) which flares distally ; at the base of this and on the second 
joint is articulated a process, which together with the terminal 
joint of the ramus and the toothed process forms a forceps. 

The abdominal segments are densely covered with fine spines 

or hairs, and the posterior margins of the segments are toothed. 

In the structure of the fifth pair of feet these male animals 

very closely resemble the males of the genus Euchaeta, but seem 

148 University of California Publications. [Zoology 

to be distinct from the latter in bearing an articulating process 
on the second joint of the outer ramus of the left foot. There is 
a muscle attached to the process which serves to move it. 

The relative lengths of the rami of the posterior antennae dis- 
tinguish the animals from Euchirella, as does the division 
(though indistinct i of the outer ramus of the first foot into 
three joints. In Euchaeta, the outer ramus of the first foot is 
distinctly 3-jointed in the male, and the rami of the posterior 
antennae are about equal in length. In several female speci- 
mens also the outer rami of the first feet are indistinctly divided 
into three joints, and the sexes correspond in this respect. 

Sars (1900, p. 59-63) has described the male and female of 
Undeuchaeta spectabilis. So far as I know, his is the first record 
of the male of the genus. In his specimens the anterior antennae 
of the female are 24-jointed, while in I Hesbreehl *s the Dumber of 
joints is 2:5. In his description of the male. Sars gives the num- 
ber of joints of the anterior antennae as 22, bu1 in his drawing 
(pi. 16, fig. 2) there are bu1 21. The fifth pair of feel in the 
mule of U. spectabilis is very different from that in the San 
Diego specimens. ;i striking point being that both the right and 
left feci are biramous. 

The description of the male of the genus given above is based 
upon the San Diego specimens. 

1. Undeuchaeta major Giesbrecht. 

Euchaeta australis Brady, 1883, p. 65, pi. 21, figs. 5-11. 
Undeuchaeta major Giesbrecht, 1888, p. 336; 1892, pp. 227, 232, 
pi. 37, figs. 56, 57, 59; 1898, p. 34. 

5 Head with median crest, genital segment with protrusion on 
right side and a hooked pointed appendage at the right of the 
genital opening (fig. 16a). 

J (new) Compare generic description. 

Coloration: Female not especially transparent, with red pig- 
ment on basals of posterior maxillipeds, and in month region. 
Male: plumose bristles of furca steel-blue; those of posterior 
antennae and mandible red; on the feet the bristles are faintly 
red on the outer ends. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 149 

Length : Female, 4.5-5.5 mm. ; male, 6-6.5 mm. 

Occurrence : Four females were taken from May 18 to June 
23, 1904; five on December 23, 1904, on the "Banks." One male 
was taken in July, 1904 ; another on November 1, 1904, off Point 

re 3. 


Fig. 16.— Undeuchaeta major. (a) Female, lateral, X20. Ab.l, first 
abdominal segment. (b) Male, lateral, X9. (c) Head of 
male, dorsal, X83. (d) Head of male, lateral, X83. (e) 
Outer ramus of maxilla of female X195. (/) Distal portion 
of left fifth foot of male. Be.2, Be.3, respective joints of 
outer ramus. 

2. Undeuchaeta minor Giesbrecht. 

Undeuchaeta minor Giesbrecht, 1888, p. 335; 1892, pp. 228, 232, 
pi. 14, figs. 31-34; pi. 37, figs. 55, 58; 1898, p. 34. 

Fig. 17. — Undeuchaeta minor. Female, (a) Lateral, X20. (&) Second 
foot X83. 

150 University of California Publications. [Zoolooy 

9 Head without crest (fig. 17a), genital segment with a spine 

on the dorsal surface. 

Coloration : Similar to that of TJ. major. The digestive tract 

of the single specimen was filled with orange red material. 
Length: Female, 3.18 mm. 
Occurrence : San Diego, June 14, 1904. 

4. Genus Euchirella Giesbrecht. 

Undina (in ]>art) Lubbock, 1856, p. 21. 

Calanus (in part) Lubbock, 1856, p. 15. 

Undina elans 1863, p. 186. 

Euchaeta (in part) Brady, 1883, p. 59. 

Euchirella Gieflbrecht, 1888, p. 336; 1892. pp. 54, 233, 743; 1898, 

p. 34. 
Euchirt lla Cleve, 1900, p. 4. 
Euchirella Wplfenden, L903, p. 267. 

Rostrum present in mosl species, simple: lateral angles of 
last thoracic segmenl Dot pointed, [nner ramus of posterior 
antennae \ L . to ( j as long as the outer ramus, the two proximal 
joints of which are fused, [nner and outer rami of the maxilla 
short, the former provided with heavy hooked bristles. Outer 
ramus of the firsl pair of feet 2-jointed, that of the second to 
fourth pairs 3-jointed. [nner ramus <>f first and second pairs 
1-jointed, of the third and fourth 3-jointed. Right foot of fifth 
pair of male with shear-like formation of distal portion, the left 
foot stylet-like (cf. figs. 18d, 196). 

5 1 not always distim-t from thorax, last two thoracic 
segments fused. Abdomen with four segments, genital segment 
and f ureal bristles symmetrical <>r asymmetrical. Anterior an- 
tennae with 23 joints, reaching to the end of the thorax or some- 
what beyond. Second basal of posterior maxillipeds twice as 
long as the 5-jointed inner ramus. Feathering on the inner 
margin of the proximal basal joint of the fourth pair of feet 
replaced by spines. Fifth pair of feet absent. 

J Head occasionally with a median crest; abdomen with five 
segments, anal segment very short. Joints 20 and 21 of right 
anterior antennae fused, inner ramus of posterior antenna rela- 
tively longer than in the female. Blade of mandible, appendages 
on inner border of maxilla, and anterior maxillipeds reduced; 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 151 

posterior maxilliped slender. Spines on second basal of fourth 
foot unusual. Right foot of fifth pair biramous, with forceps; 
left stylet-like, with rudimentary inner ramus. 


Fifth foot absent 5 

Fifth foot present £ 

$1. Head without crest, and rostrum one-pointed (fig. 19a) 2 

1. Head with crest, rostrum present (fig. 20a) 4 

2. Outer ramus of posterior antenna about twice as long as inner. . 

E. rostrata 

2. Outer ramus nearly four times as long as inner (fig. 20c) 3 

3. Genital segment with long sac-like appendage on left side (fig. 

186) E. messinensis 

4. Head with low crest (fig. 20a) E. pulchra 

4. Head with high crest (fig. 22a) E. galeata 

J Head with low crest (fig. 18c) 2 

Head without crest (fig. 21a) 3 

2. Forceps-like terminal portion of right fifth foot longer than the 

basal portion (fig. 18d) '. E. messinensis 

2. Terminal portion (forceps) of right fifth foot shorter than the 

basal portion (fig. 206) E. pulchra 

3. Fifth foot short, the right about four times as long as the second 

basal is broad (fig. 216) E. amoena 

3. Eight fifth foot six times as long as the second basal joint. .E. rostrata 

1. Euchirella messinensis Claus. 

Undina messinensis Claus, 1863, p. 187, pi. 31, figs. 8-18. 
Euchirella messinensis Giesbreeht, 1892, pp. 232, 244; pi. 15, figs. 
12, 16, 21, 24; pi. 36, figs. 14, 15, 18, 24, 25; 1898, p. 35. 

5 Forehead with rostrum, without crest (fig. 18a) ; genital 
segment asymmetrical, with sac-like appendage on left side of 
dorsal surface (fig. 186) ; third terminal bristle on right side of 
furca elongated. Inner ramus of posterior antenna 14 as long 
as outer, second joint of former with 5-4 bristles. First basal 
of fourth pair of feet with one or two spines on inner border, 
the longer of which reaches beyond the end of the joint. 

J Forehead with a low and rather long crest (fig. 18c) ; fifth 
foot slender, the right foot (fig. 18d) over seven times as long 
as the second basal is broad, the forceps longer than the basals. 

Coloration : Not very transparent ; red pigment in body and 
on bristles of posterior antennae, and basals of swimming feet. 

Length : Female, 4.5 mm. ; male, 4 mm. 


University of California Publications. [Zoology 

Occurrence: July 9, 1903, one male: July 22, 190°,, one 

Fig. 18.— Euchirella messinensis. (a) Eead of female, Lateral, X15. (b) 
A i"lmiicn of female ■ L5. (c) Eead of male X30. (d) 
Fifth foot of male X20. 

2. Euchirella rostrata Claus. 

Undina rostrata Claus, L866, p. 11, pi. 1. fig. 2. 

Euchaeta hessei Brady, L883, p. 63, pi. 20, figs. 1-13; pi. 2.3, figs. 

1 II 1. 
Euchirella rostrata Giesbrecht, 1892, pp. 2:;::, 245, pi. 15, figs. 3, 

L3, 25; pi. 36, figs. L9, 20; L898, p. 36. 
Euchirella rostrata Cleve, L900, p. 1. pi. 2, figs. 1-12. 

$ Front without crest, with rostrum, abdomen symmetrical. 
Inner ramus of posterior antennae ' - ;is long .is outer ramus; 
second joinl of inner ramus with 8-6 lu-istles. Firsl basal of 
fourth pair of feel i fig. 196 ; B. 1) with 6 or 7 triangular Lamellae 
on the inner border. Bristle on outer margin of second joint of 
outer ramus of the second pair reaches ;it least to the point of 
the first bristle on the outer border of the third joint of the 

J Head without crest, with rostrum. Fifth foot six times as 
long as its second basal joint. Margin of second joint of outer 
ramus not denticulate, third joint smooth. Inner ramus of pos- 
terior antenna % as long as outer. First basal of fourth foot 
without triangular lamellae. (Cleve 1900). 

Vol. 2] Estcrhj. — Copepoda of the San Diego Region. 153 

The male was described by Cleve, 1900, and is identical with 
Euchaeta hessei Brady. 

Coloration : Red pigment as in E. niessiiwiisis, but more abun- 
dant, especially on swimming feet. 

Fig. 19.— Euchirella rostrata. (a) Female, lateral, X18. (&) Fourth 
foot, female, X83. B.l, first basal, showing lamellar pro- 

Length : Female, 2.97-3.1 mm. 

Occurrence : San Diego, July 14, 16, 21, 1903, females ; May 
24, 1904, two females; June 2, 1904, one female. 

3. Euchirella pulchra Lubbock. 

Undina pulchra $ Lubbock, 1856, p. 26, pi. 4, figs. 5-8 ; pi. 7, fig. 6. 
Calanus latus $ Lubbock, 1856, p. 15, pi. 2, fig. 12; pi. 11, figs. 


University of California Publications. 


Euchaeta pulchra Brady, 1883, p. 63, pi. 14, fig. 7; pi. 20, figs. 
15, 17, 19. 
Euchirella pulchra Giesbreeht, 1892, pp. 233, 244, pi. 15, figs. 22, 
23, pi. 36; figs. 13, 27; 1898, p. 36. 

Fig. 20.— Euchirella pulchra, (a) Female, lateral, X9. (b) Fifth foot 
of male X45. (c) Rami of posterior antennae, to show 
relative lengths; bristles omitted, X83. 2?!., inner ramus. 
(d) First basal of fourth foot of female X140. 

$ Front with low crest (fig. 20a) and small rostrum. Genital 
segment asymmetrical ; left side strongly convex in front of the 
middle of the segment, right side indented. Inner ramus of pos- 
terior antennae about % as long as outer ramus; second joint of 
inner ramus with 6-5 bristles. First basal of fourth pair of feet 
(fig. 20c?) with one or two thorns about equal in length on the 
inner border, which do not reach the distal margin of the joint, 
Outer bristle of the second joint of the outer ramus of the second 
pair at most as long as the first outer bristle of the third joint. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 155 

J Considerably like E. messinensis, the chief difference being 
in the structure of the fifth pair of feet (fig. 206). The claw of 
the right foot is shorter than the basal (in messinensis longer). 

Coloration : About as in E. messinensis. 

Length : Female, 3.4-4 mm. ; male, 3.5 mm. 

Occurrence : San Diego, May 31, 1904, two immature males, 
one female ; June 23, 1904, one female adult ; December 23, 1904, 
"Banks," eleven females, all adult; one male adult, December 
30, 1904, on the "Banks." 

4. Euchirella amoena Giesbrecht. 

Euchirella amoena Giesbrecht, 1888, p. 336; 1892, pp. 233, 244; 
pi. 15, fig. 20; 1898, p. 36. 

Fig. 21. — Euchirella amoena. 

Male, (a) Lateral X20. (b) Fifth foot 

J Unknown. 

5 Front without crest. Fifth pair of feet shortened, the right 
about four times as long as the second basal is broad. 
Length: Male, 3.02 mm. 
Occurrence : San Diego, May 28, 1904, one male. 

5. Euchirella galeata Giesbrecht. 

Euchirella galeata Giesbrecht, 1888, p. 336; 1892, p. 233, 244; pi. 
15, fig. 18; pi. 36, figs. 22, 26; 1898, p. 36. 

5 Head with high crest, and rostrum ; genital segment asym- 
metrical, strongly protruding on the posterior portion of the 
dorsal surface. Inner ramus of posterior antennae about % as 


University of California Publications. [Zoology 

long as the outer: basals of fourth foot about as in E. pulchra, 
the spines not reaching to the distal border of the joint. 

J Head as in the female. 

Coloration: Opaque, without pigment. 

Length : Female, 6.5 mm. 

Occurrence: San Diego, November 18, 1904, one adult female. 
two immature males. 

Pig, 22.- Euchirella galeata. Female, (a) Lateral X9. (6) Firsl basal 

of fun rlli toot X83. 

Sub-fam. Euchaetinae, 

Euchaetina Giesbrecht, 1892, p, 55. 

$ Rostrum with one point; a pouch-like appendage in front 
of the upper labium. Inner marginal bristle of furca very Long. 
Distal hooked bristles of anterior maxillipeds Longer than the 
proximal. Outer ramus of first pair of feel 2-jointed, of the 
second to fourth 3-jointed; inner ramus of firsl and second pairs 
1-jointed, of the tliir<l and fourth pairs 2-jointed. 

J Abdomen as in the Clausocdlaninae. Outer ramus of firsl 
pair of feel 3-jointed; fifth foot on each side with 2-jointed basal, 
and biramous; inner ramus of Lefl stylet-like, of right truncate: 
1 < ■ it outer ramus 3-jointed, right 2-jointed. 

1. Genus Euchaeta Philippi. 

lutclith fa I'hilippi, 1843, p. 54, pi. 4, fig. 5. 
F.achinis Dana, 1S40, p. 1S3. 

Euchaeta Dana, 1848, p. 20; 1849, p. 279; 1852, p. 1084. 
Euchaeta Clans, 1863, p. 163. 

Euchaeta Giesbrecht, 1892, pp. 55, 245, 740; 1895, p. 251; 1898, 
p. 37. 

5 Cephalothorax with five segments, the last two thoracic 
segments fused; abdomen with four segments, genital segment 
more or less asymmetrical. Anterior antennae of varying rela- 
tive lengths, but of characteristic form, with 23 joints. Rami 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 157 

of posterior antennae abont equal in length, outer ramus with 
seven joints. Blade of mandible with few but strong teeth. Sec- 
ond basal joint of the posterior maxilliped at least three times 
as long as the inner ramus of five joints. Inner marginal bristle 
of first basal of the swimming feet long and richly plumose, 
terminal bristle of outer rami finely toothed ; fifth pair absent. 

J Head fused with thorax ; abdomen with five segments, anal 
segment short ; innermost bristle of f urea shortened and bent at 
an angle. Blade of mandible, appendage of inner border of max- 
illa, and anterior maxilliped stunted ; less obvious differences also 
in the posterior antennae and maxillipeds and swimming feet ; 
outer ramus of first pair of feet 3-jointed. Feet of fifth pair 
long, strongly built, and of rather complicated structure (fig. 



Fifth foot absent $ 

Fifth foot present J 

£1. Terminal joint at each foot of fifth pair, with long straight or 
slightly curved stylet; elevation for frontal organ not pro- 
truding E. acuta 

$1. Hairs of frontal organ on a low elevation (fig. 25b) 2 

1. Hairs of frontal organ on an elevation which extends toward the 

front (fig. 21b) 4 

2. Genital segment with asymmetrical outgrowths (figs. 25c, d) ; no 

bristle in middle of outer border of first joint of outer ramus 
of the first foot; terminal bristles of furca about equal in 
length, the dorsal (inner) bristle much longer and thicker 
(fig. 25g) 3 

3. Genital segment with a knobdike protuberance in front on the 

left side E. acuta 

3. Genital segment without such an outgrowth E. media 

4. Middle spine on outer border of terminal joint of second foot 

longer than the others, and the distal indentation in the border 
deeper (fig. 24c) 5 

5. Anterior antennae longer than body E. spinosa 

1. Euchaeta acuta Giesbrecht. 

Euchaeta acuta Giesbrecht, 1892, pp. 246, 262, pi. 16, figs. 6, 10, 
14, 18, 21, 27, 39 ;pl. 37, figs. 47, 48, 52; 1898, p. 38. 

5 Elevation on front of head flat ; genital segment asymmet- 
rical, more strongly convex on the right side than on the left, 
and with more prominent process at the right of the opening ; a 
knob-shaped outgrowth on anterior part on left side. Furca 


University of California Publications. 


with four terminal bristles nearly equal in Length, inner bristle 
of furea much thicker than end bristles. Anterior antennae 
reach a little beyond the posterior end of the thorax. 

Fig. 23. — JSuchaeta acuta, (a) Fifth foot of male X37. /.'<. dx. f inner 

ramus of right foot. /•'< . :'>. '/./•., third joint of (inter ramus. 
i:< . 1. •_'. sn., first and Becond joints of outer ramus of left 
foot. /'/"<■.. process. Sph. } spermatophore. (b) Second and 
third joints of outer ramus of left fifth foot of male X140. 
Tarts as in a. (<■) Second foot of male X60. (d) Maxilla 
of male X60. B.l, first basal. B.2, second basal. Le.l, first 
lobe of outer margin. Et., inner ramus. Re., outer ramus. 

First lobe of outer border of maxilla (cf. fig. 25/) with six 
bristles ( one very small), second basal with three, fused second 
and third joints of inner ramus with four. Outer border of 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 159 

first joint of outer ramus of first pair of feet concave ; outer 
border of third joint of outer ramus of second pair and its outer 
bristles different than in the following pairs of feet ; outer mar- 
ginal bristle of second joint of outer ramus reaches almost to end 
of the first outer marginal bristle of the third joint. Third joint 
of outer ramus of left fifth foot of male (fig. 23a) with a stil- 
etto-like process ; second joint with a finely dentate, pyramidal 
and pointed process (proc). 

Coloration: Rather opaque, a fleck of red pigment in the 
mouth; most of the pigment is found on the back and sides of 
the cephalothorax, and on the posterior maxillipeds. 

Length : Female, 4 mm. ; male, 3.5-4 mm. 

Occurrence : July 31, 1903, one male ; June 23, 1904, one male 
and one female. A good many (12-15) males were taken at one 
time on December 23, 1903, on the ' ' Banks. ' ' 

2. Euchaeta spinosa Giesbrecht. 

Euchaeta spinosa Giesbrecht, 1892, pp. 246, 263, pi. 16, figs. 12, 
26, 34, 47; pi. 37, figs. 31, 34, 35, 50; 1898, p. 39. 

5 Elevation in front of head (fig. 24a) produced anteriorly; 
genital segment almost symmetrical, with large, flap-like projec- 
tion at each side of the orifice (fig. 24&). Second terminal bristle 
of the furca longer than the other terminal bristles, dorsal bristle 
much thicker than the terminal ones. Anterior antennae reach 
beyond the end of the furca by more than the end joints. First 
outer marginal lobe of maxilla with eight bristles, second basal 
with three, fused second and third joints of inner ramus with 
four. Outer border of first joint of outer ramus of first foot 
concave ; outer border of third joint of outer ramus of second 
pair and its outer bristles different than in the following pairs; 
outer marginal bristle of second joint reaches to the end of the 
first marginal bristle of the third joint (fig. 24c). Basals and 
rami of posterior pairs of feet covered in places with short 
spines (fig. 24cZ). 

J Unknown. 

Coloration : Red in cephalothorax, sometimes on furca. 
plumose bristles of maxillipeds same color. Eggs blue. 

Length: Female, 6 mm. or over. 


University of California Publications. [Zoology 

Occurrence: July 21, 1903, one female: May 26, 1904, one 
female; July 5, 1904, four females; May 28, 1904, two females, 
one with egg cases, one without. 


Pig. 24. — Kurlnii in spinosa. Female, (a) Bead, lateral, Xlo. (M Abdo- 
men, Lateral, X48. Gen. seg., genital segment. (<■) Outer 
ramus of second foot • 45. (d) Basals, inner ramus, proximal 
joints of outer ramus of fourth foot X45. 

3. Euchaeta media Gtesbrecht. 

Kiirliui tn media Giesbrecht, 18S8, p. 337; 1892, pp. 246, 263, pi. 
16, figs. 13, 36; pi. 37, figs. 39, 40; 1898, p. 39. 

5 Elevation on front of head low I fig. 256) : genital 
segment asymmetrical with processes in the region of the orifice 
and a flap on the right side of the segment behind the orifice 
(fig's. 25c, d) ; furea (fig. 2og) as in E. acuta. Anterior antennae 
extend a little beyond the posterior border of the genital segment. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 


First outer marginal lobe of the maxilla (fig. 25/) with eight 
bristles, second basal with three, fused second and third joints 
of the inner ramus with four. Outer border of first joint of 
outer ramus of first foot concave ; outer border of third joint of 
second pair and its outer bristles different than in the following 

Ie.1 li.1 ri.2,3 

Fig. 25.— Euchaeta media. Female, (a) Lateral, X18. (b) Head, lateral, 
X83. (c) Genital segment, dorsal, X45. (d) Genital seg- 
ment, from right side X45. (e) Second foot X83. (f) Max- 
illa X83. Le.l, first lobe of outer margin. Li.l, first lobe of 
inner margin. Bi.2, 3, fused second and third joints of inner 
ramus, bristles not shown, (g) Furca, dorsal, X45. Si., inner 
marginal bristle. 

J Unknown. 

Coloration: Rather transparent; there is no pigment in the 
preserved specimens I have seen. 

Length: Females average about 3.3 mm. 

Occurrence: Forty or fifty females, many with eggs, were 
taken December 23, 1903, on the "Banks" with males of E. 

162 University of California Publications. [Zoology 

acuta. Three or four females were taken during June and July, 

The specimens which I have placed in this species correspond 
to Giesbrecht's descriptions; bul the females have hairs on the 
ventral sides of the abdominal segments, and Giesbrecht does 
nut mention these nor figure them (1892, pi. 37, figs. 39, 40). The 
outer marginal lobe of the maxilla is always provided with eight 
bristles, bul one of these is very inconspicuous and much shorter 
than the others. As these animals correspond wry closely in 
other respects to Griesbrechl 's specimens, especially in the form of 
the genital segment, 1 have thoughl besl to include them under 
his species, even though there are slighl differences. 

Sub-fam. Scolecithricin \i . 

Scolt cithru ina Giesbn cht, 1 892, ; 

$ Head commonly fused with first, and fourth with fifth 
thoracic segmenl ; rostrum with two usually sofl filaments; abdo- 
men .\ith four segments, symmetrical. Eighth and ninth joints 
of anterior antennae always fused, and occasionally other joints. 
Outer ramus of posterior antennae 6-jointed. Blade >*\' mandible 
with weak teeth; inner ramus of maxilla fused with second basal. 
The distal bristles of the anterior maxillipeds are modified into 
sac-like structures (fig. 30b), which occasionally arc pencillate 
at the end : lobes <>f appendages closely crowded together. Enner 
ramus of posterior maxillipeds a1 mosl only as Long as the second 
basal. Inner rami <>f swimming-feel jointed as in the Clauso- 
calanina< and se1 with spines; fifth fool rudimentary or absent. 

£ Abdomen with shortened anal segment, number of joints 
of anterior antennae reduced, the twentieth and twenty-firsl 
often fused only in one. Other head appendages like those of 
the female, nr specifically modified. Left fifth foot 5-jointed, 
occasionally with inner ramus, the righl 4-jointed (rudiment of 
inner ramus sometimes present I or lacking. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 163 
1. Genus Scolecithrix Brady. 

i'nclina (in part) Lubbock, 1856, p. 21. 

Scolecithrix Brady, 1883, p. 56. 

Scolecithrix Giesbrecht, p. 337; 1892, pp. 56, 265, 264; 1898, p. 4a. 

Lophothrix Giesbrecht, 1895, p. 254. 

Jmallophora (in part) Scott, T., 1893, p. 54. 

Neoscolecithrix Canu, 1896, p. 426. 

Scolecithrix Wolfenden, 1904, pp. 119, 120. 

Cephalothorax ellipsoidal, head fused with thorax, abdomen 
of female with four segments, of male with five; anal segment 
commonly short. Anterior antennae in female 19- to 24- jointed, 
in male 17- to 24- jointed, end joints (24 and 25) fused or dis- 
tinct, aesthetasks well developed, more numerous in male. Biting 
part of mandible and maxilla rather weak, inner ramus of max- 
illa mostly unsegmented and fused with the second basal. Distal 
bristles of anterior maxilliped thick, soft, in appearance some- 
thing like the aesthetasks of the antennae; these may be vermi- 
form, end in tufts (pencillate), or be pestle-shaped, and are 
usually present in both sexes. Head appendages of male like 
those of the female, but may in special cases be modified in par- 
ticular ways. Outer rami of first four feet 3-jointed, inner 
ramus of first foot 1-jointed, of second 2-jointed, of third and 
fourth 3-jointed; surfaces of both often set with spines and 
points. Fifth foot in female uniramous, 1- to 3-jointed, seldom 
absent ; fifth foot of male uniramous on each side, or the left 
biramous and the right uniramous, or both biramous. 


1. Head without crest 2 

1. Head with crest (fig. 28a) 4 

2. Anterior antennae of female 19-jointed; right of the male 17-, 

the left 18-jointed 3 

2. Anterior antennae of female 23-jointed S. subdentata 

2. Number of joints unknown; for characters compare description 

and fig. 30 S. pacifica 

3. First joint of outer ramus of first foot with a thorn-like bristle 

on outer margin (fig.' 26c) S. danae 

3. This joint without the bristle S. bradyi 

4. Anterior antennae of female 23-jointed; fifth foot (fig. 26e) ; 

right antenna of male 17-jointed, left 18-jointed; fifth foot 
(figs. 26c, d) S. persecans 


University of California Publications. [Zoology 

1. Scolecithrix danae Lubbock. 

Undina danae Lubbock, 1856, p. 21, pi. 4, figs. 6-9. 
Scolecitln i.r iliincic Brady, 1883, p. 57, pi. 17, figs. 1-12. 
Scolecithrix danae Giesbrecht, 1888, p. 333; 1892, pp. 265, 283, pi. 
pi. 13, figs. 4, 9, 14, 17: pi. 37, fig. 6; 1898, p. 42. 


Fig. 26.— Scolecithrix danae. (a) Female, lateral, X20. (b) Genital seg- 
ment, female, Lateral, XS3. (c) Outer ramus of first foot 
of female X140. <,/) Fifth foot of male X83. Ee.l dx., 
first joint of Outer ramus of tighl foot. .niter ramus of 

left foot., inner ramus of lefl foot. 

$ Fourth thoracic segment separate from fifth, latter with 
rather flat, rounded lateral angles. Third and fourth segments 
of the abdomen broader than long, genital segmenl with ventral. 
shovel-shaped process (fig. 266), anal segment short. Anterior 
antennae with nineteen segments, reaching beyond posterior 
border of the thorax but little. Outer ramus of posterior antennae 
9 7 as long as the inner ramus, seventh joint of outer ramus 
without proximal bristle. Second basal of maxilla with five, 
inner ramus with six, outer with five bristles (cf. fig. 29c). First 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 165 

basal of fourth pair without inner marginal bristle, first joint of 
outer ramus of first pair (fig. 26c) with outer marginal bristle. 
Fifth pair of feet absent. 

J Mouth parts not retrograded ; left fifth foot biramous, right 
uniramous, terminal joint very short (fig. 26d). 

Coloration : In formalin, both males and females have a light 
red or pink color. 

Length : Both sexes, 2-2.2 mm. 

Occurrence : June 28, 1904, one female ; December 29, 1903, 
one female, surface tow at 2 a.m. One male, October 20, 1904. 

2. Scolecithrix bradyi Giesbrecht. 

Scolecithrix bradyi Giesbrecht, 1888, p. 337; 1892, pp. 266, 283, 
pi. 4, fig. 7; pi. 13, figs. 1, 3, 7, 11, 21, 28; pi. 37, figs. 1, 
2, 9; 1898, p. 42. 

Fig. 27. —Scolecithrix bradyi. Female X31. 

$ Line of separation between fourth and fifth thoracic seg- 
ments visible only on the back; lateral portions of last thoracic 
segment elongated into two flaps, on the right more than on the 
left. Third and fourth segments of the abdomen much broader 
than long, genital segment asymmetrical, anal segment as long 
as the preceding ones, furca twice as long as broad. Anterior 
antennae 19-jointed, not reaching the posterior end of the thorax. 
Outer ramus of posterior antennae longer than the inner, seventh 
joint of the outer ramus without a proximal bristle. Maxilla as 
in S. danae, except that outer ramus has four bristles. First 
basal of fourth pair without bristle on inner margin, first joint 
of outer ramus of first pair without outer marginal bristle ; fifth 
foot very small. 

J Right anterior antennae with 18 joints, left with 17. Left 
fifth foot longer than the right by the last joint. Third joint of 
the outer ramus of the right large and with a prong. 

Coloration : Yellowish pigment in body, mouth region, and 
on feet. 


University of California Publications. [Zooloo- 

Length : Female, 1.4 mm. 

Occurrence: June 14, 1904. one female. 

3. Scolecithrix persecans Giesbrecht. 

Scolecith ans Giesbrecht, 1895, p. -53. pi. 3, figs. 6-12; 

1898, p. 48, fig. 9. 

Pig. 28.- Scolecithrix persecans. (a) Male, lateral, X9. (b) Second foot, 
male, ■ 15. (c) Lefl fifth fool of male X83. (<h Righl 
fifth foot, male. • 83. Be.3, third joint of outer ramus. 
(f) Fifth fool of female. 

J Head with rather high cresl (fig. 28a . Last two thoracic 
segments fused: left anterior antenna 18-jointed, right 17- 
jointed, reaching beyond cephalothorax. Outer ramus of pos- 
terior antennae at least 114 times as long as inner; second basal 
of maxilla with five, outer ramus with eight, inner with seven 
bristles, appendages of anterior oaaxilliped in part pencillate. 
First basal of fourth foot with feathered inner border: middle 
of outer border of first basal of second and third feet with a 
small tooth, outer border of second basal of second to fourth feet 
with a tooth i fig. 286) ; spines on outer margin of the two prox- 

Vol. 2] Esterhj. — Copepoda of the San Diego Region. 167 

imal joints of outer ramus of first foot shorter and more slender 
than on the third joint; terminal saw of outer ramus of third 
foot indented at base, inner ramus of foot with three spines on 
posterior surface of second and joints ; no spines on posterior 
surface of inner ramus of fourth foot. Anterior surface of outer 
ramus of second to fourth feet without spines, few on the anterior 
face of the inner ramus. Fifth foot fig. 28c, d. 

5 Anterior antennae 23-jointed, reaching 1 to end of furca; 
abdomen symmetrical, ventral surface of genital segment convex. 
Posterior antennae mandible, maxilla and maxilliped and swim- 
ming feet as in the male. Fifth foot symmetrical, rather well 
developed (fig. 28e). 

Coloration : Opaque white in formation, eye spots red. 

Length : Male, 5.3 mm. ; female, 4.6 mm. Giesbrecht gives the 
length of the male as 4.5 mm. 

Occurrence : Two males, one female collected at San Diego, 
May 31, 1904; obtained also May 18 and June 23, 1904. 

The female was not obtained by Giesbrecht, and has not since 
then been described, as far as I am aware. There can be little 
doubt that the outer ramus of the right fifth foot in the male 
is 3- jointed, and that the terminal joint in Giesbrecht 's single 
specimen was broken off. I have seen a considerable number of 
males, and in all the outer ramus is 3-jointed as shown (fig. 28rZ). 

4. Scolecithrix subdentata n. sp. 

5 Last two thoracic segments fused, each side with a small in- 
dentation in the lateral margin. Anterior antennae 23-jointed, 
not much longer than the cephalothorax. Inner ramus of the pos- 
terior antenna % as long as the outer; second basal of maxilla 
with four bristles, rami each with five (fig. 29c). Appendages 
of anterior maxilliped vermiform. First basal of fourth foot 
with a small, non-plumose bristle on inner margin; inner mar- 
ginal bristle of second basal of third and fourth pairs long and 
plumose; outer margin of first basal of first, second and third 
pairs with a small tooth in the middle, inner margin with promi- 
nent rounded process bearing the inner marginal bristle. First 
joint of outer ramus of first pair with outer marginal bristle. 
Fifth foot 2-jointed, leaf-like; terminal joint broad, oval, with 


University of California Publications. [Zoology 

a short distal spine on the outer border, and a longer proximal 
spine in the middle of the outer border (fig 1 . 29&). 
J Unknown. 

Fig. 29.— Scolecithrix subdentata, n. sp. (a) Female, Lateral, X31. (b) 
Fifth foot, female, • 195. (c) Maxilla X140. B.I, second 
basal. A''., inner ramus. /•''.. outer ramus. 

Approaches 8. dentata Giesbrechl in form of lasl thoracic 
segment, bu1 the indentation is qo1 as deep as in that species. 
The fifth fool is much as in dentata, bu1 more oval and rounded. 
Distinct from dentata in possessing an outer marginal bristle 
on the firsl joini of outer ramus of firsl foot, and in the number 
of joints of the anterior antennae. The bristles of the maxilla 
distinguish S. subdentata most sharply. 8. subdentata has the 
same number of joints in the antenna as S. longicornis Scott and 
8. auropt ctt n Giesbrecht. 

Length: Female, 1.48 mm. 

Occurrence: San Diego, May 31, June 14. June 23, 1904. 

5. Scolecithrix pacifica n. sp. 

5 Fourth and fifth thoracic segments fused, rounded later- 
ally. First segment of abdomen about as long as second and 
third together; the latter two are equal in length. Outer ramus 
of posterior antenna a little longer than the inner ramus. Sec- 
ond basal of maxilla with five bristles, inner ramus with eight, 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 169 

outer with five (fig. SOd). First basals of fourth feet without 
inner marginal bristle, inner border of second basal in second to 
fourth pairs ending in a sharp point. First joint of outer ramus 
of first foot with short, curved outer marginal bristle ; first joint 
of outer ramus of fourth pair without outer marginal bristle. 
Fifth foot (fig. 30c) 2-jointed, with a short distal bristle and a 
very long proximal one. 

Fig. £0. —Scolecithrix pacifica, n. sp. (a) Female, lateral, X31. (&) 
Anterior maxilliped X140. (c) Fifth foot X195. (d) Max- 
illa X83, parts as in fig. 29c. 

J Unknown. 

This specimen approaches S. porrecta closely in general char- 
acter, but is distinct in the length of the rami of the posterior 
antennae, form of the maxilla, bristle on outer margin of first 
joint of the outer ramus of the first foot, and in the form of 
the fifth feet. The anterior antennae are broken, but have prob- 
ably not over twenty joints. 

Length : Female, 2.3 mm. 

Occurrence: June 23, 1904, San Diego, one female. 


University of California Publications. 


6. Scolecithrix similis T. Scott. 

Amallophora dubia var. similis Scott, T., 1893, p. 56, pi. 4, figs. 

Scolecithrix .similis Giesbrecht, 1898, p. 46. 
S. similis ( .') Wolfenden, L904, p. 119, pi. 9, figs. •'. 6. 

Pig. 31.- Scolecithrix similis. (a) Male, lateral, <31. I b > Fifth foot, 

male, • \'<. 

J First abdominal segment short, second Long, twice the 
length of the third, which is shorter than the fourth. Right 
anterior antenna 18-jointed, lefl 23-jointed (Scott). Lasl two 
thoracic segments fused. First basal of fourth fool with a 
plumose inner marginal bristle; both feel of fifth pair biramous 
(fig. 316). 

Length : Male, 2.6 nun. 

Occurrence: San Diego, June 23, 1904. 

The antennae of the single specimen were broken, bu1 the 
form of the abdomen and fifth feel warranl one in identifying 
it with Scott 's species, a1 least provisionally. 

I tropagidai Giesbrecht, L892, p. 58; 1898, p. 52. 
J Head always distinct from thorax; rust ruin with two, usu- 
ally soft, filaments, sometimes plumose. Anterior antennae as 
in the Calanidae, but the second joint is more often divided into 
two parts, never less than twenty-three joints. Outer ramus of 
posterior antennae at least % as long as the inner. The suc- 
ceeding four pairs of appendages as in Calanus and like forms. 
In the three anterior ones are found peculiarities (in the Hete- 

Vol. 2] Esterly. — Copcpoda of the San Diego Region. 171 

rorhabdinae) : stunting of the second and third inner marginal 
lobes of the maxilla through lengthening of outer ramus; pre- 
ponderance of distal bristles of anterior maxilliped over the prox- 
imal. The four anterior pairs of feet with 3-jointed rami ; 
but in Temora the number of joints is reduced through fusion. 
The fifth pair of feet is like the others (inner marginal bristle 
of second joint of outer ramus of special form, sword-shaped, 
awl- or thorn- like) or rudimentary, inner ramus 1-jointed or 
lacking, outer ramus 1- to 3-jointed. 

J Abdomen with five segments, anal segment rarely short- 
ened; genital orifice and grasping antenna on opposite sides of 
the body. Grasping antenna right or left, joints 19 to 21, and 22 
to 23 fused. Both feet of fifth pair present, inner rami complete 
or reduced to absence; outer rami forming hooks or forceps. 
Slight sexual differences occasionally in form of last thoracic 
segment and swimming feet. 

Sub-fam. Centropaginae. 
Centropagina Giesbreeht, 1892, p. 59. 

9 Cephalothorax with six segments, abdomen with three ; ros- 
tral filaments soft. Anterior antennae (24th and 25th joints 
fused), mandibles and maxilla as in Calanus; the length of the 
distal curved bristles of the anterior maxillipeds and the heavily 
bristled first basal of the posterior maxillipeds is characteristic. 
All five pairs of feet with 3-jointed rami. 

J Grasping antenna on the right side; outer ramus of left 
fifth foot 2- jointed; the right foot with forceps. 

1. Genus Centropages Kroyer. 

Centropages Kroyer, 1849, p. 602. 

Catopia Dana, 1848, p. 25; 1849, p. 280; 1852, p. 1172. 

Hemicalanus DaEa, 1852, p. 1103. 

Ichthyophorba Lilljeborg, 1853, p. 184. 

Diaptomus Lubbock, 1857, p. 403. 

Ichthyophorba Clans, 1863, p. 198. 

Centropages Brady, 1883, p. 81. 

Centropages Giesbreeht, 1892, pp. 59, 303, 731; 1898, p. 53. 

Centropages Wheeler, 1899, p. 172. 

Centropages Thompson and Scott, 1903, p. 247, pi. 1, figs. 19-25. 

5 Head separate from thorax, fourth thoracic segment from 
fifth. Abdomen with three segments, genital segment asymmet- 

172 University of California Publications. [Zoology 

rical. Anterior antennae 24-jointed; joints 24 and 25 fused. 
Outer ramus of posterior antennae 7-jointed and almost l 1 /* 
limes as long as the inner ramus. The distal bristles of the ante- 
rim- maxillipeds sickle-shaped, with spinous feathering, and much 
longer and thicker than the proximal bristles. First basal of the 
posterior maxillipeds with strongly protruding lobes, both the 
middle ones set with bristles, which have ;i spinous feathering; 
inner ramus well developed, 5-jointed. Rami of swimming feet 
usually 3-jointed, bul the inner ramus is exceptionally 2-jointed. 
First basal with bristle on inner border in first to fourth feet, 
second basal thus equipped in the first pair* First basal of fifth 
pair without inner marginal bristle; inner marginal hristle of 
second joint of outer ramus thorn-like and fused with the joinl 
i fig. 32. . 

£ Sexual peculiarities in the form of the abdomen, righl 
anterior antenna, and fifth pair of feet. The abdomen is com- 
posed of five segments; anal seg nt in mosl species very shorl ; 

genital opening on the Left. Righl anterior antenna a grasping 
organ. Joints 19-2] and 22-23 fused, geniculation between the 
18th and 19th. Inner marginal bristles lacking on outer ramus 
of lefl fifth foot, joints 2 and •'> fused. The outer ramus of the 
right foot 3-jointed, both distal joints metamorphosed into ;i for- 
ceps, one blade of which is the terminal joint, while the other is 
the thickened inner marginal bristle of the second joint. 

1. Centropages bradyi Wheeler. 

oliii'i iis Brady, lss:;, p. s.",, pi. .",7. figs. 111. 
Centropages bradyi Wheeler, L899, p. 17 1. fig. 12. 

$ Second joint of outer ramus of fifth fool with a stoul 
smooth spine (fig. 32c). Sides of inflated genital segmenl with- 
out spines or knob-shaped projections. Furca symmetrical, with 
a peculiar short, truncated, peg-shaped projection (fig. 326) 
between insertions of the two outer bristles. Wheeler, 1899). 

J Joint 17 of right anterior antenna with smooth anterior 
border, not seriate: joints 19 and 20 fused, separated from joint 
21 ; joint 18 with accessory series of teeth on lower surface 
(Wheeler, 1899). 

Vol. 2] Estcrly.—Copepoda of the San Diego Region. 173 

Coloration : Opaque, with a large purplish spot in middle of 

Length: Female, thorax, 1.6 mm.; abdomen ? 
Occurrence: June 10, 1904, one female. 

Fig. 32.— Centropages bradyi. (a) Female, thorax, lateral, X45. (6) 
Abdomen, ventral, after Wheeler 1899. (c) Fifth foot 

Wheeler, 1899, p. 174, does not mention the spine-like pro- 
tuberance on the dorsal surface of the first segment of the ceph- 
alothorax in the female, but since the other characters as given 
by him (especially the furca) agree with the San Diego speci- 
men, I have not made a new species of the latter. This agrees 
in possessing the dorsal spine, with C. dorsispwatus (Thompson, 
1903, p. 247, pi. 1, figs. 19-25), but differs widely in other 

Sub-fam. Temorinae. 
Temorina Giesbrecht, 1892, p. 60. 

5 Cephalothorax with five segments; fourth and fifth thor- 
acic segments fused; rostral filaments soft, sometimes plumose. 
Anterior antennae 23- or 24-jointed; the second joint is either 
not divided, or, if it is divided into two parts, the proximal 
portion is fused with the first joint. Outer ramus of posterior 
antennae 7-jointed, and, with the following four appendages, is 

174 University of California Publications. [Zoology 

like those of the Calanidae. The first four pairs of feet usually 
with 3-jointed rami, in which, however, the two proximal joints 
may be fused: inner ramus absent in fifth pair, or small and 
1-jointed; the outer ramus is 1- to 3-jointed. 

J Grasping antenna usually the right; distally from the 
geniculation, the nineteenth and twenty-first and twenty-second 
and twenty-third joints are fused: sexual peculiarities often in 
the swimming feet as well as in the form of the body, anterior 
antennae and fifth pair of feet. 

1. (onus Pleuromamma Giesbrecht. 

Diaptomus Lubbock, 1856, p. 27. 

/'// uromma < 'laus, 1S63, p. 195. 

/'/< wromma Brady, 1883, p. 45. 

I'l, ilium urn Giesbrecht, 1892, pp. 61, . -, >47, 757. 

Ph wromma Dahl, L893, p. 105. 

Pleuromma Wheeler, L899, p. L76. 

I'h iiinmiimmii Cirslircrht. 1898, p. b's. 

Easily recognizable by a dark-pigmented knob on the right 
or left side of the first thoracic segmenl I figs. 33a, 34a I. Furca 
at mosl twice as long as broad. Rami of the firsl to fourth pairs 
of feel 3-jointed, firsl joinl of outer ramus of third pair with a 
deep Dotch in the outer border; terminal bristle of outer ramus 
of third pair shorl and benl outward; first joinl of inner ramus 
of second pair with hooks on inner border, on righl and lefl font 
in the female, usually on one side in male. Fifth pair in female 
rudimentary, '2- to 4-jointed, in male 5-jointed on each sidi . 
without forceps. Grasping antenna of male on right or left 
side. Abdomen of female with three segments; of male with 
five, sometimes asymmetrical. 

1. Pleuromamma abdominalis Lubbock. 

Diaptomus abdominale Lubbock, 1856, p. 28, pi. 10, figs. 1-8. 
Pleuromma abdominale Claus, 1863, p. 197, pi. 5, figs. 1-6, 13, 

14; pi. 6, fig. 1-10. 
Pleuromma abdominale Brady, 1883, p. 46, pi. 11, figs. 1-13. 
Pleuromma abdominale Giesbrecht, 1892, pp. 347, 357, pi. 5, fig. 

8; pi. 32, figs. 3, 5, 13, 22, 25-30; pi. 33, figs. 43, 44, 48, 

49, 52. 
Pleuromamma abdominalis Giesbrecht, 1898, p. 109. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 175 

5 Pigment knob on right or left side ; proximal joint of first 
antenna with several smaller and two larger (one straight and 
one curved) teeth on anterior border. Fifth pair of feet 4- 
jointed, with three apical bristles (fig. 336). 

Fig. 33.— Pleuromam ma abdominalis and P. gracilis, (a) P.a., female, 
lateral, X31. (6) P.a., fifth foot X45. (c) P.g., fifth foot 

J Pigment knob, genital opening and hooks on inner border 
of first joint of inner ramus of second foot, on left side. Proxi- 
mal joint of anterior antennae with small teeth only, grasping 
antenna on right side. Abdomen symmetrical. End joint of 
left fifth foot broadened. 

Coloration: Transparent, except for a small amount of red 
in the mouth region. 

Length: Female, about 2.4-3 mm.; male, 3.3 mm. 

Occurrence : A common species, both summer and winter, but 
males are very infrequently found; one male was taken June 2, 

2. Pleuromamma gracilis Clans. 

Pleuromma gracile Glaus, 1863, p. 197, pi. 5, figs. 7-11. 
Pleuromma abdominale (in part) Brady, 1883, p. 47, pi. 2, figs. 

1-16; pi. 21, figs. 13, 14. 
Pleuromma gracile Giesbrecht, 1892, pp. 347, 357, pi. 5, fig. 7; 

pi. 32, figs. 6, 18-20; pi. 33, fig. 41-47. 
Pleuromamma gracilis Giesbrecht, 1898, p. 110. 

5 Pigment knob on right side; anterior border of anterior 
antennae with only small teeth ; fifth pair of feet 2-jointed, three 
prongs at the end (fig. 33c). 


University of California Publications. [Zoology 

J Pigment knob on right side; abdomen symmetrical-, ante- 
rior antennae as in female as regards armature; grasping 
antenna on left side; first joint of inner ramus of second foot 
with hooks only on right side ; third and fourth feet as in female. 

Coloration: As in P. abdominal is. 

Length : Both sexes, 1-2 mm. 

Occurrence: More abundant than P. abdominalis, but males 
are rare. 

Brady, 1883, p. 47, considers that P. gracilis Claus is an 
immature form of P. abdominalis Claus, but Giesbrecht, 1892, 
does not favor this view, and the San Diego specimens of P. gra- 
cilis present such differences when compared with P. abdominalis 
that there can be no doubt of the distinctness of the species. The 
forms represented by /'. gracilis are without doubt mature, since 
females are often found with attached spermatophores. 

3. Pleuromamma xiphias Giesbrecht. 

Pleuromma xiphias Giesbrecht, 1889, p. 6; 1892, pp. 347, 367, 
pi. 32, fig. 14; pi. 33, figs. 42, 45, 50. 

Plewomamma xiphias Giesbrecht, 1898, p. 110. 

Fig. 34.— Pleuromamma xiphias. (a) Female, lateral, X18. (b) Head 
of female, lateral, X48. (c) Second basal, and proximal 
joints of rami of second foot, X48. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 177 

Allied to P. abdominalis, but the front of the head anterior 
to the rostrum is prolonged into a prominent process (fig. 346). 

Coloration : As in the other species. 

Length: Female, 4.1-4.5 mm. 

Occurrence: San Diego, July 31, 1903, one female; June 23, 
1904, one female; taken also December 23, 1903, on "Banks." 
The occurrence of the male is uncertain ; large male animals have 
been taken Avith the characteristic shape of xiphias, but I cannot 
say definitely whether they belong to this species or not. 

2. Genus Metridia Boeck. 

Metridia Boeck, 1864, p. 13. 

Paracalanus Brady and Bobertson, 1878, p. 126. 

Metridia Giesbrecht, 1892, pp. 61, 339, 749; 1897, p. 254; 1898, 

p. 105. 
Metridia Dalai, 1894a, p. 10. 
Metridia Wheeler, 1899, p. 175. 
Metridia Wolfenden, 1904, p. 125. 
(See also T. Scott, 1S93, p. 42, pi. 3, figs. 8-20.) 

Closely allied to Pleuromamma, but is without the lateral pig- 
ment knob. Terminal bristle of outer ramus of third pair of 
normal form; swimming feet of the male (especially the second 
pair) corresponding with those of the female. Furca 2 to 5 times 
as long as broad. 

1. Metridia lucens Boeck. 

Metridia lucens Boeck, 1864, p. 14. 

Paracalanus hibernicus Brady and Robertson, 1873, p. 126. pi. 8, 

figs. 1-3. 
Metridia armata Brady, 1878, p. 42. 
Metridia Mbernica Giesbrecht, 1892, pp. 340, 357, pi. 33, figs. 2, 

12, 16, 22, 28, 36, 39. 
Metridia lucens Dahl, 1894, p. 11. 
Metridia lucens Giesbrecht, 1898, p. 106. 

5 Cephalothorax 1% times as long as the abdomen ; lateral 
angles of fifth thoracic segment slightly pointed. Genital seg- 
ment somewhat shorter than the two last abdominal segments 
together, the anal segment about % as long as the preceding. 
Furca shorter than the last abdominal segment and twice as 
long as broad. The anterior antennae reach back hardly to 
posterior margin of the genital segment. Terminal bristle of 
end joint of outer ramus of fourth foot little over 14 as long 


University of California Publications. [Zoology 

as the joint. Fifth foot 3-jointed and with three rather long 
bristles on end joint (fig. 35c i. 

Pig. 35.- Metridia lucens and M. boeckii. (a) .1/./.. female, lateral, X20. 
(b) M.L. second basal and lirst joint of inner ramus of second 
fool of female to shovi hooks, X195. (c) .1/./.. fifth foot 
of female X195. (d) M.b„ fifth foot of female X195. 

J Grasping antenna on righl side. Fifth foot: second joint 
of outer ramus of lefl fool without, firsl joinl of outer ramus 
of righl fool with a long, thorn-like bristle. 

Length : Female, '■>.- mm. 

Occurrence: Very common, summer and winter. 

2. Metridia boeckii Giesbrecht. 

Metridia boeckii Giesbrecht, L889, p. 5; 1892, pp. 340, 3-46, pi. 33, 
figs. 8, 19, 31, 37; 189S, p. in?. 

$ Like .1/. lucens, but furca is ;is long as the fifth abdominal 
segment, and twice as long as broad. Anterior antennae reach 
a little beyond the posterior border of the thorax. Fifth foot 
with four joints < tit;-. 35<2). 

J Unknown. 

Length : Female, 2.5 mm. 

Occurrence : One or two in catches with M. lucens. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 179 

It should be noted that not a male specimen of Metridia has 
been taken in any catch, so far as I have examined them, and 
rather particular attention has been paid to this point. 

Sub-fam. Lucicutiinae. 
Leuckartiina Giesbrecht, 1892, p. 62. 

5 Cephalothorax with five segments, fourth and fifth thoracic 
segments fused, rostral filaments thin and usually soft ; abdomen 
with four segments, symmetrical. The second joint of the ante- 
rior antenna is divided, and the twenty-fourth joint is separate 
from the twenty-fifth. Outer ramus of posterior antennae 8- 
jointed, the four following appendages like those in the Cala- 
nidae. The first four pairs of feet almost always with 3-jointed 
rami, the fifth pair like the preceding ones and that of Centro- 
pages, with 3-jointed outer ramus and 2- to 3-jointed inner 

J (Known only in Lucicutia.) Grasping antenna the left; 
distal to the geniculation the nineteenth to twenty-first, and 
twenty-second and twenty-third joints are fused; fifth pair of 
feet without forceps, basals 2-jointed, the right with 2-, the left 
with 3-jointed rami ; no other sexual differences except in form 
of body. 

1. Genus Lucicutia Giesbrecht. 

Leuclartia Claus, 1863, p. 182. 

Leuckartia (in part) Brady, 1883, p. 50. 

Leuclartia Giesbrecht, 1892, pp. 62, 358; 1895, p. 258. 

Lucicutia Giesbrecht, 1898, p. 110. 

Lucicutia Steuer, 1904, p. 596. 

Lucicutia Wolfenden, 1904, p. 121. 

Head broad ; furca symmetrical. First lobe on outer border 
of maxilla with five bristles. 

$ Five segments in cephalothorax, abdomen with four, sym- 
metrical. Rostral filaments slender, situated on a papilla. Pos- 
terior antennae like those of Centropages, but with eight joints 
in outer ramus. Blade of mandible weakly built, outer ramus 
bent rather far proximally. Outer border lobes of maxilla with 
only five bristles ; inner border lobes well developed, the proximal 
one, however, with weak masticatory bristles. Inner ramus 2- 
jointed, articulated with basal; outer ramus large, oval. Distal 


University of California Publications. [Zoolog 

bristles of maxillipeds little longer than the proximal; bristles 
of outer border of posterior maxilliped slender, without hairs. 
Outer rami of the five pairs of feet 3-jointed ; inner ramus of first 
pair 2-jointed (second and third joints fused), of second to fifth 
pairs 3-jointed; first basal with bristle on inner margin in second 
to fourth pairs, second basal in first pair with inner marginal 
bristle, and sometimes with a lulu-like process. The bristle on 
the inner margin of the second joint of tbe outer ramus of the 
fifth pair has the form of ;i curved awl I fig. 366). 

£ Sexual peculiarities in the form of tbe abdomen, posterior 
antennae and fifth pairs of feel. Abdomen with five segments, 
genital opening on righl si.le. The Lefl ;m1enna is a grasping 
organ, geniculating between joints I s and 19; joints 19 to 21. 
22 ami i':: fused. Fifth pair of feet with 2-jointed basals; rami 
of left foot 2-jointed, of righl 3-jointed. Tbe distal joint of the 
outer ramus of tbe righl foot is booked and may be moved toward 
the proximal joints. 

1. Lucicutia flavicornis Claus. 

Leuckartia flavicornis clans. L863, p. l v :'.. pi. 32, fig. 17. 
Leuckartia flavicornis Giesbrecht, ls'.ci, p. 358, pi. 5, fig. 4; pi. 

in. figs. 2, 3, 15, 17. 21, 23, 29, 38; pi. 38, fig. 38, 40. 
Lucicutia /<<< iesbrecht, 1898, p. 111. 

Fig. 36.— Lucicutia flavicornis. (a) Female, lateral, X18. (b) Fifth 
foot, female. XS3. Ri., inner ramus, (c) Outer margin of 
miter ramus of third foot X140. 

$ Anal segment shorter than the preceding; second terminal 
bristle of furca thick, twice as long as abdomen. The anterior an- 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 181 

tennae reach beyond middle of the furca, joint 19 as long as tenth 
to twelfth, inclusive. Second basal of maxilla with four bristles. 
Inner ramus of first pair of feet 3-jointed, with eight bristles ; 
inner ramus of fifth pair reaches almost to the distal border of 
the second joint of the outer ramus; first joint of outer ramus 
much shorter than the third, which is twice as long as the term- 
inal bristle. 

J Terminal portion of grasping antenna (joints 19-25) some- 
what longer than joints 14-18. Inner ramus of right foot of fifth 
pair straight, with five bristles, which are at the end of the 
terminal joint. 

Coloration: Transparent, with light yellowish pigment in 
various locations. The San Diego specimens showed this to a 
very small extent. 

Length: Female, 1.6 mm. ; male a little less. 

Occurrence : June 8, 1904, one male, one female ; June 10, 
1904, one male (?). 

Sub-fam. Heterorhabdinae. 
Heterocliaetina Giesbrecht, 1892, p. 63. 

Cephalothorax with five segments; fourth fused with fifth 
thoracic segment ; rostral filaments slender, sometimes plumose ; 
last thoracic segment in some cases with pointed lateral angles. 
Abdomen with three or four segments, not always symmetrical. 
Second joint of anterior antennae divided, the two terminal 
joints usually distinct. Second joint of outer ramus of posterior 
antennae divided into two, so that there are as a result eight 
joints in the ramus (which, however, may be reduced by fusions). 
Blade of mandible with few teeth, inner ramus small, sometimes 
lacking. Inner ramus and both distal lobes of inner margin of 
maxilla small, occasionally absent; outer ramus always present, 
and usually much lengthened. Anterior maxilliped elongate, 
lobes small, the proximal ones usually rudimentary ; bristles of 
distal lobes, and usually those of the inner ramus, almost always 
long, thick and hooked. The four anterior pairs of feet with 
3-jointed rami, fifth pair like the others, rami almost without 
exception 3-jointed. 

J Sexual differences in form of body, anterior antennae, fifth 
pair of feet, seldom in structure of mouth parts. Grasping an- 

182 University of California Publications. [Zoolouy 

tenna usually the left ; first and second joints fused, as well as 
the nineteenth to twenty-first, twenty-second and twenty-third 
(or twenty-second to twenty-fifth) ; fifth pair of feet with 3- 
jointed outer and 1- to 3-jointed inner ramus; forceps incomplete 
or absent. 

1. Genus Heterorhabdus Giesbrecht. 

Heterochaeta Claus, l v ti:;. p. 180. 

II, t, rochai ta Brady, 1883, p. 48. 

Heterochaeta Giesbrecht, 1892, pp. 64, 372, 745; 1895, p. 259. 

Heterochaeta, Aurivillius, 1899, p. 38, figs. 4, 5. 

II, i, rorhabdus Giesbrecht, 1898, p. 113. 

Heterorhabdus Wolfenden, 1904, p. 124. 

$ Cephalothorax with five segments, rostra] filaments soft, 
situated on a papilla. Abdomen with four segments, Left half of 
I'niva not articulating with anal segment, Larger than the right 
and with much longer bristles. Rami of posterior antennae about 
equal in length, outer ramus with eighl bristles. Anterior maxil- 
liped straight, terminal portion and proximal lobes with their 
bristles strongly suppressed, while the distal lobes are provided 
with strong hooked bristles. Posterior maxUlipeds distinguished 
by shortness of the bristles on the inner ramus and by the Length 
and thickness of one bristle on the inner margin of the firsl basal 
joint. All the feet have 3-jointed rami: inner marginal bristle 
of the firsl basal in pairs one to four, and of the distal basal joint 
in the firsl pair, well-developed and plumose. Terminal joint 
of outer ramus of third pair usually different in form from that 
joint in the other pairs, being broad and oval (fig. 38d). The 
inner marginal bristle of the second joint of the outer ramus of 
the fifth pair is thickened and sword-shaped I fig. 38e). 

J Sexual peculiarities in form of posterior portion of body, 
left anterior antenna and fifth pair of feet. Abdomen with five 
segments, genital opening on right side. Grasping antenna 
slender. Fifth pair of feet with 3-jointed rami; terminal joints 
of both outer rami hooked: processes on the distal basal joint. 


Abdomen 4-segmented, fifth foot symmetrical $ 

Abdomen 5-segmented, fifth foot asymmetrical J 

•1. Third joint of outer ramus of third foot of same form as in 

second and fourth pairs (fig. 40a) H. longicornis 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 


1. This joint in third pair broad and oval, terminal bristle short- 

ened (fig. 38d) 2 

2. First basal of posterior maxilliped with a very long, heavy- 

bristle in the middle of the inner border; rostral papilla with 

a point (fig. 37a) H. spinifrons 

2. As above, but rostral papilla without point (figs. 38b, c) 3 

3.$ Inner marginal bristle of second joint of inner ramus of fifth 
foot much shorter and more slender than those of the third 
joint; first joint of outer ramus with thorn-like inner marginal 

bristle. J Fifth foot (fig. 39) H. clausi 

3.$ Inner marginal of second joint of inner ramus of fifth foot but 
little shorter than those of third joint; first joint of outer 
ramus without inner marginal bristles H. papilliger 

1. Heterorhabdus spinifrons Clans. 

Heterochacta spinifrons Claus, 1863, p. 182, pi. 32, figs. 8-9, 

14, 16. 
Heterochaela spinifrons Giesbrecht, 1892, pp. 372, 382, pi. 10, 

figs. 1, 3, 11, 16, 19, 31; pi. 39, figs. 42, 43, 51, 52, 54. 
Heterorhabdus spinifrons Giesbrecht, 1898, p. 114. 

b.1 ... 

Fig. 37.— Heterorhabdus spinifrons. Female. («) Head, lateral, X83. (b) 
Posterior maxilliped X167. B.I, first basal. 

The papilla on front of head ends in a sharp point ; anterior 
antennae reach beyond the end of the fnrca by the last four or 
five joints. The fourth lobe of the anterior maxilliped has two long, 
thick bristles, and a small, slender one which is hardly y± as long 
as the other two; the fifth lobe has two bristles, one of which 
is longer and thicker than the other. A spine-like bristle at the 
end of the inner margin of the first basal of the posterior maxil- 
liped is 14 the length of the long bristle in the middle of the 
margin (fig. 37&). Hooks at the end of both outer rami of the 


University of California Publications. [Zoology 

fifth foot of the male relatively longer than in H. papilliger, the 
left over twice as long as the first and second joints of the outer 

Coloration; Transparent and colorless. 

Length : Female, 3.4 mm. 

Occurrence : June 23, 1904, one female. 

2. Heterorhabdus papilliger clans. 

Heterochacla papilligt ra Claus, 1863, p, 182, pi. 3, figs. 10-13, 15. 
Heterochaeta papilligera Giesbrecht, 1892, pp. 372, 382, pi. 20, 

figs. 4, 7, 10, 15, 17, 23, 35, 36; pi. 39, figs. 40, 53. 
Heterorhabdus papilliger Giesbrecht, 1898, p. 114. 


Fig. 38. — Heterorhabdus papilliger. (a) Female, lateral, X31. (6) Head, 
female, lateral, X83. (c) Head, female, dorsal, X83. (d) 
Outer ramus of third foot of male X83. (e) Fifth foot of 
female X140. Ee.2, second joint of outer ramus. (f) 
Eight mandibular blade of male X83. (g) Anterior maxil- 
liped, female, X83. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 185 

Papilla on front of head elongated bnt not ending in a point 
(figs. 38b, c). Anterior antennae when at the sides of the body 
reach about to the end of the furca. Fourth lobe of the anterior 
maxillipeds (fig. 38g) with two long, thick bristles and a shorter, 
thinner one, which is over half as long as the others; fifth lobe 
with two bristles of about equal length and thickness. On the end 
of the inner border of the first basal of the posterior maxilliped 
is a spine-like bristle, which is hardly one-eighth as long as the 
bristle in the middle of the margin, and the latter one is almost 
twice as long as the second basal (cf. fig. 376). Hooks at the 
end of both outer rami of the fifth foot in the male relatively 
shorter than in spinifrons ; the left little longer than the first 
and second joints of the outer ramus together. 

Coloration : As in spinifrons. 

Length : Female, 2.2 mm. ; males slightly smaller. 

Occurrence : A few specimens of both sexes were taken during 
May and June, 1904. 

3. Heterorhabdus clausi Giesbrecht. 

Eeterochacta clausii Giesbrecht, 1889, p. 2 ; 1892, pp. 372, 382, pi. 

20, fig. 2, 28, 37, 38. 
Heterorhabdus clausi Giesbrecht, 1898, p. 115. 

Pig. 39. — Heterorhabdus clausi. Fifth foot, male, X83. 


University of California Publications. [Zoology 

Like papiUi<i< r, but the anterior antennae reach somewhat 
beyond the end of the fnrea. Inner ramus of anterior maxilli- 
peds with longer bristles; inner marginal bristle of first and 
second joints of inner ramus of fifth foot in the female short and 
slender, that of the first joint of the outer ramus thick and 
booked; second basal of the right fifth loot in the male with a 
Long Lamellar process, the second joint of the outer ramus with 
a shorter projection on the inner border, third relatively longer. 
especially on the Lefl side. 

Length: Male, 2-2.5 mm. 

Occurrence: San Diego. July 22, 1903, one male; June 23, 
1904. one male. 

4. Heterorhabdus longicornis Griesbrecht. 

Hi h mi Inn in longicornis Giesbrecht, 1889, p. --, 1892, pp. :'>73, 383, 
pi. 20, figs. II. 21, -•". 26; pi. 39, fig. 44. 

Heterorhabdus longicornis (Jiesbnvht, lsits, p. 116. 

Heterorhabdus zetesios Wolfenden, L902, p. 367. 

Heterorhabdus longicornis (male) Wolfenden, 1904, p. 124, pi. 
9, fig. 34. 

Fig. 40. — Heterorhabdus longicornis. Male, (a) Third foot X140. (6) 
Right fifth foot X140. (c) Left fifth foot X140. 

5 Anterior antennae reach beyond the end of the furca for 
the last eight or nine joints; inner ramus of maxilla with five 
bristles, first and second inner marginal lobes relatively long; 
anterior maxillipeds with a greater number of bristles, but with 

Vol. 2] Esterhj. — Copepoda of the San Diego Region. 187 

less strongly developed hooked bristles than in the other species ; 
inner ramus clearly with three joints, and with seven long 
bristles; bristles of first basal of the posterior maxillipeds and 
third joint of outer ramus of third swimming foot of usual form. 
Inner marginal bristle of second joint of outer ramus of fifth 
pair more slender, and inner marginals of first and second joints 
thicker than in the other species, distal border of second joint of 
outer ramus of ordinary form. 

J Like female in structure of maxillipeds and terminal joint 
of outer ramus of third and fourth swimming feet. Fifth foot 
(figs. 406, c) : right with stiff upright process on second basal 
(inner margin), covered with stiff spines, second joint of outer 
ramus with a projection having four teeth at end. 

Length: Male, 3 mm. 

Occurrence : San Diego, June 23, 1904, one male. 

2. Genus Augaptilus. 

Hemicalanus (in part) Clans, 18C3, p. 176. 

A-ugaptilus Giesbrecht, 1889, p. 3; 1892, pp. 65, 400, 724; 1898, 

p. 120. 
(See also T. Scott, 1893, p. 36, pi. 2, figs. 25-37; Steuer, 1904, 

p. 597.) 

$ Cephalothorax composed of five segments; rostral filaments 
short and sometimes feathered. Abdomen with 3 segments, geni- 
tal segment usually not wholly symmetrical. Anterior antennae 
25-jointed, outer ramus of posterior antennae rarely longer than 
the inner ramus. Mandibular blade with two teeth (mandible 
sometimes uniramous and with a stylet-like blade). Inner ramus 
of maxilla lacking; both maxillipeds with reduced proximal lobes 
and peculiarly equipped bristles (cf. fig. 41&). Feet with spines 
on outer border of outer rami reduced in part, the third bristle 
on the inner border of the last joint of the outer ramus of the 
fifth foot not elongated, the inner marginal bristle of the middle 
joint awl-shaped or lacking; both rami 3-, rarely 2-jointed. 

J Sexual differences in the form of the abdomen, anterior 
antennae and fifth feet. Abdomen with 5 segments, genital open- 
ing right or left. The right or left anterior antenna may be the 
grasping organ. Rami of both feet of fifth pair 3-jointed ( fig. 
41c, d). 


University of California Publications. [Zoology 

1 Augaptilus longicaudatus Claus. 

//' micalanus longicaudatus Claus, 1863, p. 179, pi. 29, fig. 3. 
Augaptilus longicaudatus Giesbrecht, 1S92, p. 414, pi. 27, fig. 31; 

pi. 28, figs. 1], 19, 23, 31, 35, 38; pi. 2, fig. 22; pi. 39, figs. 

37, 48; 1898, p. 123. 
Augaptilus longicaudatus Scott, 1894, p. 34, pi. 1, figs. 24-26; pi. 

2, fig. 5. 

Fie. 4}. 

Augaptilus longicaudatus. Male. (a) Dorsal X18. (b) An- 
terior maxilliped XS3. (<•) Right fifth foot X83. (d) 
Left fifth foot X83. 

5 Genital segment not entirely symmetrical, twice as long as 
both the following segments together; furca as long as the anal 
segment, and about 5 times as long as broad. Anterior antennae 
longer than trunk by about the last 6 joints. Inner ramus of 
posterior antennae ~% longer than the outer ramus; first and sec- 
ond joints of outer ramus not fused; mandible uniramous. 
Anterior maxilliped: First and second lobes lacking, third with 
1 bristle, fourth and fifth with 2, sixth with 1. First basal of 
posterior maxilliped with 0, 0, 1, 2 bristles. Length of first 
and second basals and inner ramus as 7 :6 :5. Outer ramus of 
tifth foot 2-jointed. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 189 

J Grasping antenna on left. Fifth foot, fig. 41c. <7. 
Coloration : Transparent, without pigment. 
Length: Male, 3.39 mm. 
Occurrence: June 10. 1904, 1 male. 

3. Genus Arietellus Giesbrecht. 

Arietellus Giesbrecht, 1892, pp. 6(3, 415. 
Hhincalanus (part) T. Scott, 1893, p. 31. 
Arietellus Giesbrecht, 189S, p. 124. 

Last two thoracic segments fused, elongated into a strong 
spine on each side (fig. 42a), front with wedge-shaped process, 
rostral filaments slender. Abdomen of female with 4 segments, 
symmetrical ; furca, and appendages with long, richly plumose 
bristles. Anterior antennae of female and the right one of male 
at most 20-jointecl, joints 1 and 2, 21-25 fused; grasping 
antenna on the left, 19- jointed, terminal portion 2-jointed. 
Inner ramus of posterior antenna straight, longer than outer ; 
mandible uniramous, inner ramus lacking; inner ramus and 
third inner marginal lobe of maxilla lacking, outer ramus long 
and characteristic. Anterior and posterior maxillipeds as in 
Augaptilus except in appendages of bristles (fig. 42c). Rami 
of first to fourth feet 3-jointed; fifth foot of female (fig. 426) 
3-jointed, basals 2-, outer ramus 1-jointed, inner ramus rudi- 
mentary. Fifth foot of male without forceps, basals 2-, outer 
ramus 3-jointed, inner ramus 1-jointed. 

1. Arietellus setosus Giesbrecht. 

Arietellus setosus Giesbrecht, 1892, p. 415, pi. 29, figs. 1, 3-7, 
9-13; pi. 39, figs. 34-36; 1897, p. 254; 1898, p. 124. 

With the characters of the genus. 

Coloration : Terminal expansions of plumose f ureal bristles 
red, the remaining portion black. Body orange red, bristles on 
posterior antennae and mouth parts, deep red. 

Length : 5.5 mm. 

Occurrence : One female was taken at San Diego, Dec. 
22, 1903. 


University of California Publications. [Zoology 

Fig. 42.— Arietellus setosus. Female, («) Dorsal X9. (b) Fifth foot 
X83. (c) Dist.-il portion of one of the bristles of the an- 
terior maxillipod > 1 I". 

4. Genus Phyllopus Brady. 

Phyllopus Brady, 1883, p. 78. 

Phyllopus Giesbreeht, L892, pp. 66, 419; 1898, p. 124. 

Phyllopus Wolfenden, 1904, p. 124. 
$ Last thoracic segment not entirely symmetrical ; abdomen 
with 4 segments, genital segment asymmetrical. Anterior 
antennae with 24 joints. Inner ramus of posterior antennae 
about half as long as the outer ramus of 8 joints. Blade of 
mandible strong, with four teeth. Anterior maxillipeds elon- 
gated, posterior with short, broad first basal. First to fourth 
pairs of feet with 3-jointed rami, second basal with inner mar- 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 191 

ginal bristle in first pair and with outer marginal in first and 
fourth. Fifth pair with basal of two joints and 3-jointed outer 
ramus. Inner ramus lacking, inner marginal bristle of middle 
joint of outer ramus thick and long; terminal joint shortened, 
its distal margin toothed (fig. 43&). 

J Like female except in structure of anterior antennae and 
fifth feet. Abdomen with 5 segments. Left anterior antenna 
20-jointed, geniculating between joints 17 and 18. Fifth feet 
each with 2 basals, and 3-jointed outer rami, the right foot has 
a rudimentary inner ramus, broad and without spines. Second 
basal of each foot with a long, slender plumose bristle. 

1. Phyllopus bidentatus Brady. 

Phyllopus bidentatus Brady, 1883, p. 78, pi. 5, figs. 7-16. 
Phyllopus bidentatus Giesbrecht, 1892, p. 419, pi. 18, figs. 25-33; 

pi. 38, fig. 35; 1898, p. 124. 
Phyllopus bidentatus $ Wolfenden, 1904, p. 124, pi. 9, fig. 16. 

Fig. 43.— Phyllopus bidentatus. Female, (a) Lateral X18. (b) Fifth foot 

With the generic characters. Both Giesbrecht (1892) and 
Wolfenden (1904) state that the "bidentate" lateral portion of 
the last thoracic segment does not exist as in Brady's descrip- 
tion. The San Diego specimen agrees with the description of 
the two former authors. The male of the species is described 
by Wolfenden as cited, and the above description is taken 
from him. 

Coloration : Transparent, without pigment. 

192 University of California Publications. [Zoology 

Length : Female, 2.2 mm. 

Occurrence: San Diego, May 31, 1904, one female. 


Candacidae Giesbrecht, 1892, p. 67. 
Candaciidae Giesbrecht, 1898, p. 126. 

5 Cephalothorax with 5 segments, rostrum absent, abdomen 
with 3 segments. In posterior antennae the second basal and first 
joint of inner ramus is fused, outer ramus slender, end joints 
shortened. Blade of mandible with few teeth. Second lobe of 
inner margin of maxilla very long, third and fourth absent. 
Anterior maxilliped without lobes, bristles on distal portions 
sickle-shaped ;in<l hooked. Posterior maxilliped as in Calanus 
1 n it small and weak. Inner ramus of anterior pairs of feel 2- 
jointed ; fifth pair rudimentary. 

J Genital orifin lefl ; grasping antenna the left, seven- 
teenth and eighteenth and nineteenth and twentieth joints fused; 
fifth foot without inner ramus, the left 4-jointed, the right 3- 
jointed ending in a forceps or bristle. 

1. Genus Candacia Dana. 

Candacia I (ana, I - 16, p. 184. 
Ifionyx Kroyer, 1848-49, p. 601. 
Candaa Dana, L849, p. 279; L852, p. 1109. 
Conduct Lubbock, L856, p. 29. 
Candace < Slaus, L863, p. 189. 
Candaa Streets, L877, p. 139. 
Candace Brady, 1883, p. 66 
Candaa Thompson, 1888b, p. 148. 
Candaa Giesbrecht, 1892, pp. 67, 423, 729. 
Candaa Wheeler, 1S99, p. 177. 
acta Giesbrecht, 1898, p. 126. 

5 Fourth and fifth thoracic segments fused: front of head 
rectangular, lateral angles of last thoracic segment pointed: 
abdomen with 3 segments, genital segment often asymmetrical. 
Anterior antennae 23- or 21-jointed, proximal segments thick- 
ened, anterior border toothed. Rami of posterior antennae 
short, outer ramus slender, second joint elongated, terminal 
ones very short. Basal of mandible large, rami short, blade 
with 2 teeth. Anterior maxilliped elongate, without lobes, dis- 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 193 

tal bristles strong, sickle-shaped ; posterior maxilliped small and 
weak, second basal and inner ramus suppressed. First to fourth 
pairs of feet with 3-jointed outer rami, inner rami relatively 
small, 2-jointed ; first basal with inner marginal bristle in second 
and third pairs. Outer border of outer ramus toothed, fifth 
pair stunted, 3-jointed on each side. 

J Last thoracic segment more often asymmetrical, the point 
on the right side noticeable for form, size, and color. Abdomen 
with 5 segments, genital segment often asymmetrical with out- 
growths on the right side. Right anterior antenna a grasping 
organ. Fifth foot on right side 3-jointed, on left side 4-jointed ; 
the right foot terminates in a forceps or bristle. 


1. Terminal bristle of outer ramus of third foot with outward bent 
point, at least as long as the distance between the distal spines 
of the outer border of the joint (fig. 44c) 2 

1. This bristle much shorter than the designated portion of the outer 

border ( fig. 4.7 d) 

2. Genital segment of female longer than broad C. pectinata 

2. Genital segment of female broader than long, male not known. . 

C. bipinnata 

3. The thick proximal portion of the anterior antennae is 7-jointed 

(fig. 45b) 4 

3. This portion 6-jointed 5 

4. Terminal joint of fifth foot of female without bristles on inner 

border; joint of grasping antenna proximal to geniculation 
with deep teeth on anterior border. Fifth foot of male (fig. 

4(j c ) C. curta 

4. Terminal joint of fifth foot of female with three bristles, apical 
teeth slender and sharp. Teeth on grasping antenna fine; 
genital segment with flat outgrowth on right side; (fig. 47&). . 

C. aethiopica 

1. Candacia pectinata Brady. 

Candace pectinata Brady, 1878, p. 49; 1883, p. 67, pi. 30, 

figs. 1-13. 
Candace pectinata Giesbrecht, 1892, pp. 424, 439, pi. 4, fig. 3; pi. 

21, figs. 2, 12; pi. 22, figs. 9, 17, 18, 31, 43-46; pi. 39, figs. 

1, 21, 22, 24, 25. 
Candacia pectinata Giesbrecht, 1898, p. 128. 
Candace pectinata Wheeler, 1899, p. 177, fig. 15. 

Genital and following segment in female asymmetrical, the 

latter protruding posteriorly ; last thoracic segment in the male 


University of. California Publications. [Zoology 

asymmetrical. Anterior antennae with 23 joints, pectinate part 
of joints of grasping antenna deeply toothed, the segments on 
either side of the articulation suppressed. Proximal hooked 
bristles of second basal of anterior maxilliped as thick and 
almost as long as the distal ones. Terminal joint of fifth foot 
of female long and claw-like (fig. 446), without inner marginal 
bristle ; right fifth foot of male with forceps. 

Fig. 44.— Candactfl pectinata. (a) Las! thoracic segment and first and 
second segments of abdomen, male, X45. (6) Fifth foot of 
female X83. (c) Third joint of outer ramus of third foot 
of male X49. (d) Fourth foot of female X83. 

Coloration: Rather transparent, lateral prolongations of last 
thoracic segments, genital orifice, rami and bristles of feet and 
mouth parts, joints 18 and II) of grasping antenna, usually a 
blackish brown. 

Length: Females average 2 mm., males 1.9 mm. 

Occurrence: Rather common; both sexes are taken in sum- 
mer and winter. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 195 

2. Candacia bipinnata Giesbrecht. 

Candacc bipinnata Giesbrecht, 1889, p. 5; 1892, pp. 424, 439; 

pi. 22, fig. 20; pi. 39, figs. 27, 29. 
Candacia bipinnata Giesbrecht, 1898, p. 129. 

Fig. 45.— Candacia bipinnata. (a) Cephalothorax and genital segment 
of female, dorsal, X31. (&) First eight joints of anterior 
antennae of female X83. (c) Last thoracic segment and 
abdomen of female X20. 

5 Like C. pectinata, but genital segment (fig. 45a) is broad 
and has a wing-like expansion on each side. J Unknown. 

Coloration : Much as in C. pectinata. 

Length : Female, 2.6 mm. 

Occurrence: Taken usually with C. pectinata, but in fewer 


University of California Publications. [Zoology 

3. Candacia curta Dana. 

C, ni.lnr, curt, i Dana, 1849, p. 279; 1852, p. 1116; 1855, pi. 78, 

figs. 6 a-<l. 
Candaci curta Giesbrecht, 1892, pp. 42-4, 439, pi. 21, fig. 15; pi. 

22, fi^rs. 12, 24; pi. 39, figs. 8-10, 12. 
Cumin, -'in carta Giesbrecht, 1898, p. 129. 

Fig. 46.— Candacia curta. Mai.'. < a \ Last thoracic segment, ami genital 
segment, lateral, X83. (b) Same, dorsal, X31. (c) Fifth 
foot X83. Ri<:lit fool at ri-l.t of figure. 

Allied to C. pectinata, but right side of genital segment in 
female has a ventral projection; fifth fool of female with two 
heavy teeth on the end, and one oil the inner border. Proximal 
joint of inner ramus of first foot with bu1 two inner marginal 

Coloration: As in preceding species, with very slight varia- 

Length: .Male. 1.5 mm. Giesbrecht gives 2.4-2.65 nun. 

Occurrence: San Diego, Jan. 4, 1904, one male. 

4. Candacia aethiopica Dana. 

Candace ethiopica Dana, 1848, p. 23. 

Candace melanopus clans. 1863, p. 191, pi. 33. 

Candace ethiopica GiesbrecM, 1892, pp. 424, 439, pi. 4, fig. 13, 

pi. 21, figs. 1, 9; pi. 22, figs. 1, 6, 13, 14, 32, 40-42; pi. 39, 

figs. 7, 11, 13. 
Candacia aethiopica (Hesbreeht, 1898, p. 128. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 


Genital segment of female slightly asymmetrical, with a 
process on the left side ; last thoracic segment of male asymmet- 
rical. Anterior antennae 23-jointed, denticulation of the geni- 
culating joints of grasping antenna fine, joints proximal and 
distal to the geniculation long and slender. Proximal hooked 
bristle of the second basal of the anterior maxilliped as thick 
and almost as long as the distal (fig. 47/;). Terminal joint of 
fifth foot of female with one tooth on the outer margin, 3 apical 
teeth, and 3 bristles on the inner border ; right fifth foot of male 
with a forceps (fig. 47#). 

Fig. 47. — Candacia aethiopica. (a) Female, dorsal, X165. (b) Last 
thoracic segment, and genital segment of male, dorsal, X60. 
(c) Abdomen of female, lateral, X37. , (d) Second and third 
joints of outer ramus of third foot, female, X60. (e) 
Fifth foot of female X130. (f) Left fifth foot of male 
X130. (g) Forceps of right fifth foot of male X130. (7i) 
Anterior maxilliped of female X130. 

Coloration : Dorsal surface of cephalothorax, excepting an- 
terior portion of head and the last thoracic segment, black brown, 

198 University of California Publications. [Zoology 

distinguishing the species a1 once. Appendages colored about as 
in the other forms. 

Length : Female 2.9 mm, male 2-2.5 mm. 

Occurrence: Several males and females were taken October 
20, 1904. 

Pontellidae Giesbrecht, 1892, p. 68; 1898, \>. 131. 

$ Head and thorax distinct, fourth thoracic segment usually 
fused with fifth. Rostrum forked, usually ending in two very 
strong prongs; rarely absent. Eyes Large, sometimes with one 
or two paifs of cuticular lenses and one unpaired lens. Anterior 
antennae 16- to 24-jointed, the two terminal joints always fused. 
Second basal and first joinl of inner ramus fused, terminal joints 
of outer ramus shortened. Mandible od the whole as in the 
Centropagidae. First basal of maxilla large, second basal and 
rami relatively small. Anterior maxillipeds as in the Centro- 
pagidae, long, hooked bristles on distal portion and commonly 
mii tic proximal. First basal id' posterior maxillipeds large with 
Long bristles on Lobed inner border, second basal and inner ramus 
relatively small. Inner ramus of four anterior pairs of feet or 
second to fourth, 2-jointed; fifth pair rudimentary, outer ramus 
1-jointed (rarely 2-jointed), inner ramus 1-jointed or lacking. 

J Distinct from female in form of abdomen, anterior 
antennae and fifth pair of feet, at times also in form of eyes, 

rostrum and last thoracic segment. Genital orific i left side, 

grasping antenna on right, middle joints much or slightly 
broadened: joints 1!) and 21 and 22 to 2.1 fused. Fifth pair of 
feet rarely with rudimenl of inner ramus; forceps of right foot 
incomplete or very powerful. 

Sub-fam. Ponteliinae. 
Pontellina Giesbrecht, 1892, p. (58. 
5 Cephalothorax with five or six segments: last thoracic seg- 
ment ends in one (seldom two) sharp points on each side and is 
at times asymmetrical. Rostrum ends in two strong chitinous 
prongs or in two filaments. One pair of cuticular lenses is 
occasionally found on the dorsal side, seldom two pairs, ventral 

Vol. 2] Esterly. — Copepocla of the San Diego Region. 199 

eye strongly protruding (fig. 48c). Abdomen with from one 
to three segments, never symmetrical. Anterior antennae 16- 
to 24-jointed, at least two terminal joints fused, usually also a 
number of proximal joints. Posterior antennae with reduced 
number of terminal joints in outer ramus which is often more 
slender and thinner than the inner ramus. Mandible as a 
whole as in Centropages, blade with at least five teeth. Maxilla 
with relatively large proximal basal, second inner marginal lobe 
large, second basal, rami and first outer marginal lobe accord- 
ingly relatively smaller. Anterior maxilliped as in Centropages, 
with very strong hooked bristles; posterior maxilliped short, 
first basal with indented or folded inner margin, set with long, 
strong bristles ; inner ramus 3- to 5-jointed, bristles short. Outer 
ramus of four anterior feet 3-jointed, inner ramus of second 
to fourth pairs or of all 2-jointed. Basal of fifth pair 2-jointed, 
inner and outer rami usually 1-jointed; outer ramus seldom 

J Sexual peculiarities in form of body, more often in eyes, 
anterior antennae and fifth foot. Last thoracic segment as a 
rule asymmetrical, right posterior angle more strongly de- 
veloped ; abdomen with 5 segments, in cases with asymmetrical 
processes on right side. Right anterior antenna with broadened 
middle joints ; beyond the geniculation either the nineteenth 
and twenty-first joints only are fused (besides the twenty-fourth 
and twenty-fifth) or also the twenty -second and twenty-third; 
fifth foot without inner ramus (perhaps a rudiment on left 
foot), four jointed on each side, the right foot with forceps. 

1. Genus Labidocera Lubbock. 

Pontella (part) Dana, 1846, p. 184; 1848, p. 26; 1849, p. 280. 

Pontellina (in part) Dana, 1852, p. 1135. 

Labidocera (snb-genus) Lubbock, 1853a, p. 25; 1853&, p. 202. 

Pontella Clans, 1863, p. 207; 1893, p. 233. 

Pontella Brady,, 1878, p. 73; 1883, p. 87. 

Pontella Thompson, 1887, p. 34. 

Labidocera Giesbrecht, 1889, p. 7; 1892, pp. 70, 444, 746; 1897, 

p. 254; 1898, p. 132. 
Labidocera T. Scott, 1893, p. 82. 
Labidocera Wheeler, 1899, p. 178. 

200 University of California Publications. [Zoology 

Head usually without hooks on side ; one pair of dorsal eye 
lenses, larger in the male than in the female; rostral hooks 
strongly chitinized. Cephalothorax of 5 segments, ending in 
points laterally, more strongly developed on right side in the 
male Abdomen of female with 2 or 3 segments, of male 
with •">, sometimes asymmetrical in the female. Anterior 
antennae of Pemale 23-jointed; terminal section of grasping 
antenna (the right) of male 4-jointed. Mandibular blade with 
5-7 hooked, pointed teeth. Second basal of maxilla bent toward 
outside, about twice as Long as the second lobe of the inner 
margin: anterior maxilliped stunted and provided with strong 
hooked bristles especially on the distal half; posterior maxilliped 
with 4-jointed innei ramus. Inner ramus of swimming feet 
2-jointed, outer ramus 3-jointed; fifth font of female on each 
side with 2-jointed basal portion, rami 1-jointed, though the 
innei ramus may be rudimentary. Fifth fool of male 4-jointed 
on each side, righl withoul inner ramus, left a1 times with nidi 
mentary inner ramus. The two terminal joints of the right foot 
form a powerful forceps 

1. Labidocera trispinosa a.sp. 

5 Cephalothorax symmetrical, evenly rounded in front; 
crest, and hooks on side of head absent; rostrum bifid, very 
long; last thoracic segment produced on each side into sharp 
points (cf. fig. 48a. left side). Abdomen with :> segments, 
genital Longer than the last two, asymmetrical, with a prominent 
blunt wing-like process on the righl side (fig. 48d) ; middle 
abdominal segment with a knob-like projection on the left in 
front. Purca symmetrical, about 3 times as long as broad. 
Anterior antennae extending back to the posterior border of 
fourth thoracic segment. Fifth pair of feet symmetrical (fig. 
4%), outer ramus ending in two teeth, the inner one twice 
as long as the outer: inner ramus about one-half as long as the 
outer, articulating with basal: outer ramus Longer than the first 
and second basals together. 

J Eye lenses larger, and more nearly contiguous (fig. 48a). 
Last thoracic segment on right side with a long slender spine, 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 201 

curving dorsally, (fig. 486), a shorter straight dorsal spine, 
and a very short spine directed toward median line. On the left 
side the thorax is about as in the female. Abdomen with five 
segments, genital segment in some cases slightly asymmetrical; 
orifice on right side ; middle segment as long as the first two, 
and longer than the last two. Anterior antennae reaching to 
base of furca; joints 16 and 17 about of equal length; teeth 
on joint 18 directed toward distal end of antenna, those on the 
next joint larger and straight (fig. -iSli). Fifth foot, fig. 48e, f. 

Fig. 48.— Labidoeera trispinosa, n. sp. (a) Male, dorsal, X18. (&) Lat- 
eral portion of last thoracic segment from right side X60. 
(c) Head of male, lateral, X60. (d) Abdomen of female, 
dorsal, X60. (e) Left fifth foot of male, (f) Right fifth 
foot of male, (g) Fifth foot of female. (/*) Joints 17, 18, 
19 to 21, 22, of grasping antenna of male, X160. 

202 University of California Publications. [Zoology 

This species is distinct from any of the seventeen described 
species of Labidocera in the form of the last thoracic segment 
of the male, and the genital segment of the female. It approaches 
L. lubbocki Giesbrecht and L. bruncscens Giesbrecht, more 
closely than any others, but differs distinctly from them in 
the above-named features as well as in the structure of the fifth 
feet of the sexes, etc. 

Coloration: Rather transparent, intestinal contents light 
green, thorax and abdomen yellowish with given tinge in places. 

Length: Female, 1.6 mm. Male, 1.7-2.2 mm. 

Occurrence: .May 24. 1904. one female. .June 16, 1904. four 
males, five females. 

Sub-fam. Parapontellinae. 
Parapontellina Giesbrecht, L892, p. 7:'.. 

Cephalothorax usually with five, seldom with six segments, 
rostral filaments slender or Lacking, last thoracic segmenl with 
rounded or pointed sides. Eyes without dorsal chitin lenses. 

5 Abdomen usually with three, seldom two segments, at 
times asymmetrical. Anterior antennae 17- to 19-jointed; sev- 
eral proximal joints fused in addition to the terminal points. 
Second basal of mandible elongate, usually cylindrical, blade 
narrow with from five to seven teeth. Maxilla elongate, lobes 
slightly protruding and not articulating; rami more often 
stunted; the entire second basal may be absent. Distal honked 
bristles of anterior maxillipeds Long and strong, seldom so on 
proximal portion of the appendage. Posterior maxilliped as in 
the I'oiiti lliiiio bui inner ramus is only 1- or 2-jointed. Outer 
ramus of anterior four pairs of feet 3-jointed, inner ramus of 
second to fourth or of all 2-jointed. Fifth pair stunted, basal 
1- or 2-jointed, outer ramus 1-jointed usually claw-like and with- 
out inner ramus ; not always symmetrical. 

J Sexual peculiarities in form of body, anterior antennae 
and fifth foot. Last thoracic segment and abdomen at times 
asymmetrical as in many Pontellinae; abdomen with five seg- 
ments. The right grasping antenna simulated in some respects 
in the left; the right antenna with but little broadened middle 
joints and differences in the segmentation of the joints in the 

Vol. 2] Esterhj. — Copepoda of the San Diego Region. 203 

proximal portion ; joints nineteen and twenty-one and twenty- 
two and twenty-five fused. Right fifth foot 3- to 4-jointed, 
forceps not complete, the left 3-jointed, seldom with rudiment of 
inner ramus. 

1. Genus Acartia Dana. 

Acartia Dana, 1846, p. 183; 1852, p. 118. 

Bias Brady, 18 S3, p. 72. 

Bias Lilljeborg, 1853. 

Bias Glaus, 1863, p. 191. 

Bias Brady, 1883, p. 72. 

Acartia Thompson, 1888a, p. 149; 1888ft, p. 141. 

Acartia Giesbrecht, 1892, pp. 75, 506, 721; 1898, p. 150. 

Acartia Dahl, 1894c, p. 13. 

Acartia Wheeler, 1899, p. 182. 

Fifth thoracic segment and abdomen of male symmetrical; 
latter with shortened anal segment. Antennae of female with 
seventeen segments, of the same diameter throughout the length ; 
grasping antenna of male with very slightly thickened middle 
joints. Outer ramus of posterior antenna much shorter than 
the inner; second joint of inner elongated, first joint with nine 
bristles on the inner border. Outer ramus of mandible articu- 
lates in the middle of the margin of the second basal. First 
outer marginal lobe of maxilla with long bristles, outer ramus 
rudimentary, its place supplied by two bristles. Proximal lobes 
of anterior maxillipeds well developed, with long bristles; pos- 
terior maxilliped with four joints. Inner ramus of first pair of 
feet with two joints, fifth pair of female without inner ramus, 
with long outer terminal bristle on second basal. 

5 Cephalothorax with five segments, last two fused. Pos- 
terior antenna very slender, second basal joint fused with prox- 
imal joint of inner ramus, mandible with weak blade, which 
has seven teeth. Posterior maxilliped without outer marginal 
bristles, and with three inner marginals on third joint. Outer 
ramus of first to fourth pairs of feet 3-jointed, first basal 
without bristles, second with rather long outer marginal bristle 
in fourth pair. The very much stunted fifth pair (fig. 49cZ) 
consists on each side of two or three joints; the end joint (outer 
ramus) is a thick stylet-shaped bristle, and on the outer border 
of the second basal is a slender feathered bristle. 


University of California Publications. 


J Sexual differences in form of body, anterior antennae and 
fifth pair of feet. Abdomen with five segments, genital orifice 
on left side; the fourth segment and furca shortened. The 
right anterior antenna is a grasping organ and joints 19-21 and 
22-25 are fused. The fifth pair of feet (fig. 49c) consists of a 
common middle pari and ;i right foot of four joints, a left of 
three, each uniramous. The right especially has the form of a 
claw, which, because of ,-i process od the joint preceding the 
terminal, becomes ;in incomplete forceps. 

1. Acartia tonsa Dana. 

Acartia tonsn I tana, L848, p. 26. 

Acartia tonsa Giesbrecht, 1892, pp. 508, 522; \>\. 30, figs. 7, 24, 
::»; pi. 43, figs. 6, L0; L898, p. L54. 

Fig. 49. — Acartia tonsa. cm Female, dorsal, X45. (b) Abdomen of male, 
dorsal. X55. (c) Fifth tout of male X138. Dx., right foot. 
(d) Fifth foot of female X138. 

Rostral filaments present, last thoracic segment rounded 
laterally: abdomen of male (fig. 49b) with spines on the second 
segment; anal segment with lateral hairs. Anterior antennae of 
female without thorns, not reaching to the posterior border of 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 205 

the genital segment. Middle joint of fifth foot of female about 
as broad as long (fig. 49<i) ; terminal portion of foot as long 
as the rest of the appendage, straight, toothed posterior to 
middle ; plumose bristle as long as terriiinal claw. Second joint 
of right fifth foot of male (fig. 49c) without process on inner 
margin; process of third and fourth joints broader than in 
A. clausi. 

Coloration: Very transparent, without pigment. 

Length : Female, 1.2-1.5 mm. Male, 1-1.1 mm. 

Occurrence : Enormous quantities may be obtained in Glor- 
ietta Bight, San Diego Bay, especially at night; the species 
occurs rather infrequently outside. 

II Sub-order!— PODOPLEA. 
Body divided into an anterior and posterior portion but 
the line of separation falls in front of the last thoracic seg- 
ment (fig. 50a) ; the posterior portion of the body has as the 
first segment the fifth thoracic segment, which bears almost 
without exception a rudimentary pair of feet (figs. 50a, 57b) ; 
these are never of service to the male in pairing. The sper- 
matophores are placed directly upon the genital orifice of the 
female, without the use of a pair of appendages. The genital 
organs of the male are usually paired, their orifices always 
symmetrically placed. The female carries the eggs until the 
young are set free. 


(The genera marked with an asterisk have been found in the 
San Diego region.) 

1. Inner rami of third and fourth feet 3-jointed, or lacking in 

fourth pair 2 

1. Inner rami of third and fourth feet 1-jointed Mormonilla 

2. Inner ramus of first foot 1-jointed 3 

2. Inner ramus of first foot 2-jointed (fig. 53c) 14 

3. Head with two large chitinous lenses (figs. 62, 57b) 4 

3. Head without chitinous lenses 7 

4. Inuer ramus of fourth foot 2- or 3-jointed (fig. 59b) 5 

4. Inner ramus of fourth foot 1-jointed or replaced by bristles 

(fig. 63b) 6 

5. Abdomen with four or five segments which are broadened later- 

ally (figs. 57a, 59a) *Sapphirina 

206 University of California Publications. [Zoology 

5. Abdomen with two segments, not broadened Corina 

6. Eye lenses separated by at least their diameter; the last two 

thoracic segments without lateral sharp prolongations Copilia^ 

6. Eye lenses placed close together, last two segments of the 

anterior portion of the body prolonged into lateral pointed 
processes (figs. 61a, 62b) *Corycaeus 

7. Entire body much flattened, furca very long and stylet -like .. CopiliaJ 

7. Body of various shapes but more rounded; if at times depressed, 

never leaf like 8 

8. Outer ramus of first foot 1-jointed, postern lateral angles of 

fourth segment of body prolonged into processes ( ii^ r . 54a).... 

8. Outer ramus of firsl fool 2- to 3-jointed 9 

9. Outer ramus of posterior antenna I -jointed; furca very short, 

each ramus with a very long bristle twice as long as the body at 
least; rami of furca and the two bristles fused in the median 

line; remaining furca] bristles stunted Aegisthus 

St. Outer rami's of posterior antenna 3 -jointed; fuTCS short, rami 
separate ( fig. 52c), each with a long bristle, at leasl us long 
the body, and at least twice us long as the other bristles 
(fig. 52a ) *Microsetella 

9. Outer ramus of posterior antenna lacking; furca longer than 

broad, rami separate HI 

10. Anterior and posterior maxillipeda alike in structure, both with 

long, spinous bristles *Oithona 

10. Posterior maxilliped with tew or no short bristles and a terminal 

hook (Oncaeidae) ( li^s. 556, 566) 11 

11. Fifth pair of feet l-jointed, with two lancet-shaped appendages 

at the end which have dentate borders; body elongate. .Lubbockia 

11. Fifth pair 1- or 2 jointed or knob-like, with naked or plumose 

bristles; body more robuSl 12 

12. Anterior antennae with very long and thick aesthetasks on the 

terminal joints; fifth feet 2-jointed Ratania 

ll'. Anterior antennae with numerous pencillate aesthetasks on the 

proximal joints; fifth foot a protuberance Pachysoma 

ll'. Anterior antennae with few and very delicate aesthetasks; fifth 

foot reduced to a small rod or knob, or at times to one 

bristle 13 

13. Terminal hooked bristles on the posterior antenna of medium 

length; inner ramus of rear feet at least as long as the outer, 
terminal joint in fourth pair at least l 1 - times as long as the 
first and second joints together *Oncaea 

13. Hooked bristles on the much elongated terminal joint of posterior 

antennae very long; inner ramus of rear feet shorter than 

outer, its terminal joint in the fourth pair no longer than each 

of the proximal joints Conaea 

14. Front of head with two great chitinous lenses Miracia 

14. Head without lenses 15 

15. Forehead conical, rounded in front; body very narrow; outer 

ramus of posterior antennae lacking Setella 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 207 

15. Forehead pointed (fig. 53a) ; body broad; outer ramus of pos- 

terior antenna 1-jointed 17 

16. Furca with separate rami (about twice as long as broad) and 

bristles much shorter than body (fig. 53a) *Euterpe 

16. Kami of the fuca very short and with the two unusually long 

bristles fused in the median line Aegisthus 


1. Genus Oithona Baird. 

Oithona Baird, 1843. 

Scribella Dana, 1847, p. 279; 1848, p. 19. 

Oithona Dana, 1852, p. 1097. 

Oithona Claus, 1863, p. 104. 

Oithona Brady, 1883, p. 97. 

Oithona Giesbrecht, 1892, pp. 77, 537, 753; 1896, p. 324. 

Oithona Wheeler, 1899, p. 186. 

5 Anterior and posterior parts of body composed of five 
segments, first and second abdominal segments fused (fig. 50a). 
Genital opening lateral. Anterior antennae rather obscurely 
jointed, bristles long; posterior antennae 3-jointed, outer ramus 
absent. Inner ramus of mandible small, 1-jointed, outer ramus 
4-jointed; blade dentate. Rami of maxilla. 1-jointed, inner 
ramus small. Maxillipeds slender, bristles strong, spinous; 
inner ramus of posterior maxilliped 2-jointed. Rami of swim- 
ming feet 3-jointed. Fifth pair very rudimentary, being reduced 
to two bristles on each side. 

J Front of head blunt (fig. 51a) ; first and second abdominal 
segments not fused (fig. 506, 51a), bristles of furca short. 
Anterior antenna are grasping organs, geniculating at two 
places. Swimming feet somewhat irregular in number and 
arrangement of bristles. 

1. Oithona plumifera Baird. 

Oithona plumifera Baird, 1843. 

Oithona plumifera Dana, 1852, p. 1099, pi. 76, figs. 4a-e. 

Scribella scriba Dana, 1849, p. 279. 

Oithona spinirostris Claus, 1863, p. 105. 

Oithona plumifera Giesbrecht, 1891, p. 475; 1892, pp. 537, 548; 

pi. 4, fig. 10; pi. 34, figs. 12, 13, 22, 25, 27-29, 32, 33, 

44-47; pi. 44, figs. 1, 7, 12-15. 
Oithona plumifera Wheeler, 1899, p. 186, fig. 22. 

5 Front ending in a somewhat ventrally directed, pointed 
beak, but visible in dorsal view. Furca shorter than anal seg- 


University of California Publications. [Zoology 

merit, about three times as long as broad, outer marginal bristle 
about three times as long as the furea. Anterior antennae 
extend to the posterior border of the fourth abdominal seg- 
ment. Second basal of mandible with two hooked bristles ; inner 
ramus of maxilla with a minute bristle. Outer ramus of first 
pair of feet (fig. 50cZ) with one outer marginal bristle on the 
first joint, one on the second and two on the third; outer ramus 
of the second and third pairs, with one on the first joint, none on 
the second and two on the third; of the fourth pair with none 


Fig. 50.— Oithona plumifera. (a) Female, dorsal, X40. Th.5, fifth 
thoracic segment, (b) Abdomen of male X140. (c) Outer 
ramus of third foot of male X83. (d) Outer ramus of first 
foot of female X265. Se., outer marginal bristles, St.. 
terminal bristle. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 


on the first and second joints, two on the third ; proximal bristle 
of outer margin of third joint of third and fourth pairs reduced. 

J Genital segment broad (fig/ 506). Proximal joint of distal 
portion of anterior antennae with a half crescentic process on 
the inner margin. Third joint of outer ramus of first and 
fourth pairs of feet with two outer marginal bristles, the second 
and third with three (fig. 50c). 

Coloration : Giesbrecbt shows red pigment in body, and 
especially in long bristles of anterior antennae, furca, feet and 
mouth parts, while other animals may be colorless. All speci- 
mens I have seen are colorless. 

Length : Female, 1-1.4 mm. ; male, 0.75-1 mm. 

Occurrence: Not as abundant as 0. nana, but some speci- 
mens occur in all catches where the ordinary Podoplea are num- 
erous, summer and winter. 

2. Oithona nana Giesbrecht. 

Oithona nana Giesbrecht, 1892, p. 549, pi. 4, fig. 8; pi. 34, figs. 
10, 11, 20, 24, 26, 34, 35, 42; pi. 44, figs. 2, 4. 

Fig. 51.— Oithona nana, (a) Male, dorsal, X83. (6) Third joint of outer 
ramus of first foot of female X83. (c) Third joint of outer 
ramus of fourth foot of male X83. 

210 University of California Publications. [Zoology 

5 Front blunt ; furca as long as the anal segment, hardly 
twice as Long as broad, outer marginal bristle about as long as 
the furca. Anterior antennae reach about to the posterior 
margin of the third thoracic segment. Second basal of mandible 
with one booked bristle. Inner ramus of maxilla with four 
bristles. First, second and third I fig. 516 ; . joints of outer ramus 
of first to third feet respectively, with one. one. three outer 
marginal bristles, of the fourth with one. one. two. 

J Division line between the firsl and second thoracic seg- 
ments with a sharp median projection; genital segmenl nar- 
rower than in <). plumifera. Proximal joint of the distal por- 
tion of the anterior antennae without the round process. Third 
joint of outer ramus of tirst to third feel with three outer mar- 
ginal bristles, of the fourth with two i fig. 51< , 

Coloration: Transparent, without pigment. 

Length: Female, 0.7-0.8 nun.: male. 0.5-0.6. 

Occurrence: Rather abundanl in hauls taken from inside of 

the kelp beds ;it l'oint I ;i. Both sexes found, 'flic tow in 

which Oithona occurs most plentifully contains scarcely any 
other genera than Oncaea, Euterpt and Corycaeus. 

1. Genus Microsetella Bradj and Robertson. 

MicroseteUa Brady and Robertson, l^T::. p. L30, pi. 9, figs. 11-1(3, 

Harpacticus Dana, 1847, p. L52. 

Canthocamptus Dana, \^'<-, p. 1 1 ^7. 

Ectinosoma Brady, l" s -"'. p. 99. 

Ectinosoma Mbbius, L887, p. 116. 

Microsetella Giesbrecht, L892, pp. 78, 549, 750. 

$ Body cylindrical, smaller in front and behind, anterior 
poll ion with lour segments, posterior with five: furca short, 
bristles very Long. Anterior antennae 5-jointed, posterior 
3-jointed, outer ramus 3-jointed and slender. Rami of first to 
fourth feet 3-jointed. inner ramus longer than outer: fifth pair 
rudimentary and leaf-like (fig. 516). 

J Posterior portion of body with five segments, anterior 
antennae rather strong grasping organs; feet much smaller. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 211 

1. Microsetella rosea Dana. 

Harpacticus roseus Dana, 1847, p. 153. 

Canthocamptus roseus Dana, 1852, p. 1189; 1855, pi. 83, figs. 1-10. 
Microsetella rosea Giesbrecht, 1892, pp. 550, 554, pi. 44, figs. 32, 
35, 37, 38, 41, 43, 48, 49. 

Fig. 52.— Microsetella rosea. Female, (a) Lateral X83. (6) Fifth foot 
X195. Be, outer ramus, (c) Furca, ventral, X195. 

$ Longest bristle of furca almost twice as long as the body, 
third terminal bristle less than half as long as the abdomen. 
Innermost bristle of fifth foot not much shorter than the others. 

J Unknown. 

Coloration: Rather transparent, region of mouth red, and 
also long f ureal bristles ; digestive tract rosy red. 

Length : Female, 0.84—0.9 mm. 

Occurrence: June 10, 1904, five females; catch taken near 
La Jolla. 

212 University of California Publications. [Zoology 

2. Genus Euterpe Clans. 

Harpacticus Dana, 1847, p. 152; 1S52, p. 1189. 

Euterpe Claus, 1863, p. 109. 

Euterpi Giesbrecht, 1892, pp. 78, 555. 

2 Anterior portion of body with four segments, posterior 
with five; front of head pointed (cf. fig. 53a). Anterior an- 
tennae 7-jointed, posterior 3-jjointed, outer ramus 1-jointed. 
Rami of swinuning feet 2-jointed in the first pair. 3-jointed in 
second to fourth pairs-, fifth pair rudimentary (fig. 53d). 

£ First and second abdominal segments aot fused (fig. 53a): 
anterior antennae i fig. 53< I are powerful grasping organs, 
fourth and fifth joints fused and much thickened, geniculat- 
in- wiih the hooked terminal joint which is composed of the 
fused sixth and seventh joints. Rami, especially the inner, of 
the first pair of feet of peculiar form; fifth pair shorter and 

with fewer bristles than in the female. 

1. Euterpe acutifrons Dana 

Harpacticus acutifrons Dana, L847, p. L53; L852, p. 1192; 1855, 

pi. 83, fig. 11". b. 
Euterpe gracilis Claus, L863, p. 109, pi. 14, figs. 1 L3. 
Euterpt acutifrons Giesbrecht, L892, p. 555, pi. 44, figs. 16-31. 

The only species of the genus. 

Coloration: Transparent, almost without pigment, but 
digestive canal is often yellowish or green. 

Length: Male. .7:: i .; females slightly smaller. 

Occurrence: Abundanl in catches with Oithona, Oncaea and 
Corycat us. 

Vol. 2] Esterkj. — Copepoda of the San Diego Region. 213 

a hi th.5. 

Fig. 53.— Euterpe acutifrons. (a) Male, lateral, X175. Ab.l, first 
abdominal segment. Th.5, fifth thoracic segment. (6) Outer 
ramus of fourth foot of female X140. (c) First foot of male 
X265. (d) Fifth foot of female X410. (e) Anterior an- 
tenna of male X195. 

3. Genus Clytemnestra Dana. 

Clytemnestra Dana, 1847, p. 154; 1852, p. 1193. 
Clytemnestra Lubbock, 1860, p. 180. 
Goniopsyllus Brady, 1883, p. 107. 
Clytemnestra Giesbrecht, 1892, pp. 79, 565, 733. 
Clytemnestra "Wheeler, 1899, p. 188. 

5 Anterior part of body composed of four segments, pos- 
terior part of five ; furca short. Anterior antennae 7- to 8-jointed, 
bristles short, posterior antennae 3-jointed, outer ramus sup- 
plied by one or two bristles. Posterior maxilliped 2-jointed, 
slender and elongated, with short hooks at the end. Rami of 
swimming feet long and narrow, inner ramus the longer, 
3-jointed in all pairs; outer ramus 3-jointed except in first 
pair, where it is 1-jointed; fifth pair rudimentary, 2-jointed. 


University of California Publications. [Zoology 

J Posterior portion of body with six segments, furcal bristles 
sometimes lengthened. The .interior antennae are grasping or- 
gans, geniculating lid ween the last two joints; posterior maxil- 
lipeds longer, with thicker second joint and longer terminal 

1. Clytemnestra rostrata Brady. 

Clytemnestra tenuis Lubbock, L860, p. 160, pi. 29, ii^s. 6-7. 
Goniopsyllus rostratus Brady, 1883, p. 1<>7, pi. 42. figs. 9-16. 
Clytemnestra rostrata Giesbrecht, 1892, pp. 566, 572, pi. 45, figs. 

19, 20, 22, 25, 26, 31, 33. 
Clytemnestra rostrata Wheeler, 1899, p. 189, fig. 26. 

Pig. 54.— Clytemnestra rostrata. Female. (a) Dorsal, X45. (b) Forca, 
dorsal, ■ 265. 

Furca at most .-is Long as broad, bristles no1 plumose, equal 
in length in both sexes. Anterior antennae in each sex seven- 
jointed. Ias1 joint in female five limes as Long as the preceding 
one-. Lancet-shaped bristle Lacking in the male. Outer ramus 
o\' posterior antenna replaced by one bristle: second basal of 
lirst foot without bristle on outer margin, outer ramus with 
three bristles; outer ramus of second foot with one outer mar- 
ginal bristle on first and second joints, two on the third: second 
joint of inner ramus of third foot longer than the terminal 
joint. Fifth foot as long as the outer ramus of the fourth. 
with five bristles on the terminal joint, which are as long in 
the female as in the male. 

Vol. 2] Esterly.—Gopepoda of the San Diego Region. 215 

Coloration: Reddish, due to the presence of rose, brown 
or greenish oil globules in the transparent body. 

Length : Female, 1.28 mm. 

Occurrence : Rather uncommon ; one female was taken June 
14, 1904, at San Diego. Occurs also in the winter. 

Oncaeidae Giesbrecht, 1892, p. 81. 

Paired eyes with cuticular lenses and pigment bodies not 

5 Form of body in general like the Cyclopidae. Each por- 
tion of the furca has six bristles. Anterior antennae 4- to 6- 
jointed; posterior antennae 3- or 4- jointed ; mandibles reduced 
to blade, without specific form. Maxillae are bristle-bearing 
platelets, usually separated into two lobes. Anterior maxilliped 
2-jointed. Posterior maxilliped 4-, seldom 3-jointed, terminal 
hook strong. First to fourth pairs of feet with 3-jointed rami ; 
inner ramus of fourth foot longer or but little shorter than the 


J Sexual peculiarities in form of abdomen and posterior 
maxillipeds, fewer joints usually in anterior antennae, rarely in 
posterior antennae and mouth parts. 

1. Genus Oncaea Philippi. 

Oncaea Philippi, 1843, p. 62. 

Antaria Dana, 1852, p. 1227. 

Antaria Glaus, 1863, p. 158. 

Antaria Brady, 1883, p. 119. 

Oncaea Lubbock, 1860, p. 183. 

Oncaea Giesbrecht, 1892, pp. 81, 590, 755. 

Shape of body as in Oithona, Terminal joints of inner rami 
of swimming feet long and narrow, that of the fourth pair at 
least one and one-half times as long as the first and second 
together ; fifth foot rod or knob-shaped. 

5 Both portions of body with five segments (figs. 55a, 56a). 
Anterior antennae 6-jointed, posterior 3-jointed, hooked bristles 
of medium length (cf. fig. 56c). Posterior maxillipeds 4-jointed, 
rows of spines on inner border of second basal. Outer mar- 
ginal bristles of outer rami of first and second feet as follows : 

216 University of California Publications. [Zoology 

One on the first and second, three on the third joint ; of the third 
and fourth feet, one on the first and second, two on the third 

J Abdomen with five segments, genital segment Large, lips 
of the orifice with spines at the sides. Posterior maxilliped with 
more muscular second basal, and more strongly curved terminal 
hook than in the female. In the anterior antennae the three 
short terminal joints are fused into one piece. 

1. Oncaea conifera Giesbrecht. 

Antaria mediterranea Claus, l v »;:;, p. l~>'.>, pi. mo, figs. 1-7. 
Oncaea conifera Giesbrecht, 1892, pp. 591, 603, pi. 2, fig. 10; pi. 
17, figs. 1. 16, 21, 28, 34, 38, 12, 55, 56. 

BHg. 55.- Oncaea conifera. Female, (a) Lateral X45. (b) Posterior 
maxilliped ■ 265. 

$ Median portion of second thoracic segmenl protruding 
from the dorsal surface of the body (fig. 55a), genital segment 
almost one and one-half times as long as the rest of the abdomen, 
the following segments broader than Long. Furca as Long as 

the fifth abdominal segment, between two and two and one-half 
times as Long as broad, its branches strongly directed away 
from each other. Hook at end of posterior maxilliped set with 
thick spines, distal bristles of second basal heavier and longer 
than the proximal (fig. 556). Processes at end of third joint 
of inner ramus of sw immin g feet very Large, present even in the 
fourth pair, the adjacent lancetdike bristle shortened. Fifth pair 
of feet elongated, with thickened terminal bristle. 

J Lips of genital orifice long, furca short and broad. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 217 

Coloration : Often distinctly green-yellow tint to body, which 
is not very transparent. 

Length: Female, 1.2 mm. ; male, about 0.8 mm. 

Occurrence : A few were taken June 14, 1904, and in some 
cases the sexes were pairing. 

2. Oncaea minuta Giesbrecht. 

Oncaea minuta Giesbrecht, 1892, p. 603, pi. 47, figs. 3, 6, 26, 46, 59. 
Oncaea minuta (male) Aurivillius, 1899, p. 29, figs. 1-3. 

Fig. 56. — Oncaea minuta. Female. (a) Lateral X140. (&) Posterior 
maxilliped X265. (c) Posterior antenna X265. 

$ Genital segment longer than the rest of the abdomen, the 
following segments broader than long; furca shorter than the 
fifth abdominal segment, less than twice as long as broad, inner- 
most terminal bristle shorter than the outermost. Posterior an- 
tennae retrograded; terminal hook of posterior maxillipeds, and 
the distal bristles of the second basal provided with spines. Outer 
ramus of swimming feet narrow, end joint of inner ramus even 
in the fourth with terminal processes and smooth proximal outer 
marginal bristle. 

J Unknown ( ? ) 

Coloration : Reddish throughout body ; eggs reel. 

Length: Female, 0.46-0.5 mm. 

Occurence : Rather uncommon ; a few come during the sum- 

218 University of California Publications. [Zoology 

Fam. Corycaeidae. 
Corycaeidae Giesbrecht, 1892, p. 83. 

Paired eyes highly developed in both sexes or in females, 
with large cuticular lenses and pigment bodies. 

J The broad front and the two chitin lenses, sometimes con- 
tiguons and sometimes separated, are characteristic of the body 
form (fig. 576; 62.). Anterior portion of the body may be coni- 
cal (Corycaeus fig. 62) or cubical (CopiUa) or ova] and de- 
pressed (Sapphirina fig. 58a, b; 59a Corina). The number of 
segments may be Id ( Sapphirina > . S (Carina. Cnpilia) or 7 to 4 
(Corycaeus); each pari of the furca with only four or five 
bristles. Anterior antennae 3- to 6-jointed, posterior antennae 
( fig. .'tSr ! with at least a heavy terminal hook; mandibles reduced 
to blade-, maxillae oval or elongate platelets, with 3 to •"> bristles; 
anterior maxillipeds as in the Oncai idcu . posterior 3-jointed, ter- 
minal hook strong. Rami of swimming feel 3-jointed, except in 
the case of the inner ramus of the fourth pair, which shows all 
transitions from the 3- jointed ramus to a rudimenl consisting of 
a single brisl Le. 

J Sexual peculiarities m form of body and posterior maxil- 
lipeds. also in the other appendages and more striking than in 

t lie Cm a* alai . 

1. Genus Sapphirina •!. V. Thompson. 

Sapphirina J. V. Thompson, Is 29. 

Sapphirina Templeton, 1836. 

Sapphirina Dana, 1*48, p. 41; L849, p. 281; 1852, p. L234. 

Sapphirina ( 'laus, lsi',:;. p. ] }<». 
Sapphirina lla.vkcl, ls64, p. 10l'. 
Sapphirina Brady, 1SS.S, p. 1'21. 
Sapphirina Giesbrecht, L892, pp. 84, 618, 761. 
Sapphirina Wheeler, 1899, p. 190. 

Body depressed: anterior and posterior portions of body 
with 5 segments in the female, middle abdominal segments broad- 
ened. Furca leaf-like, with five bristles. Kami of feet broad, in 
first, second, and third pairs about equal in length; inner ramus 
of fourth pair with 3 joints, of varying relative size: fifth pair 
of feet with two bristles. Male with leaf-like broadened seg- 
ments in trunk, iridescent; no general sexual peculiarities in 
mouth parts and swimming feet. 

Vcl. 2] Estcrhj. — Copepoda of the San Diego Region. 219 

§ Eye lenses contiguous or close together. Genital orifice 
placed far at the side of the segment. Anterior antennae 3- to 5- 
jointed, posterior antennae (fig. 58c) with a short hooked bristle 
on the terminal joint and slender bristles elsewhere. The termi- 
nal joint of the anterior maxillipeds is drawn out into a long 
spine; hook at end of posterior maxillipeds short and thick. 
Outer rami of swimming feet with broad-edged, lancet-shaped 
outer marginal bristles ; in the first to third pairs the first, sec- 
ond, and third joints have respectively one, one, three bristles; 
in the fourth pair, one, one, two (three). The first, second and 
third joints of the inner ramus of the first foot have respectively 
one, one, six bristles ; of the second foot one, two, six ; of the 
third one, two, five, and of the fourth one, two, two, (one). 

J Abdomen with five segments; genital valves broad but 
short, with several bristles; hooks at end of posterior maxillipeds 
elongated, and articulating with the second basal by means of an 
intervening joint (fig. 60a). 

1. Sapphirina iris Dana. 

Sapphirina iris Dana, 1849, p. 41; 1852, p. 1239; 1855, pi. 87, 

figs. 1 a-d. 
Sapphirina salpae Clans, 1863, p. 152. 
Sapphirina gemma Brady, 1883, p. 127; pi. 48, fig. 6-8. 
Sapphirina salpae Giesbrecht, 1892, pp. 618, 641; pi. 2, fig. 9; pi. 

52, figs. 1, 2, 18, 19, 27, 45, 51; pi. 53, figs. 7, 23, 24, 60; 

pi. 54, figs. 9, 13, 15, 16, 19, 57. 
Sapphirina iris, Giesbrecht, 1895, p. 261. 

5 Furca more than 2% times as long as broad, inner border 
more convex than outer, a small point at end of inner margin 
(fig. 57c) ; the dorsal bristle is placed farther back than those on 
the outer border. Anterior antennae 5-jointed, 6/7 as long as the 
posterior; second joint 1% times as long as the three terminal 
joints. Inner ramus of posterior antennae about % the length 
of the second basal, end hook half as long as the second joint of 
the inner ramus. Inner ramus of fourth foot little shorter than 
the outer, third joint of latter with 3 outer marginal bristles. 
Third joint of inner ramus not much shorter than the other two 
together, with two bristles on the end. 


University of California Publications. 


J Body about 1 M L . times as Long as its greatest width (fig. 
57a). Eye lenses ventral, overhung by the margin of the front 
of the head. Furca, anterior antennae, fourth pair of feet like 
same parts in the female, posterior antennae, mandible, maxilla, 

Fig. 57. — Sapphirina iris, (a) Male, dorsal, X9. (6) Female, dorsal, 
X9. (c) One ramus of furea of female, dorsal, X83. (d) 
Outer ramus of fourth foot of male X140. 

anterior maxillipeds somewhat different; terminal joint of inner 
ramus of second pair with 3 lancet-like bristles. 

Coloration : Egg cases red ; body rather transparent and 
strikingly iridescent in the male. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 221 

Length : Female, 5-7 mm. ; male, 7-8 mm. 

Occurrence: Both sexes are rather common, in winter and 
summer collections. 

2. Sapphirina angusta Dana. 

Sapphirina angusta Dana, 1849, p. 41; 1852, p. 1240; 1855, pi. 

87, figs. 3a, &. 
Sapphirina danae Lubbock, 1856, p. 33, pi. 12, figs. 9-11. 
Sapphirina clausii Haeekel, 1864, p. 104, pi. 2, figs. 21-25. 
Sapphirina angusta Giesbrecht, 1892, pp. 619, 641 ; pi. 52, figs. 5, 
6, 53, 58, 66; pi. 53, figs. 6, 17, 29, 30; pi. 54, figs. 2, 8, 17, 

20, 60, 61. 

Fig. 58. — Sapphirina angusta. (a) Female, dorsal, furca not shown, X14. 
(h) Fourth foot of female X160. Be., outer ramus. (c) 
Posterior antenna, female, X160. B.2, second basal joint. 
Ei.2, second joint of inner ramus. (d) Furca of male, 
dorsal, X60. 

$ Head longer than broad; furca almost twice as long as 
broad, with a broad tooth at end of inner border (fig. 58d), dor- 
sal bristle placed farther back than the outer marginal bristles. 
Anterior antennae 5- jointed, 5/6 as long as the posterior, second 
joint 5/4 as long as the 3 terminal joints together. Inner ramus 


University of California Publications. [Zcolog-j 

of posterior antennae 5 7 as long as the second basal joint ter- 
minal hook -;; ;|S long as the second joint of the inner ramus 
(fig. 58c). Inner ramus of fourth foot little shorter than the 
outer: terminal joint of inner ramus about :? 4 the length of 
the first and second joints together, with two bristles on the end 
(fig. 586). 

J Length of trunk- 2' j as much as its greatesl diameter. Eye 
lenses as in iris. Furca, fourth pair of feet, anterior antennae 
as in the female, the other appendages somewhat different ; ter- 
minal joint of inner ramus of second foot with 3 lancet bristles 
and elongated teeth. 

Coloration: Egg cases bine, otherwise as 8. iris; the mules 
are brilliantly iridescent. 

Length: Female, 2.5-5 mm; male, 3-5 mm. 

Occurrence: Both sexes occur frequently in summer and 

3. Sapphirina scarlata Giesbrecht. 

Sapphirina scarlata Giesbrecht, 1892, p. 642; pi. 52, figs. 42, 60, 
6] : pi. 53, figs. L2, 39; pi. 54, figs. 25, 31, 72. 

Pig. 59.— Sapphirina scarlata. Female, (a) Dorsal, X18. (b) Fourth foot, 
X140. fit., inner ramus. 

5 Head broad, furca hardly twice as long as wide: inner mar- 
ginal bristle placed a little farther forward than the outer mar- 
ginal. Anterior antennae 5-jointed, not half as long as the pos- 
terior, second joint l 1 /! times as long as the terminal joint. 

Vol. 2] Esterhj. — Copepoda of the San Diego Region. 223 

Inner ramus of posterior antennae longer than the second 
basal ; terminal hook half as long as second joint of inner ramus. 
Inner ramus of fourth foot half as long as the outer (fig. 596), 
terminal joint of inner ramus as long as first or second joints, 
with two bristles at the end. 

J Length of trunk not quite twice its greatest breadth, eye 
lenses set back almost on the margin of the front. Furca, an- 
tennae and anterior mouth parts as in the female ; terminal joint 
of outer ramus of second foot with two lancet bristles, the three 
thick, awl-shaped teeth elongated. 

Coloration : Transparent, with bright red spots on the thorax 
and abdomen. 

Length : Female, 3.3 mm. 

Occurrence : One adult female was taken ; immature speci- 
mens have come in at other times. 

4. Sapphirina lomae n.sp. 

Fig. 60. — Sapphirina lomae, n. sp. Male, (a) Posterior maxilliped X83. (6 ) 
Anterior antenna, bristles omitted, X83. (c) Third joint 
of inner ramus of second foot X195. (d) Posterior an- 
tenna X83. 

Resembling S. nigromaculata in general; anterior antennae 
(fig. 606) nearly half as long (7/16) as posterior and 5- jointed; 
second joint shorter than the three terminal joints together. 
Inner ramus of posterior antennae (fig. 60d) nearly half again 

224 University of California Publications. [Zoology 

as long as the second basal, terminal hook not 1/5 as long as the 
second joint of the inner ramus. Inner ramus of fourth foot not 
1 2 as long as the outer ramus, third joint of former with two 
terminal bristles; third joint of inner ramus of second foot 
(fig. 60c) with 2 lancet bristles, the third notched on one side; 
projections on margin of joint much elongated. 

8. lomcu differs from S. nigromaculata most in the relative 
lengths of the joints of the posterior antennae and in the form of 
the toothed bristles on the terminal joint of the inner ramus of 
the second foot. Two males were taken on May 31, 190-1. but 
are so badly mutilated thai it is impossible to make a drawing of 
the entire animal. 

Lengl li : 3.2 nun. 

Occurrence: San Diego, .May 31, 1904, 2 males. 

XoTE. — The species of Sapphiiiiui are separated into two general groups. 
according as the inner ramus of the fourth foot is very small and narrow 
(fig. 596), compared with the outer ramus, or at least % as long as the 
outer. S. iris and S. <iiifiust<i belong in the latter group, ami flf. SCarlata ami 
8. lomae in the former. 8. iris is distinct from any other Bpecies \ u having 
,"> bristles on the miter margin of the third joint of the outer ramus of the 
fourth font; 8. angusta may lie recognized by the shape of the furcal rami. 

2. Genus Corycaeus Dana. 

Corycaeus Dana. r848, p. :;">; 1849, p. 280; L852, p. 1203. 

Corycaeus Lubbock, 1856, p. 32; Is.",;, p. 409; 1860, p. 182. 

Corycaeus elans. 1863, p. L54. 

Corycat us Giesbrecht, 1891, p. 480; 1892, pp. 85, 659, 7:'..". 

Cori/cdias Dahl, 1894b. p. ti7. 

Corycat us Wheeler, L899, p. L91. 

$ Eye lenses close together, in some e,-ises contiguous; fifth 
thoracic segment very short. Anterior antennae 6-jointed, 
bristles not plumose; second basal of posterior antennae large, 
firsl basal very short, each provided with a Long, heavy bristle, 
inner ramus with a thick, strongly curved hooked bristle. Ter- 
minal joint of anterior maxilliped ends in a strong hook; second 
basal of posterior maxilliped with one bristle on the inner mar- 
gin, terminal hook more slender than in Sapphirina. Outer 
ramns of swimming feet longer than inner rami; outer marginal 
bristles of outer ramus in first and second pairs are lanceolate 
and dentate, and are more or less suppressed in the third and 
fourth pairs. 

Vol. 2] Esterly.- — Copcpoda of the San Diego Region. 225 

J Lips of genital orifice long, with one bristle; the posterior 
antenna and maxilliped show distinct differences, especially in 
the elongation of the terminal hook. 

The genus may be readily recognized by the cylindrical shape 
of the body, with the eye lenses at the anterior end. 

1. Corycaeus venustus Dana. 

Corycaeus venustus Dana, 1849, p. 280; 1852, p. 1222, pi. 86, 

fig. 4a. 
Corycaeus limbatus Brady, 1883, p. 114, pi. 49, figs. 18-22. 
Corycaeus venustus Giesbrecht, 1892, pp. 659, 674, pi. 51, figs. 32, 

33, 34, 47. 

Fig. 61. — Corycaeus venustus. (a) Female, dorsal, X83. (b) Fourth foot, 
female, X140. 

$ Cephalothorax with 4 segments, abdomen with 2, ventral 
keel rounded, f urea almost 5 times as long as broad ( Genital seg- 
ment : anal segment : furca : : 3 :2 :2) . 


University of California Publications. [Zoology 

J Genital segment about % as long as the anal segment and 
furca together. 

Coloration : Varying amounts of red or yellow red pigment in 
mouth region, posterior thoracic segments, and genital segment; 
eye red. 

Length: Female, 0.8-1 mm.; male, not over 0.8 mm. 

Occurrence: A few were takes June 16, 1901. 

2. Corycaeus carinatus Giesbrecht. 

Corycaeus carinatus Giesbrecht, 1892, pp, 661, 675; pi. 51, figs. 

20, 26. 
Corycaeus carinatus Wheeler. lsiut, p. 19i\ fig. 30. 

Fig. 62.— Corycaeus carinatus. (a) Female, dorsal, X40. (b) Female, 
lateral, X40. 

$ Cephalothorax with 2 segments, abdomen with 1; ventral 
keel beak-like and pointing back; the abdomen tapers toward the 

posterior end, furca half as long as the rest of the abdomen, 
about 4 times as Long as broad. 

J Unknown. 

Coloration : Red or yellowish fed pigment in region of mouth, 
extensions of thoracic segments, and in the genital segment; eye 

Length: 0.86 mm. to 0.92 mm. 

Occurrence: A few specimens taken at San Diego, Dec. 30, 
1903, and Jan. 1, 1901. 

Cambridg( . Mass., 

January 1. 1V(>.~>. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 227 


Aurivillius, Carl W. S. 

1899. Animalisehes Plankton ans dem Meere zwischen Jan Magen, 
Spitzbergen, K. Karls Land und der Nord Knste Nor- 
wegens. Kongl. Svensk. Vet.-Akad. Hand., Bd. 32, No. 6, pp. 
1-71, figs. 1-5. (Heterorhdbdus angulata, new species). 

Baird, W. 

1843. Notes on British Entomostraca. Zoologist (Newman), Vol. 1, pp. 
193-197. (Description of Oithona and 0. plumifera.) 

Boeck, Axel. 

1864. Oversigt over de ved Norges Kyster jagtagne Copepoder hen- 
horende til Calanidernes, Cyclopidernes og Harpactidernes 
Familier. (Citations from Giesbrecht, 1892, 1898.) Forh. 
Selsk. Christian. (Description of Paracalanus, Metridia). 

1872. Nye Slaegter og Arter af Saltsvands-Copepoder. Vidensk.-Selsk. 

Forhandl. Christian., 28 pp. 

Bourne, G. C. 

1889. Beport on the pelagic Copepoda collected at Plymouth in 1888-89. 
Jonr. Marine Biol. Assoc, London (2), Vol. 1, pp. 144-152, 
pis. 11-12. (Description of Gynmoplea found and Oithona, 
Euterpe, Corycaeiis, Oneaea). 

Brady, G. Stewardson. 

1878-1880. A monograph of the free and semi-parasitic Copepoda of 
the British Islands. Bay Society, London. 
1S78. Vol. 1, pp. 1-148, pis. 1-33. 
1880a, Vol. 2, pp. 1-182, pis. 34-82. 
1880&. Vol. 3, pp. 1-83, pis. 83-93. 
1883. Beport on the Copepoda collected by H.M.S. "Challenger" dur- 
ing the years 1873-1876. Challenger Sep., Vol. 8, 142 pp., 
55 pis. 
1899. On the Marine Copepoda of New Zealand. Trans. Zool. Soc, Lon- 
don, Vol. 15, part 2, pp. 31-54, pis. 9-13. (New family Ento- 
molepidae; 4 new genera; new species in Acartia, Temora, 
Centropages (2), Ectinosoma). 

Brady, G. S. and David Robertson. 

1873. Contributions to the study of the Entomostraca. VIII. On 

Marine Copepoda taken in the west of Ireland. Ann. Mag. 
Nat, Hist., Vol. 12, ser. 4, pp. 126-142, pis. 8-9. 

Canu, E. 

1896. Copepodes. Besult scient. Camp. Caudan, pp. 421-437. (Abstract 
in Jour. Eoy. Mic. Soc, London, 1897, p. 124). [Neoscole- 
cithrix new genus = Scolecithrix Koehleri.~\ 

228 University of California Publications. [Zoology 

Car, L. 

1884. Ein Beitrag zur Copepoden-fauna des Adriatischen Meerea. Arch. 
Naturg., Jahrg. 50, Bd. 1, pp. 236-256, pi. 17. (Descriptive 
notes; synonymy, distribution. Good literature list and re- 
Claus, C. 

1863. Die freilebenden Copepoden mit besonderer Beriicksichtigung der 
Fauna Deutschlands, der NTordsee and dea Mittelmeeres. Leip- 
zig, Verl. W. Engelmann., 230 i>p., 37 pis. (Monograph deal- 
ing with morphology very largely; descriptions of genera and 
species I. 

1866. Die Copepoden-Fauna von Nizza. Ein Beitrag zur Charakteristik 
der Pormen and deren Abanderungen "im Sinne Darwin's." 
Schriften Ges. Naturw. Marburg, 1. Supplem.-heft, 35 pp., 5 
pis. (Discussion of variability, and completion of some de 
scriptions from 1 s (i.".. | 

1893. Ueber die Entwicklung und das System der Pontelliden. Arb. 
Zool. Inst. Wien u. Zool. St. Triest, Vol. 10, pp. 233 -282, 
pis. L-5. 
Cleve, P. T. 

L900. Notes on some Atlantic plankton Organisms. Kongl. Svensk. Vet- 
ensk.-Akad. Handl., Bd. 34. \o. 1. pp. 1-22, pis. 1-8. (New 
species in Acartia ; male of EucMrella rostrata). 

1901. Plankton from the Indian Ocean and Malay Archipelago. Kongl. 
Svensk. Vetensk. Akad. Eandl., Bd. 35, No. 5, pp. 1-58, pis. 1-8. 
(New species in dcrocalanus, Calanopia, Tort anus, Pseudodi 
aptomus; Metacalanus new genus.) 
Dahl, F. 

l s '.i.".. Pleuromma, ein Krebs mit Leuchtqrgan. Zool. Anz., Vol. !<i, pp. 
L04 L09. (Addition of several species to the genus; consid- 
eration of lateral pigmented body as a phosphorescent organ). 

1894a. Leuehtende Copepoden. Zool. An/... Vol. 17, pp. 10-13. 

1894&. Ueber 'lie horizontale und verticale Verbreitung der Copepoden 
im Ocean. Verh. Deutsch. Zool. Gesell. auf vierten Jahresvers., 
pp. 61-80, 4 ti-s. (Distribution, diagrams ami keys for 
Corycaeus, Calanus, ami Heterochaeta. Five new species in 
Corycat us, 4 in Hett rocha< ta I. 

L894c. Die Copepodenfauna des onteren Amazonas. Ber. naturf. 
Gesell. l'reili., n. ser., Vol. 8, pp. 1-11, pi. 1. (Acartia 
gu sbu clii i new. Labidoa ra flvrviatilis new, Paracalanus crass- 
irostris new.) 

Dana, J. D. 

1845. Description of a new genus of Cyelopidae. Proc. Acad. Nat. Sci. 
Phil., Vol. 2, pp. 28o-286. (Original description of Corycaeus ; 
brief descriptions of other genera and species). 
184C. Notice of some new genera of Oyclopacea. Am. Jour. Sci. and 
Arts, ser. 2, Vol. 1, pp. 225-230; Ann. Mag. Nat. Hist., Vol. 18, 
pp. 181-185. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 229 

1847. Conspectus Crustaceorum, in orbis terrarum circumnavigatione, 

C. Wilkes e Classe Eeipublicae Federatae duce, collectorum 
auctore J. D. Dana. Part 1. Proc. Am. Acad. Arts and Sci., 
Vol. 1, pp. 149-155. (Cyclopidae, Ilarpacticidae). 

1848. Conspectus Crustaceorum quae in orbis terrarum circum- 

navigatione, Carolo Wilkes e Classe Eeipublicae Federatae 
duce, lexit et descripsit Jacobus D. Dana, Part II. Proc. Am. 
Acad. Arts and Sci., Vol. 2, pp. 9-55. (Calamus, Scribella, 
Euchaeta, Undina, Candace, Cyclopina, Catopia, Acartia, Pon- 
tella, Corycaeus, Antaria, Copilia, SappMrina, Miracia.) 

1849. Same title. Am. Jour. Sci. and Arts, ser. 2, Vol. 8, pp. 276-285, 

(Citations of descriptions of genera and families; species 
named only). 

1852. Crustacea. U.S. Expl. Exped., during years 1838, 1839, 1840, 
1841, 1842, under the command of Charles Wilkes, U.S.N., 
Vol. 13, pt. 2, pp. 1019-1262. 

1855. Atlas, 96 pis.; Copepoda, pis. 70-78. (Monograph; system of 
classification here, adopted by later writers; descriptions very 
brief and unsatisfactory, figures not from high magnifica- 
Foster. E. 

1904. Notes on the free-swimming Copepods of the waters in the vicinity 
of the Gulf Biologic Station, Louisiana. 2nd Eep. Gulf Biol. 
Stat., pp. 69-79. 

Giesbrecht, W. 

1888-89-91. Elenco dei Copepodi pelagici raccolti dal tenente di vascello 
G. Chierchia durante il viaggio della E. Corvetto "Vettor 
Pisani, ' ' negli anni 1882-85 dal tenente di vascello Fran- 
cesco Orsini nel Mar Eosso, nel 1884. Atti Accad. Lincei, 
Eoma (4) Eend. 

1888. Vol. 4, 2 Sem., fasc. 10, pp. 285-338. 

1889. Vol. 5, 1 Sem., fasc, 11, pp. 1-10. 
1891a. Vol. 7, 1 Sem., fasc. 10, pp. 474-481. 
1891b. Vol. 7, 2 Sem., fasc, 8, pp. 276-282. 

(List of 229 species with localities; new genera and species 
briefly described in Latin). 

1892. Systematik und Faunistik der pelagischen Copepoden des Golfes 
von Neapel. Fauna und Flora des Golfes v. Neapel, Vol. 19, 
text 831 pp., atlas 54 pis. (Monograph; descriptions, dis- 
tribution, cause of migration, 59 genera, 299 species). 

1895. Eeports on the dredging operations off the west coast of Central 
America to the Galapagos, to the west coast of Mexico, and in 
the Gulf of California, in charge of Alex. Agassiz, carried on 
by the U.S.F.C. Steamer Albatross, during 1891, Lieut. Comm. 
Z. T. Tanner, U.S.N., commanding. XVI. Die pelagischen 
Copepoden. Bull. M.C.Z., Vol. 25, No. 12, pp. 243-261, pis. 1-4. 

230 University of California Publications. [Zoolou* 

1896. Ueber pelagisehe Copepoden des Kothes Meeres gesammelt vom 

Marinestabartz Dr. Augustin Kramer. Zool. Jahrb., Abth. 
System.. Bd. '.». Elf. 2, pp. 315-327, pis. 5-6. (Five new species 
in Scolcfilltri.r, Cent ropuijes, Monops, OiUiona, SchmacJceria) . 

1897. Notizen znr Systematic der Copepoden. Zool. An/... Bd. 29, pp. 

253-255. ( Female of Arit U Hits s< tosus, Scolecithrix similis). 

Giesbrecht, W. and O. Schmeil. 

1898. Copepoda. I. Gymnoplea. Das Tierreich (Schulze), Lief. 6, Merlin, 

189s. pp. XVI. L69, 31 figs. (5 families, 65 genera, 370 estab- 
lished ami 105 doubtful species, •"> varieties. Literature com- 
plete in end of 1897). 

Gunner, J. E. 

1765. Nogle smaa rare, mestendelen aye aorske Sodyr, beskrevene. 
Skrifter, Kjobenhavnske Selsk. (Acta Havn.), Bd. 10, p. 17.". 
(Citations from Giesbrecht, 1892, 1898). [Description of 
Monocwlus finmarchicus-Calanus finmarchicus. | 

Haeckel, E. 

L864. Beitrage zur Kenntnis der Corycaiden. Jena. Zeits. Med Naturw., 
Bd. 1. pp. iil 112, pis. 1 •"•. (Systematic; anatomy of male of 
Copilia ami Sapphit 

Herdman, W. A. 

1891. The biological results of the cruise of the s. v. "Argo" round 

the west mast of Ireland ill August. 1S90. Trans. Biol. Soc. 

Liverpool, Vol. 5, pp. L81-212. (32 species diagnosed by I. 
< '. Thompson ). 

Herdman, W. A., I- C. Thompson, A. Scott. 

1S97. On the plankton collected continuously during two transverses 
of the North Atlantic in the Bummer of l x '.'7. with descrip- 
tions of new species of Copepoda and an appendix <»n dredg- 
ing in Pugel Sound. Trans. BioL Sue. Liver] 1. Vol. 12, 

pp. 33-90, pis. 1-4, 4 figs. (39 species of Copepoda, 4 new in 
Eurytemora, Acartia, Corynura.) 

Herrick, C. L. 

1887. Contributions to the fauna of the Gulf of Mexico and the 

South. Mem. Denison Se. Assoc., Vol. 1. No. 1, pp. 1-56, 
pis. 1-7. (List of fresh water and marine Crustacea of Ala- 
bama; new species; keys). 

Kramer, Augustin. 

1895. On the most frequent pelagic Copepods and Cladoceres of the 

Hauraki Gulf. Trans, and Proc. N. Zeal. Inst., Vol. 27, 
pp. 214-2:'..'!, pis. 15-23. (New species in Corycaeus and 

1896. Zwei neue Pontella-Arten aus Neu-Stid-Wales. Zool. Jahrb. 

Abth. Syst., Bd. 9, pp. 720-724, 4 figs. 

Vol. 2] Esterly. — Copepoda of the San Diego Region. 231 

Kroyer, Henrik. 

1842-1845. Crustaces. Voyages de la commission scientifique du Nord 
en Scandinavie . . . pendant les Annees 1838, 1839, 
1840, sur la corvette "La Becherche. " Pnbl. by Paul 
Gaimard, Paris. Atlas, pis. 41-43. (Citations from Giesbrecht ; 

1892, 1895.) 

Leach, W. E. 

1819. Article: Entomostraces. Diet. Se. Nat. Strasbourg and Paris, 
T. 14, p. 524. (Description of genus Calanus.) 

Leuckart, R. 

1859. Carcinologisches. Arch. Naturg., Jahrg. 25, Bd. 1, pp. 232- 

262, pis. 6-7. 

Lubbock, John. 

1853a. Description of a new genus of Calanidae. Ann. Mag. Nat. 

Hist. (2), Vol. 11, pp. 25-29, pi. 1. (Labidocera darwinii. 

new genus, new species; first description of Labidocera.) 
1853&. On two new subgenera of Labidocera. Ann. Mag. Nat. Hist. 

(2), Vol. 11, pp. 25-29, pi. 10. 
185G. On some Entomostraca collected by Dr. Sutherland in the 

Atlantic Ocean. Trans. Entom. Soc. London (2), Vol. 4 r 

part 2, pp. 8-39, pis. 2-12. 
1857. Description of eight new species of Entomostraca found at 

Weymouth. Ann. Mag. Nat. Hist., Vol. 20, pp. 401-410, pis. 


1860. On some oceanic Entomostraca collected by Captain Toynbee. 

Trans. Linn. Soc. London, Vol. 13, pp. 173, 192; pi. 29. 

Mobius, Karl. 

1887. Systematische Darstellung der Thiere des Planktons, gewonnen 
in der westlichen Ostsee, und auf einer Fahrt von Kiel in 
den Atlantischen Ocean bis jenseit der Hebrides. Koram. 
Unters. Deutsch. Meere, 5 Ber., 12-16 Jahrg., pp. 109-126; 
pi. 7-8. (19 pelagic species; tables of distribution, etc.) 

Norman, A. M. 

1903. Copepoda Calanoida, chiefly abyssal from the Faroe Channel 
and other parts of the North Atlantic. Jour. Linn. Soc. 
London, Zool., Vol. 29, pp. 133-141. (List of species, notes 
on distribution.) 
Philippi, A. 

1843. Fernere Beobachtungen iiber die Copepoden des Mittelmeeres. 
Arch. Naturg., Jahrg. 9, Bd. 1, pp. 54-71, pis. 3-4. (Original 
descriptions of Euchaeta, Oncaea.) 

Sars, G. O. 

1900. Crustacea. The Norweg. North Polar Exped., 1893-1896. Edited 
by Fritltjof Nansen, Vol. 1; Crustacea, V; Copepoda, pp. 35- 
126, pis. 7-35. (3 new genera, 12 new species.) 

232 University of California Publications. [Zoology 

Scott, Andrew. 

1890a. Description of new and rare Copepoda. (1) Rep. Sea-Fish. 
Lab., Liverpool, pp. 32-56, 5 pis. (2) Trans. Liv. Biol. Soc, 
Vol. 10, pp. 134-158. \3) Rep. Fauna Liv. Bay, Vol. 5, 
pp. 59-86. (10 new species, 1 new genus.) 
1890b. On Scolecithrix hiberwica, a new species of Copepod, with some 
remarks on the distribution of the Crustacea. Ann. Mag. 
Nat. Hist. (6), Vol. 18, pp. 362-367. 
L902. On some Red Sea ami Indian Ocean Cope-pods. Trans. Liv. 
Biol. Soc, Vol. 16, pp. 397-428, pis. 1-3. (New species in 
Candacia, Calanopia, Stenhelia, Delavalia.) 

Scott, Thomas. 

1894. Report on Enitmiostraea from the Gulf of Guinea, Trans. Linn. 
Soc. London, Ser. 2, A T ol. 6, pp. 1-16, pis. 1-15. 
Scott, Thomas and Andrew. 

1896. A revision of the British Copepoda belonging to the genera 
Bradya Boeck and Ectinosoma Boeck. Trans. Linn. Soc. 
London (2), Zool., Vol. 6, pp. 419 146, 4 pis. (Six species 
in Bradya, 5 new ; 1 I in Ectinosoma, 9 new.) 

Steuer, Adolf. 

1904. Copepoden der Valdivia-Expedition. Zool. An/.., Bd. 27, pp. 
593-598, 4 figs. (Valdwiella new genus; new Bpeeies in 
Ludcutia and AugaptUus. > 

Streets, T. H. 

L877. Contributions to the Natural History of the Eawaiian and 
Panning Islands and Lower California, made in connection 
with the U. S. North Pacific Surveying Expedition, 
1873-1875. Smithson. Misc. Coll., Vol. L3, No. 7. pp. 138-141. 

Thompson, I. C. 

1888c/. Copepoda of Madeira and the Canary Islands. Jour. Linn. Soc. 

London. Vol. 20 (1890), pp. 145-156, pis. 10-13. (64 species; 

6 new, 3 new genera including M ecynocera.) 
188Sb. Report on Copepoda collected in .Maltese sens. Proc. Biol. Soc. 

Liverpool, Vol. 2, pp. 137-151, pis. 6-9. 
1888c. Second report on the Copepoda of Liver] I Bay. Proc. Biol. 

Soc. Liverpool, Vol. 2, pp. 63-71, pis. 1-2. 
1896a. Revised report on the Copepoda of Liverpool Bay. Rep. Fauna 

Liverpool Bay, Vol. 4, pp. 81-136, 21 pis. 
1896b. Free swimming Copepoda from the west coast of Ireland. 

Trans. Liv. Biol. Soc, Vol. 10, pp. 92-102. (Distribution 

and biology; tables of 18 genera, 22 species). 
1898. Contributions to our knowledge of the Plankton of the Faroe 

Channel. Xo. IV. Report on the Copepoda collected by 

Dr. G. H. Fowler. Proc. Zool. Soc. London, 1898, pp. 540- 

544. (Table; distribution.) 

Vol. 2] Esterhj. — Copepoda of the San Diego Region. 233 

1900. Eeport on two collections of tropical and more northerly plank- 
ton. Trans. Liv. Biol. Soc, Vol. 14, pp. 262-294, 1 pi., 3 figs. 
(Two tables, and notes on distribution.) 

1903. Report on the Copepoda ^obtained by Mr. George Murray dur- 
ing the cruise of the ." Oceana" in 1898. Ann. Mag. Nat. 
Hist. (7), Vol. 12, pp. 1-36, pis. 1-7. (Four new species 
in Euchaeta, Scolecithrix, Xanthocalanus, Isochaeta.) 

Thompson, I. C. and Andrew Scott. 

1900. Some recent additions to the Copepoda of Liverpool Bay. Trans. 
Liv. Biol. Soc, Vol. 14, pp. 139-144. 

1903. Report on the Copepoda collected by Prof. Herdman at Cey- 
lon in 1902. Report to the Government of Ceylon on the 
pearl oyster fisheries of the Gulf of Manaar, pp. 227-307, 
pis. 1-20. (283 species, 76 new, 10 new genera, including 
both parasitic and free forms.) 

Thompson, J. V. 

1829. On the luminosity of the ocean, with description of some re- 
markable species of luminous animals, Pyrosoma and Sap- 
phirina. Zoological Researches, Memoir 3; 25 pp., pis. 5-7. 
(Original description of Sapphirina.) 

Wheeler, W. M. 

1899. The free-swimming Copepods of the Woods Hole region. Bull. 
U. S. F. C, Vol. 19, pp. 157-192, 30 figs. (30 species; 4 new 
in Labidocera, Centropages, Pontella, Corynura.) 

Wolfenden, R. Norris. 

1902. The plankton of the Faroe Channel and Shetlands. Preliminary 

note on some Radiolaria and Copepoda. Jour. Marine Biol. 
Assoc, Vol. 6 (n. s.), pp. 344-371, pis. 1-4. (New species 
in Aegistlms, Gaidius, Euchirella.) 

1903. Occupation of a table at the zoological station, Naples. Report 

of the Committee. Appendix D, on the Copepod subfamily 
Aetidiinae, with a proposed revision of the classification. 
Report 72nd Meet. Brit. Assoc Adv. Sci., Belfast, pp. 263- 
267. (3 new species in Pseudaetideus = new genus for 
Chiridius armatus, Aetideus, Gaidius.) 

1904. Notes on the Copepoda of the North Atlantic Sea and Faroe 

Channel. Jour. Mar. Biol. Assoc. N. S., Vol. 7, pp. 110-146, 
1 pi., 1 fig. (21 new species in Megacalanus, new genus, 
Eucalanus, Gaetanus, Pseudaetideus, new genus, Aetideus, 
Faroella new genus, Chiridius, Candacia, Spinocalanus, Xanth- 
ocalanus 3, Scolecithrix 2, Lophothrix, Heterorhabdus, Luci- 
cutia 2, Augaptilus 2, Paraugaptilus new genus, Euchirella 
new variety. 


Vol. 2, No. 5, pp. 235-322, Pis. 4-16 December 9, 1905 












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The bryozoa of the west coast of North America constitute 
a fauna practically unknown to science. From time to time 
during the last half century some scattering work has been done, 
the earliest being by Trask ('57), who described and figured a 
number of species from San Francisco Bay. Later Gabb and 
Horn ('62), in connection with their work on the Geological 
Survey of California, identified a number of species of bryozoa, 
both recent and fossil ; and Hincks ( '82 and '84) reported on 
a large collection from Queen Charlotte Islands. Still later, the 
present writer ('00) reported on the bryozoa of the Harriman 
Alaska Expedition, having previously published on the Ento- 
procts of San Francisco Bay ('99). Besides the work of these 
investigators, mention should be made of that of Conrad ('55) 
and of Fewkes ('89), both of whom added to our knowledge of 
the bryozoa of this coast. 

The following pages constitute the first of a contemplated 
series of papers on the bryozoa of the Pacific coast of North 
America, and include about half of the sub-order Chilostomata, 
comprising, in the main, those Chilostomes which grow as free, 
flexible colonies. The inclusion of the adherent genera Aetea 

236 University of California Publications. [Zoology 

and Eucratea is somewhat arbitrary since they are more or less 
incrusting. In reality the forms treated comprise the first seven 
families of the Chilostomata as given by Hincks, and these with 
the exception of the two genera mentioned above are erect, flexi- 
ble and non-incrusting. Even these two genera send off erect 
free branches, and are thus partially non-incrusting. The 
bryozoa here identified comprise only a small part of the ma- 
terial on hand, the greater portion of which was collected at San 
Pedro and San Diego in 1901, 1902. and 1903 on dredging ex- 
peditions conducted by the Zoological Department of the Uni- 
versity of California; another portion was dredged in Puget 
Sound during the summers of 1903 and 1901 by the Department 
of Zoology of the University of Washington, and kindly placed 
at my disposal by Professor Trevor Kim-aid. 

In the endeavor to make this work as useful as possible to 
th«' general student as well as to the expert, original figures are 
given of all the species mentioned. These include a habil sketch 
which is a photograph wherever possible, and one or more other 
figures giving details and variations. It is thought besl even 
though a species is cosmopolitan in its range, and even though 
it has already been sufficiently well represented for diagnostic 

purposes, to refigure it here: and this for two reasons: first, 

bryozoan literature, consisting as it does, excepl in two or three 

instances, of detached papers scattered through scientific jour- 
nals is not readily accessible to the general student. Second, 
the oumber of cosmopolitan species, or of species easily identi- 
fied, is surprisingly small, while the oumber new to science is 
correspondingly large; hence as far as illustration at least is 
concerned, it is desired to make this work as complete as pos- 
sible for the forms found on the Pacific Coast. The diagnoses 
are somewhat Lengthy descriptions, with references to figures 
wherever clearness seemed to demand it. Further, in order to- 
assist in the identification of species, simple keys have been de- 
vised for the families, and where more than one genus or species 
occur under a family, separate keys are given for each. The 
keys for species cover only those treated in this paper and are 
intended only for the bryozoa of this coast. Since Hincks' ( '80) 

Vol. 2] Robertson. — Non-Incr listing Bryozoa. 237 

monograph upon the British Polyzoa is the most complete and 
exhaustive of any recent work, and its method is generally fol- 
lowed by other workers, it is here adopted as the basis of classi- 
fication, although his conclusions are not always accepted. 
Free use is made of the diagnostic definitions given by that 
author, especially for families and genera, supplementing and 
amending where necessary for greater accuracy, or in order to 
include forms peculiar to the region. Synonymy is based upon 
that of Hincks, further supplemented by Miss Jelly's invaluable 
Synonymic Catalogue ('89)1 No attempt has been made at re- 
vision of the classification. It were useless to undertake such 
a task except after careful study and comparison of all large 

It is needless here to enter into an exhaustive discussion of 
the bryozoa as a class. Such can be fouud in any of the more 
recent treatises on zoology, and the reader desirous of obtaining 
a full, clear, and delightfully written article on the group, is 
referred especially to that by Dr. Sidney F. Harmer ( '96). For 
the sake of clearness and convenient reference, however, a few 
definitions of terms used in this paper are given herewith. 

Bryozoa and Polyzoa are synonymous terms. These are 
colonial animals, and the technical term used to designate the 
colony as a whole is zoarium. An illustration of a bryozoan 
colony or zoarium may be found in any of the habit sketches, 
especially the photographs (PI. XVI). The units of which a 
zoarium is composed consist of the zocecia and their contained 
polypides. A zocecium is a chamber or sac, in which the poly- 
pide, consisting of a digestive canal and a circlet of tentacles, is 
lodged. It may be calcareous and opaque, or semi-calcareous 
or chitinous and transparent. The contents of a zocecium whose 
walls are transparent are easily made out. Below is given a 
figure of a zooecium of Beania mirabilis with its contained pohj- 
pide in a state of retraction folded within it. The zooecium 
(zce.)is seen to be a sac or bag whose front or ventral face is 
bordered with a number of spinous processes, some erect {e. sp.), 
others curved (c. sp.). Within the zocecium is the polypide 
{pd.) consisting of a bent tube, the intestinal canal, having 


University of California Publications. [Zoology 

a circlet of tentacles (tent.) around one extremity. Various 
regions of the tubular portion have specific names. In 
the middle of the membranous floor from which the tentacles 
arise is the mouth (m.). This opens into a short tube known 

Fig. 1. — Beanin mirdbilis. A zooecium and its inclosed polypide shown in 
profile view. c. .v/i. crossed spines; cur. nil. ciirdinc valve; r. */'. 
erect spine; i. intestine; m. mouth; ces. o3sophagus; or. orifice; 
par. mus. parietal muscle fibres; pd. polypide; ph. pharynx; 
py. val. pyloric valve; r. f. root fibre; re. mus. retractor muscle; 
st. stomach; tent, tentacles; tent. sh. tentacle-sheath; see. 

as the pharynx [ph.) which is really a portion of the (esophagus 
(o?s.) ; this in turn opens by a narrow valve, the cardiac valve 
(car. val.) into a stomach (st.). The stomach is a bag of a 
yellow or brown color due to the gland cells in its walls; it opens 
into the intestine (/.), by the so-called pyloric valve (py. val.). 
Visible above the tentacles when the polypide is retracted is a 
delicate membrane, the tentacle-sheath (tent, sh.). Near the 
distal end where the tentacle-sheath approaches the orifice (or.), 
a few muscle fibres may be seen on each side of the sheath ex- 
tending to the walls of the zooecium. These are part of the 
parieto-vaginal muscles which assist in retracting the tentacular 

Vol. 2] 

Robertson. — Non-Incr listing Bryozoa. 


sheath (par. mus.). Likewise, extending from the base of the 
tentacles to the basal wall of the zocecinm another band of re- 
tractor muscles is visible whose contraction draws the polypide 
within the zooecium (re. mus.). Fig. 2 represents a zocecium of 

Fig. 2.- — Bower~ban~kia imbricata. A zooecium and inclosed polypide. a. 
anus; giz. gizzard; int. intestine; m. mouth; ces. oesophagus; py. 
vol. pyloric valve; st. stomach; tent, tentacle. 

Bowerbarikia imbricata, in which the polypide is expanded. 
Here the oesophagus (oes.) is stretched out to its fullest extent, 
the mouth (m.) being at the upper margin, and the tentacles 
outside the zocecium. This polypide possesses a region between 
the oesophagus and stomach known as the gizzard lined with 
large cells (giz.). The intestine (int.) is much elongated, the 
anus (a.) reaching almost to the summit of the body wall. The 
tentacles are commonly arranged in the form of a bell, but have 

240 University of California Publications. [Zoology 

the power of independent motion as shown in this figure drawn 
from a living specimen. Many zoaria grow erect and free, and 
if calcareous, their branches frequently possess flexible joints at 
definite points along the length (PI. V, fig. 14, j.). The zooecia 
included between these joints form an internode. the number 
of zooecia in an internode being rather definite for any given 
species. In Mcnipea ternata e.g., there are, as a rule, three 
zooecia in an internode (fig. 14). From the lower zooecia small 
fibres arc given off known as rootlets, because they serve to 
anchor the colony (fig. 14, r.). Zooecia, especially if calcareous, 
often possess appendages of various kinds which serve as diag- 
nostic marks. There are also certain well marked regions to 
which special names are applied: 

Aperture the chitinous front wall of the zocecium. This may occupy 

a part of the front wall only ;is in l'l. IV, fig. 2, ap. ; or almost 

tin- whole of it as in PI. X, fig. 50, ap.). 
Area — tin' calcareous wall inclosing the whole or part of the aperture. 

(PI. XV, fig. 88, a.). 
Avicularium — an appendage of the zooecium more or less resembling a 

bird's head. This may be sessile as in IT. V, fig. 14, a/0., or 

pedunculated as in l'l. 1 1, fig. 60, «<•. 
Epistome a ciliated lobe which overhangs the mouth, and is present 

only in the Phylactolsemata or fresh water bryozoa. 
Internode — the zooecia included between the flexible joints of a branch 

or stem | IT V. fig. 1 1 I. 
Introvert- the thin cuticle at the anterior end of the polypide which 

may he retracted into the interior of the zocecium. 
Joint, or articulation — a noil calcified portion of the wall of the zoce- 
cium permitting more or less flexibility in the stem or branch 

(PI. V, fig. 14, j.). 
Lophophore — the membranous floor or rim surrounding the mouth of 

the polypide from which the tentacles spring. 
Ocecia or Ovicell synonymous terms for the chamber above the zoce- 
cium in which the embryo develops (PI. V, fig. 14, 02.). 
Operculum — the chitinous lip by which the orifice of the zooecium is 

closed (PI. IV, fig. 2, and IT. XIV, fig. 86, op.). When the 

aperture is entirely membranous, the operculum is inconspicuous. 

(PI. X, fig. 50). 
Orifice — the opening at the summit of the aperture through which the 

polypide emerges. (Text fig. \, or.). 
Scutum — a modified spine overhanging the aperture (fig. 14, sc.), often 

being large and flabellate (PI. VI, fig. 19, sc). 
Stolon — a creeping tubular stem from which the individuals of a colony 

grow. It is not found in the Chilostomata, but is characteristic 

of many of the Ctenostomata. 

Vol. 2] Robertson. — Non-Incr listing Bryozoa. 241 

Spine — a jointed or unjointed process found on the margin of the 

aperture (fig. 1 of the text, sp.). 
Tentacle-sheath — the delicate membrane of the introvert which incloses 

the tentacles when the polypide is retracted. (Text fig. 1, tent sh.) 
Vibraculum — a chitinous seta of varying length, depending on the 

species, extending from a chamber on the dorsal side of the 

zocecium (PI. IX, figs. 41, 45, v.). 
Vibracular chamber — a chamber on the dorsal side of the zocecium in 

some of the Cellulariidce, resembling an avicularium, from which 

extend a hairlike process called the vibraculum, and the rootlet 

(PI. IX, figs. 42, 46, v. ch, and v.). 

For purposes of orientation, it must be explained that the 
front or ventral side of a zocecium is that on which the aperture 
with its operculum occurs ; the side opposite is the dorsal. The 
top of a zocecium, and the ends of the growing tips of branches 
or colony are referred to as the distal or anterior end of zocecium, 
or extremity of branch or colony; the lower part or end nearest 
the root or point of origin, as the proximal or posterior extremity. 

Thirty-four species and one subspecies belonging to thirteen 
genera are here recognized. Of the genera, Stirparia has as 
yet been reported only from Australia; Synnotum only from the 
Adriatic ; and Corynoporella only from Greenland. Of the 
whole list, thirteen species and one subspecies are new, while 
twenty species and one subspecies are restricted to the Pacific 
Coast. The ovicells of Aetea anguina are here described and 
figured for the first time, as are the avicularia and ovicells of 
Stirparia, and the ovicells of Corynoporella. 


Aetea anguina Linnaeus. Bugula pacifica Eobertson. 

Aetea truncata Landsborough. Bugula flabellata Thompson. 

Eucratea chelata Linna?us. Bugula pugeti sp. nov. 

Gemellaria loricata Linnaeus. Bugula curvirostrata sp. nov. 
Menipea ternata Ellis and Solander. Bugula longirostrata sp. nov. 

Menipea gracilis Busk. Bugula laxa sp. nov. 

Menipea occidentalis Trask. Beania mirabilis Johnston. 

Menipea occidentalis catilinensis Beania longispinosa sp. nov. 

subsp. nov. Stirparia ciliata sp. nov. 

Menipea erecta Eobertson. Stirparia occidentalis sp. nov. 

Menipea pribilofi sp. nov. Stirparia calif omica sp. nov. 

Scrupocellaria calif omica Trask. Corynoporella spinosa sp. nov. 

Scrupocellaria varians Hincks. Synnotum aviculare Pieper. 

Scrupocellaria diegensis sp. nov. Cellaria borealis Busk. 

Caberea ellisi Fleming. Cellaria mandibulata Hincks. 

Bugula neritina Linnaeus. Cellaria diffusa sp. nov. 

Bugula murrayana Johnston. Flustra lichenoides Eobertson. 

Bugula calif omica sp. nov. Flustra membranacea-truncata Smitt. 

242 University of California Publications. [Zoology 

Phylum Molluscoida Milne-Edwards. 
Class BRYOZOA Ehrenberg. 
Sub-Class Ectoprocta Nitsche. 
Colonial bryozoa with anal orifice outside the lophophore; 
a well developed introvert, and a spacious coelome. 


Polypide destitute of an epistome; lophophore circular. 

Sub-Order I. Chilostomata Busk. 

Gymnolaemata with calcareous or chitinous zooecia whose 
orifice is closed by a movable chitinous lip or "operculum"; 

ova usually matured in globular cia situated above the orifice 

of some of the zooecia; vibraeula or avicularia, or both, fre- 
quently present. 

Of the fifteen or more families included by Hincka in the 
Chilostomata, all but the firsl seven, viz.. the Membraniporidce, 
Microporidce, CribrilinidcB, Microporellidce, Porinidce, Myrio- 
zoidce, Escharidce, are omitted here. These are incrusting, or 
if erect, are for the mosl pari non-flexible in habit of growth. 
Below is given a key to the firsl seven families of the Chilostomes 
which are. for the most part, erect and flexible in habit of 

1. Colony creeping 2 

1. < lolony erect •. 3 

2. Colony Bending up ereel branches from a point just below the 

aperture Eucratiidae 

'_'. Colony imt sending up erect branches Aeteidse 

3. Colony articulated 4 

3. Colony not articulated 5 

4. Zoa-eia multiserial, arranged round an imaginary axis Cellariidse 

4. Zooecia biserial Cellulariidae 

5. Zooecia arranged back to back 6 

5. Zooecia nut hack to back 7 

6. Zocecia with avicularia Notamiidae 

6. Zocecia without avicularia Eucratiidae 

7. Zocecia biserial, avicularia pedunculated Bicellariidae 

7. Zocecia multiserial, avicularia sessile Flustridae 

Vol. 2] Robertson. — Non-Incrusting Bryozoa. 243 

Aeteidae Hincks. 

Zoarium composed of creeping branches more or less ad- 
herent to the substratum, often growing in free tufts adherent 
only part of their length. Zoozcia uniserial, arising from each 
other in a tubular prolongation of greater or less length. 
Aperture terminal, orifice at its summit. 

The definitions of family Aeteidce, of genus Aetea, and of 
species Aetea angnina are here much changed from those given 
by Hincks, whose diagnoses are based upon the erroneous notion 
which that observer entertained concerning the true nature of 
the Aetean zocecium. The adherent portion of the zocecium he 
regarded as a stolon which together with the circle of setae in 
which the tentacle-sheath terminates (Smitt, '67) seemed, he 
thought, to relate Aetea to the Ctenostomes, and to place it in 
the position of a form transitional between these and the Chilos- 
tomes. Whatever may be the relationship between these sub- 
orders, Aetea cannot be said to reveal it, since as will be shown 
in the discussion under the species A. anguina, the adherent 
"stolonic portion" so-called, is not a stolon, but an important 
part of the zooecium. 

Aetea Lamouroux. 

Aetea Lamouroux, 1812. 
Anguinaria, Johnston, 1847. 
Aetea, Hincks, 1880. 
Aetea, Jullien, 1S88. 

Zoarium, adherent to the substratum. Each zocecium par- 
tially adherent, partially erect, the erect portion carrying at its 
distal extremity the membranous aperture with its operculum. 


1. Tubular portion of zocecium ringed, upper third spatulate, bent 

A. anguina 

1. Tubular portion of zooecium not ringed, upper extremity not bent, 

but erect and truncate A. truncata 

244 University of California Publications. [Zoology 

1. Aetea anguina (Linnaeus) Lamouroux. 

PI. IV. Figs. 1, 2, 3, and 4. 
Sertidaria anguina Linn., 1758, ed. 10, p. 816. 
Cellularia anguina, Pallas, 1766, p. 78. 
Corallina anguiformis Ellis, 1767, Ger. ed., p. 50, PI. XXII, figs. 

c, C. D. 
Cellularia anguina, Ellis, 1767, p. 434, PI. 19, fig. 10. 
Cellaria anguina, Ell. and Solander, 1786, p. 26. 
Aetea anguina, Lamouroux, 1812, Vol. Ill, p. 184. 
Anguinaria spatulata, Johnston, 1847, ed. II, p. 290, PI. L, figs. 

7, 8. 
Aetea anguina, Busk, 1852, pt. I, PI. XV, fig. 1. 
Aetea anguina forma spatulata, Smitt, 1867, p. 280, PI. XVI, 

figs. 2, 4. 
Aetea anguina, Hincks, 1880, p. 4, PI. I, figs. 4, 5. 

Zoarium composed of creeping branches consisting of a 
single series of zocecia growing irregularly over a stem, frond, 
or other substratum (PI. IV. figs. 1. 2). Branches arising at 
right angles to the zooecia from each side of the creeping por- 
tion. Zooecia composed of a creeping posterior portion (fig. 2, 
ad.), and an anterior, erect, tubular portion (tu.). The pos- 
terior creeping portion of the zooecium narrow and tubular 
where it arises from the neighboring zooecium (ad.) gradually 
widening anteriorly or toward the growing point (flrr.)', then 
turning upward almost at a right angle and becoming erect 
{tu.), the remainder creeping on to give rise to a new zooecium. 
The erect portion is tubular, the lower two-thirds being minutely 
ringed, the upper one-third being somewhat inflated, and in 
many eases bent forward thus forming the so-called snake's 
head (s. //.). At the base of the tubular portion and anterior 
to it. a septum (sep.) separates the zooecium from the one next 
following. The membranous aperture {mem. ap.) is situated 
on the ventral side of the widened upper third of the erect tube 
and at its upper edge is the movable lip or operculum (op.). 
The dorsal surface of the widened upper end of the tube is 
minutely granulated. The polypide is simple in structure, very 
minute, and when retracted is drawn downward into the hori- 
zontal, adherent portion of the zooecium (fig. 3, pd.). The 
tentacle-sheath terminates above in a circle of seta? which are 

Vol. 2] Robertson. — Non-Incrusting Bryozoa. 245 

everted during the expansion of the polypide. Ocecia globular, 
membranous, situated at the upper extremity on the ventral side 
of the zooeeium just below the operculum (fig. 4, a?.). 

As Smitt ('67), Waters (79), and Jullien ('88) have 
already shown, there has been much misconception in regard to 
the zocecium of Aetea, Busk ('49) considered the erect, tubular 
portion alone to be the zocecium, and in this error was followed 
by Hincks ( '80) who regards the erect tube as the zocecium, 
the horizontal portion as a stolon, which he compares with the 
stolon of the Ctenostomes. Jullien, apparently unaware of the 
observations of Smitt or Waters on this point, criticizes the 
statement of Hincks as to the stolonic nature of the adherent 
portion. This he considers the true zocecium, since into it he 
finds the polypide withdraws. itself on retraction and on its walls 
the retractor muscle fibers are inserted ; while the upright 
tubular portion he regards a peristome. Smitt and Waters 
had previously shown, however, that the polypide inhabits the 
creeping portion, and that this is in no sense a stolon. Later, 
Waters ('96) has shown that in the species known as Aetea 
anguina forma recta, the ovary is situated in the creeping part, 
thus affording conclusive proof of the zocecial character of this 
so-called ' ' stolon. ' ' 

In the colonies of this species found on the California coast 
(figs. 1, 2), the creeping and erect portions are continuous. 
The polypide when expanded occupies the upright tube, but on 
retraction retreats into the creeping portion almost completely, 
only the tips of the tentacles and the long, delicate sheath with 
its muscles remaining in the tube (fig. 3). In this figure, part 
of the upright tube is represented as broken and the polypide 
is not in a state of complete contraction, but the insertion of 
the retractor muscles on the wall of the creeping part (mus.), 
and the presence of the ovary (ou.) on the adnate ventral wall 
are shown. From the evidence presented it seems to be estab- 
lished that the zocecium of Aetea consists of both the creeping 
and the erect portions and not merely of either one of these. 

In our specimens of Aetea the ocecia are abundant. As 
shown in fig. 4, each consists of a membranous bag situated on 

246 University of California Publications. [Zoology 

the ventral side of the zooecium below the operculum but exterior 
to the aperture, and contains an ovum in the early cleavage 
stage (a?.)- In all cases in which the ooecium is present the 
tubular part of the zooecium is distinctly curved, as shown in 
fig. 4, as if affording protection to the delicate ocecium and its 
contents. This is the first instance, so far as known, of the de- 
lineation of the ooecium of Aetea anguinaj Waters has figured 
the ocecium of the so-called variety recta in which it is on the 
dorsal side of the erect portion. The difference of location of 
the ocecia in these two forms is probably sufficient to separate 
them into distinct species. 

There is a possibility that the species found here is an un- 
deseribed one peculiar to this coast, but the ouvium of Aetea 
anguina not having been hitherto known, and no other distinc- 
tions being apparent between our form and that found in for- 
eign localities, tbis identification must stand for the present. 
The distribution of Aetea anguina is world wide, and this fact 
adds to the probability that we have here the older well known 

This species is abundant at San Pedro and San Diego, grow- 
ing over kelp, hydroid stems, shells, and other bryozoa. 

2. Aetea truncata (Landsborough) Bnsk. 

l'l. IV, figs. 5, 6. 
Anguinaria truncata Lamlsliorough, 1852, p. 228, PI. XVT, figs. 

57, 57*. 
Salpigia Hassallii Coppin, 1848, p. 273, PI. X, fig. 3. 
Aetea truncata, Busk, 1852, p. 31. 

Aetea truncata, Smitt, 1865, PI. II, figs. 5-14; PI. Ill, figs. 1-8. 
Aetea truncata, Smitt, 1867, 279 and 295, PI. XVI, fig. 1. 
Aetea truncata, Hincks, 1880, p. 8, PL I, figs. 8-11; PI. II, fig. 3. 

Zoariwm growing irregularly over the substratum. Zooecia 
rather widely separated, the posterior creeping portion fre- 
quently lengthening into a long slender fiber (PL IV, fig. 5 
ad.) ; the tubular erect portion varying in length (tu.), the 
membranous aperture (ap.) occupying on an average a little 
more than one-third of its length; slightly wider at the top 

Vol. 2] Robertson. — Non-Incrusting Brxjozoa. 247 

than at the point of union with the adherent portion ; truncate, 
granular, no part of the zocecium annulated as is the erect 
portion of the zooecium of the preceding species. Oozcia not 

No colonies of the branching form described by Hincks occur 
in our material, the simple form of Aetea truncata alone pre- 
vailing. The creeping portion of the zocecium is rather closely 
adherent to the substratum so that it is not easily removed. The 
polypide resembles that of Aetea anguina, being of simple struc- 
ture and when retracted is drawn almost completely into the 
creeping part. 

This species was obtained in considerable quantity at La 
Jolla growing over the older stems of a kelp hold-fast. 

Eucratiidae Hincks. 

Zoarium forming slender, branching, phytoid tufts. Zoozcia 
uniserial, or in two series placed back to back; expanding from 
the base upwards, with a terminal or subterminal and usually 
oblique aperture. Neither avicularian nor vibracular append- 
ages known. Oozcia globose. 


1. Colony creeping Eucratea chelata 

1. Colony erect Gemellaria loricata 

Eucratea Lamouroux. 

Cellularia (part) Pallas, 1766. 

Cellaria (part), Ellis and Solander, 1786. 

Eucratea Lamouroux, 1812. 

Catenaria (part), d'Orbigny, 1850. 

Eucratea, Johnston, 1847. 

Scruparia, Busk, 1852. 

Eucratea, Hincks, 1880. 

Zoarium composed of creeping adherent branches and of 
erect free shoots. Both creeping and erect branches composed 
of zooecia arranged uniserially, the erect branches arising from 
the ventral wall of the zocecia. Zooecia prolonged into a tube 
of greater or less length. Aperture large. 

248 University of California Publications. ■ [Zoology 

3. Eucratea chelata (Linnaeus) Lamouroux. 

PI. V, figs. 7, 8, 9. 
Sertularia chelata Linn., 1758, ed. 10, p. 816. 
Cellularia chelata, Pallas, 1766, p. 77. 

Bull's Horn Coralline Ellis, 1767, p. 42, PL XXII, figs, b, B. 
Cellaria chelata, Ellis and Solander, 1786, p. 25. 
Eucratea chelata, Johnston, 1847, ed. II, p. 288, fig. 64. 
Catena ria chelata, d'Orbigny, 1S50, Vol. V. p. 43. 
Scruparia chelata, Busk, L852, pt. I. p. 29, PI. XVII, fig. 2. 
Eucratea chelata, Smitt, 1867, pp. 281 and 301, PL XVI. figs. 7-9. 
Eucratea chelata, Bincks, 1880, p. 11, PL III, figs. 9, 10. 

Zoarium composed of creeping branches consisting of a 
single series of zocecia growing irregularly over the substratum 
and adherent to it, and of ereel branches (PI. V. bus. 7 and 8). 
Creeping branches {cr. br.) arising from the sides of the ad- 
herent zooecia; the erect, l'wi> branches (e. br.), from a point on 
the zooecia! wall just below the aperture. Zmrcia narrowed pos- 
teriorly into a tube, widening gradually anteriorly; the polypide 
even in contraction remaining in the anterior portion of the 
zooecium. Aperturt (ap.) oval, surrounded by a thin, raised. 
unarmed margin; operculum at its summit (op.). O&cia 
mitriform, somewhat pointed above, with a keel down the center, 
borne on an imperfectly developed zooecium arising just below 
the aperture (fig. !>. 03.). Tentacle-sheal h terminating above in 
a circle of set;e which are everted during expansion of the poly- 

The presence of seta' both in Eucratea and Aetea is given 
only on the authority of others. Few of our specimens of 
Eiicratm possessed ocecia. and none perfect ones. Figure 9 is 
taken from 1 [incks. 

Although not abundant at any one point, Eucratea chelata 
has been found at several localities on the coast of southern 

Gemellaria Saviguy. 

Gemellaria Savigny, 1811. 
Notamia Farre, 1837. 
Gemellaria, Hincks, 18S0. 

Zoarium erect, branching- dichotomous, each branch given 
oft' from the sides of the zooecia close to their upper extremity. 

Vol. 2] Robertson. — Non-Incr listing Bryozoa. 249 

Zoo&cia joined back to back, each pair arising from the anterior 
extremity of the preceding pair. Aperture large, sloping 
slightly upward. Ooceia? 

4. Gemellaria loricata (Linnaeus) Savigny. 

PI. V, figs. 10, 11. 
Sertularia loricata Linn., 1758, ed. 10, p. 815. 
Cellularia loricata, Pallas, 1766, p. 64. 

Coat of Mail Coralline Ellis, 1767, p. 40, PI. XXI, figs, b, B. 
Cellaria loricata, Ell. & Sol., 1786, p. 24. 

Notamia loriculata, Farre, 1837, p. 413, PI. XXVII, figs. 6-9. 
Gemellaria loriculata, Savigny, 1811, Egypt. Polyp. 
Gemellaria loriculata, Van Beneden, 1844, p. 33, PI. V, figs. 1-7. 
Gemellaria loricata, Johnston, 1847, ed. II, p. 293, PI. XLVII, 

figs. 12, 13. 
Gemellaria loriculata, d'Orbigny, 1850, p. 46. 
Gemellaria loricata, Busk, 1852, pt. I, p. 34, PL XLV, figs. 5, 6. 
Gemellaria loricata, Smitt, 1867, pp. 286 and 324, PI. XVII, fig. 54. 
Gemellaria loricata, Hincks, 1880, p. 18, PI. Ill, figs. 1-4. 

Zoarium forming densely bushy masses, 50 or 75 mm. in 
height, made up of numerous long, straight branches; branching 
dichotomous (PL V, fig. 10). Zooscia narrowed below, widen- 
ing above; aperture (fig. 11, op.) oval, occupying about half the 
front, sloping somewhat toward the top, with a thin, raised, un- 
armed margin; operculum (op.) semicircular, at the anterior 
edge of the aperture. Colony attached by numerous rootlets. 
Main stem of colony thicker below. 

This species is found only on the northern shores. It is 
abundant at Orca, Prince Williams Sound, where it was taken 
from shore rocks at low tide; at Juneau, it was dredged at 20 
fathoms. Hincks reports it from Queen Charlotte Islands. 

Cellulariidae Bnsk. 

EscharidoB (part) Johnston, 1847. 
Cabereadce Busk, 1852. 
Cellulariadce Busk, 1852. 
Cellularia? Smitt, 1867. 
Cellulariidm, Hincks, 1880. 

Zoarium erect, branching dichotomous. Zocecia in two or 
more series, closely united in the same plane ; avicularia and 

250 University of California Publications. [Zoology 

vibracula, or avicularia only, almost universally present, sessile. 
This is a well marked group all of whose members have a 
strong family resemblance. The walls of the zooecia are strongly 
calcified, being generally variously adorned with spines and 
sessile avicularia, and many with waving vibracula. In all the 
embryos mature in globose, hood-like ocecia. This family is 
well represented in our fauna, there being at least six species of 
Menipea, four of which are new; and three of Scrupocellaria, all 
of which are peculiar to this region. Cabcrea is represented by 
but one species, the northern C. ellisi. 


1. Zooecia three or more in an internode without vibracula Menipea 

1. Zooecia many in an internode, vibracula present 2 

2. Vibracular chamber not covering dorsal surface of zocecium 


2. Vibracular chamber directed obliquely and covering the dorsal sur- 
face of the zocecium Caberea 

Menipea Lamouroux. 

PI. V, fig. 16. 
Menipea Lamouroux, 1816. 

Cellularia, Johnston, 1847. 
Cellarina, Van Beneden, 1849. 
Emma, Husk, 1852. 
Menipea, Busk, 1852. 
Cellularia, Smitt, 1867. 
.1/- uijH a, lliiicks, 1880. 

Zooecia oblong, widest above, attenuated below, often elon- 
gated downward; lateral avicularia generally present, sometimes 
wauling; frontal avicularia generally present on the zoopcium 
at the bifurcation of a branch. Xo vibracula. Oacia present; 
large, globose. Root fibers characteristically located, being found 
in two positions: first, on the front, or on the lateral wall of a 
zocecium just below or to one side of the aperture (fig. 16, r. d. 
and r.) : second, in a definite dorso-lateral chamber situated just 
above the lateral avicularium (r. ch. and u. r.). In certain mem- 
bers of this genus, different functions are performed by these 
differently located fibers. Those found to one side, or below the 
aperture invariably turn downward and serve to anchor the 

Vol. 2] Robertson. — Non-Incru sting Bryozoa. 251 

colony (r.) ; those arising: in the dorso-lateral chambers may 
assist in anchoring the colony when they occur on zooecia in the 
lower part of the colony; usually, however, they turn upward, 
and twining about the other portions of the colony, function as 
tendrils (u. r.). 

Waters ('96) calls attention to the diagnostic importance of 
the root fibres, their position being fairly constant so that it may 
be used as a specific character. Of the Menipeas of this region, 
M. ternata and M. gracilis have root chambers in identical posi- 
tions and root fibers performing identical functions. The other 
three species differ from these two and from each other in the 
number and shape of the root chambers and in the function of 
the root fibers arising from them. 


1. Zooecia commonly three in an internode 2 

1. Zocecia many in an internode M. erecta 

2. Spines two or three 3 

2. Spine more than two or three 5 

3. Scutum half way down on the margin of the aperture 4 

3. Scutum two-thirds of the way down on the margin of the aperture 6 

4. Scutum small, spine-like M. ternata 

4. Scutum large, flabellate M. gracilis 

5. Scutum simple 6 

5. Scutum divided 7 

6. Scutum half way down on the margin of the aperture M. pribilofi 

6. Scutum two-thirds of the way down on the margin of the aperture 

M. occidentalis 

7. Scutum two-thirds of the way down on the margin of the aperture 

M. occidentalis subspecies catalinensis 

5. Menipea ternata (Ellis and Solander) Busk. 

PI. V, figs. 12, 13, 14, 15, 16; PI. VI, fig. 17. 
Cellaria ternata Ell. and Sol., 1786, p. 30. 
Celhdaria ternata, Johnston, 1847, p. 335, PI. LIX, figs. 1, 2. 
Cellarina gracilis, Van Beneden, 1849, p. 67, PI. X, figs. 1, 2. 
Menipea ternata, Busk, 1852, pt. I, p. 21, PI. XX, figs. 3-5. 
Cellularia ternata, Smitt, 1867, p. 282, PI. XVI, figs. 10-26. 
Menipea ternata, Hincks, 1880, p. 26, PI. VI, figs. 1-4. 

Zoarium consisting- of loosely spreading, straggling branches, 
or sometimes of rather large dense masses, frequently 30 

252 University of California Publications. [Zoology 

to 35 mm. high and from 60 to 80 mm. broad (PI. V, 
figs. 12, 13). Branching dichotomous. Intemodes consisting of 
three zooecia, as a rule, although there are many instances of 
five or seven zooecia to an intemode. The longer internodes are 
generally terminal and bear the ovicells. Joints light colored, 
arising in distinct tubes (fig. 14, ./'.). Zooecia elongated, narrowed 
below; aperture oval (op.), occupying less than half the front. 
with two or three spines, one on the inner margin just above the 
operculum, and one or two on the upper, outer margin; zocecium 
at bifurcation of branch having sometimes one, sometimes two 
cither long spines ;it its summit. Towards extremity of branches 
the spines increase in length, often very considerably. Scutum 
varying in shape from a mere spine to a somewhat broad flabel- 
late process (.st.). Lateral avicularia (av.) Large, and promi- 
uent, generally developed on all the zooecia of an internode 
excepl the one a1 the bifurcation of a branch; sometimes, how- 
ever, missing. Frontal avicularia (fr. av.) on the zocecium 
at bifurcation only; small, raised, with mandible directed for- 
ward, often obliquely to right or left. Ocecia smooth, somewhat 
globose, reaching not quite to the lower margin of the next upper 
zooecium {oe.). From a flat disk in front or to one side of the 
aperture of many of the lower zooecia in a colony, root fibers (r.) 
extend downward, assisting in anchoring the colony to the sub- 
stratum; from zooecia higher in the colony other upward ex- 
tending fibres arise in a dorso-lateral chamber (r. ch.) which is 
slightly raised and which projects from the zooecia! wall. 

M. ternata is widely distributed, being found on Channel 
Rocks, and San Juan Island in Puget Sound: and at Dillons 
Beach, Lime Point. Mendocino, and Lands End on the coast of 
California. It is also reported by Hincks from Queen Charlotte 

In the species from these different localities considerable 
variation occurs. As a rule, the colonies from California grow 
in more compact masses, and the triads of zooecia forming the 
internodes are shorter than are those of the Puget Sound species. 
Thus the length of an internode in the Puget Sound species 
varies between 1,100 and 1,500 /x, and that of the species found 

Vol. 2] Robertson. — Non-Incru sting Bryozoa. 253 

farther south, slightly less. In the main points, however, the 
Menipeas from these various localities strongly resemble each 
other and in no points more constantly than in the position and 
form of the chambers from which the root and climbing fibres 

6. Menipea gracilis Busk. 

PI. VI, figs. 18, 19, 20, 21. 
Menipea gracilis Busk, 1881. 
Menipea ternata, forma gracilis, Smitt, 1867. 

Zoarium forming a loose, tangled mass due to the great num- 
ber of tendril-like fibres which twine around neighboring 
branches. (PI. VI, fig. 18.) Branching dichotomous; internodes 
consisting of three zocecia, except terminal internodes and those 
possessing ovicells, which may have five, seven, or nine zooeeia; 
internodes of three zooeeia only, often very long, ranging from 
1,600 to 1,900 [x in length. Zocecia enlongated, aperture large, 
(fig. 19). Zooeeia with two or three spines, depending upon 
the presence or absence of the lateral avieularia. If the latter 
are absent there will usually be two spines on the outer margin ; 
if present, then but one (sp.). There is always a spine on the 
inner margin of the aperture just above the scutum. Zooecium 
at bifurcation possesses one or two spines, more often one, at 
its summit. Sent ion (sc) large, flabellate, projecting outward 
and arching over the aperture in well developed specimens, pro- 
jecting beyond the outer edge of the zooecium. Lateral avieularia 
frequently absent, when present often minute. Frontal avi- 
eularia (fr. av.) occur on the zooecium at the bifurcation of tin- 
branch, but these too, are frequently absent; on ooecial inter- 
nodes,. however, they are often found on each zooecium, situated 
a little to one side of the aperture and so close to the ooecium 
as to seem perched upon its upper margin (fig. 20, av.). 
Omcia (oe.) high, rounded, with striations radiating from a 
thin place at the base of the ocecial wall. Rootlets originating 
at two places. Those which anchor the colony springing, as a 
rule, from a disk to one side and below the aperture (fig. 19. 
r. d.) ; the others, more tendril-like, and for the most part ex- 

254 University of California Publications. [Zoology 

tending: upward, arising from a definite chamber just above the 
lateral avicularium, the chamber projecting somewhat and being 
not quite circular especially as seen from the dorsal side (fig. 
21, r. ch.). 

This species is abundant at Orca, Prince Williams Sound; 
reported by Hincks from Cumshewa Harbor, Queen Charlotte 

7. Menipea occidentalis Trask. 

PI. VI, figs. 22, 23, 24, 25. 
Menipea occidentalis Trask, 1857, p. 113, PI. 4, fig. 4. 
Menipea compacta Hincks, 1882, vol. 10, p. 461. 
Menipea compacta Bincks, 1884, vol. i: 1 .. p. 208, PI. IX. fig. v - 

Zoarium forming bushy tnfts from 15 to 50 mm. in height, 
attached by a Large number of root fibres. Root fibres not de- 
veloped throughoul the colony, this species being in no sense a 
• limber. Branching extremely regular and characteristic (fig. 
22), each tuft or frond consisting of a main rib or primary 
branch (pr. br.) from which secondary branches arise alter- 
nately {sec. br.), these again giving off tertiary branches (ter. 
br.). Intern-odes consisting of three zooecia (fig. 23). Joints (j.) 
yellow or brown arising from definite chambers on each side of 
the most anterior zooecium of an internode. Zooecia elongated, 
narrowed below: apertun {ap.) occupying about half the front. 
surrounded by six jointed spines, sometimes by five or seven; 
two arising on the margin of the upper half of the aperture 
opposite each other and meeting across it (c. sp.) ; two other 
Longer ones extending from the upper margin of the aperture 
sp.), and between these two and the two crossed spines, two 
oilier opposite flaring spines (fl. sp.) ; on the terminal zooecia 
these flaring spines often grow very long so that each zooecium 
is then bordered anteriorly with a bristling array of four long 
spines. Scutum [sc.) sometimes a mere spine, sometimes broader 
than a spine, arising on the lower half of the inner margin of 
the aperture. Avicularia large, lateral, sessile, found typically 
on all the zooecia except the one at the bifurcation of a branch. 
Frontal avicularia wanting. Ocecia usually developed on the 
tertiary branches, large, globose, covering the zocecial wall below 

Vol. 2] Robertson. — Non-Incrusting Bryozoa. 255 

the apertures of the zocecia just above (fig:. 24, oe.), their front 
walls perforated by a small number of pores. Rootlets of one 
sort only, extending downward, serving to anchor the colony; 
arising in root chambers just above the lateral avicularia (fig. 25, 
r. ch.) ; the root chambers situated on the lower zooecia only, 
and never appearing unless a root fibre is developed ; rounded, 
projecting dorsally and laterally, the root fibre springing from 
the lower side and extending downward keeping close to the 
branch and running parallel with the other root fibres of the 
branch, thus forming a sort of cable or rope for the attachment 
of the colony. 

This species was first described by Dr. John Trask ('57), 
but his description was apparently unknown to Hincks ('82), 
who redescribed it under the name of M. compacta. It is per- 
haps the most abundant species of bryozoa on the shores of San 
Francisco Bay and Golden Gate Straits. It grows between tide 
marks on rocks and sea weed forming bushy tufts of a white, 
yellowish or dirty grey color, depending on its age. It is quite 
easily recognized by its excessive spininess. It is found from 
Queen Charlotte Islands to San Diego, but is most abundant 
above Point Concepcion. South of that point it is noticeably 
scarce on the shore and in dredgings, and the colonies are small 
and delicate. 

8. Menipea occidentalis catalinensis. Subsp. nov. 

PI. VII, figs. 26, 27. 

South of Point Concepcion, notably at Santa Catalina and 
San Pedro, M. occidentalis shows considerable variation. The 
internodes more often consist of five or seven zocecia, and 
the scutum which in the type form is a mere spine, or at most 
is slightly flabellate, in the subspecies catalinensis is decidedly 
fan-shaped, the edge being divided and extended into five, six, 
or more spinous processes, making it so large as to cover the 
lower half of the aperture (fig. 26, sc). One or both of the 
spines that meet over the upper part of the aperture may be 
bifid (c. sp.). The bifid spines and the large and much divided 
scutum are very noticeable features and constitute the chief 

256 University of California Publications. [Zoology 

differences between the subspecies catalinensis, and the typical 
M. occidentalis. While these variations from the type occur 
rather constantly in specimens from the south, affecting as they 
do such variable appendages as spines and scuta, they are not 
considered of sufficient importance to establish a new species. 
The drawing- (fig. 26) gives the impression, unfortunately, that 
the individual zooecia of the subspecies catalinensis arc larger 
than those of the type M. occidt ntalis, but it was unintentionally 
made on a larger scale of magnification than was that of figure 23. 

9. Menipea erecta Robertson. 

PI. VII. figs. 28, 29, 30, 31. 

Menipea erecta Robertson, 1900, p. 317, PI. XIX. figs. 1. 2. 
Scrupocellaria scabra, Robertson, 1900, p. 318, PL XIX, figs. 3, 4. 

Zoarium composed of numerous stiff, dichotomously divided 
branches from 20 to 25 mm. in Length (fig. 28). Inter- 
nodes consisting of three, live, or seven zooecia, the number in- 
creasing toward the extremities of the branches: articulations 
extending through the zooecia just below the aperture, some- 
times including its Lower border (fig. 29, art.). Zooecia biserial, 
alternate, narrowed below ; apt Hun oblong, occupying more than 
half the front: margin raised, crenulate, with one or two blunt 
-pirns at the upper outer angle; senium (sc.) a flattened spine. 
sometimes growing broad and bifid. Lal< nil avicularia fre- 
quently wanting (fig. 301, sometimes feebly developed (fig. 31), 
and again rather large (fig. 29). When avicularia are present 
there is usually bul one spine at the upper, outer angle, although 
this is not the invariable rule. Frontal avicularia (fr. or.) gen- 
erally present on each zocecium, though sometimes lacking; when 
present, they are situated to one side of the aperture, at the 
base of the scutum of the adjacent zocecium. Ocecia large, glo- 
bose, more or less striated. Root fibres mainly upon the lower 
zooecia. springing directly from the zooecial wall and extending 
downward (fig. 30, v.). 

Examination of material from several localities, especially 
of that from Puget Sound, leads me to unite the species for- 

Vol. 2] Robertson. — Non-Incr listing Bryozoa. 257 

merly identified as Scrupocellaria scabra Van Beneden with 
Mcnipea erecta Kobertson ( '00). 

This species is obtained from three localities, Kadiak and 
Sitka, Alaska, and San Juan Island, Puget Sound. The range 
of variation is considerable, but is not greater than frequently 
occurs in a species found in localities so remote, and consists 
mainly in the presence or absence of spines or avicularia. The 
specimens from Sitka, for example (fig. 30), show irregular 
development of frontal avicularia with entire absence of lateral 
avicularia. The colonies from Kadiak (fig. 29) show frequent 
absence of lateral avicularia and their replacement with spines; 
while a few zooecia from San Juan specimens (fig. 31) show an 
entire absence of spines and considerable variation in the size 
of the lateral avicularia. 

10. Menipea pribilofi sp. no v. 

PI. VII, figs. 32, 33; PI. VIII, fig. 34. 

Zoarium forming a compact mass 25-50 mm. high, attached 
by a large number of root fibres; the upward tendril-like fibres 
being very slightly developed (PI. VII, fig. 32.). Branching 
dichotoinous, main branches long, possessing a number of shorter 
inward curving secondary branches. Internodes consisting of 
three zooecia except the ooecial internodes which generally con- 
sist of five ; separated by dark colored chitinous joints. Zooecia 
relatively short and stout, broad at the top, attenuated below, 
aperture occupying less than half the front surface, with three 
spines on its upper margin ; the bifurcating zooecium may have 
but two spines, but it is frequently found bristling with three 
or four. Scutum (PI. VIII, fig. 34, sc.) simple, often a mere 
spinous process ; when better developed, it broadens at the free 
extremity. Spines and scuta distinctly jointed. Lateral avi- 
cularia generally present, large (lat. av.). Frontal avicularia 
(/>. av.) few, when present found only on the zooecium at the 
bifurcation of a branch, large, raised, beak set obliquely, some- 
times transversely to the length of the zooecium. Ocecia globose, 
smooth (PI. VII, fig. 33, oe.). Rootlets in two positions. Those 

258 University of California Publications. [Zoology 

lower in the colony develop on the front surface of the lower 
zocecia, below and to one side of the aperture. These rootlets 
invariably proceed straight downwards (PI. VIII, fig. 34, r.). 
Often on a zooeeinm in the same internode, and on zocecia at 
higher levels, just above the lateral avieularia, other rootlets 
curving upward, develop from a circular chamber slightly raised 
above the zocecial wall (u. r.). These upward growing fibres are 
of much less frequent occurrence than in .1/. tcrnata and M. 
gracilis, and the chambers from which they proceed are of much 
simpler construction. On the upper half of the colony neither 
fibres nor chambers are found. 

In the habit of growth, greater compactness, greater develop- 
ment of spines, and scarcity of tendril-like fibres, this species 
resembles .1/. occidentalis rather than M. ternata. 

Menipea pnbilofi is known only from the shores of Alaska, 
and the islands of Bering Sea. having been obtained in consid- 
erable quantity from St. Paul Island. Pribilof Islands; and in 
small quantity from Homer, Qnalaska, and Yakutat, Alaska. 

Scrupocellaria Van Beneden. 

Cellularia Pallas, 1766. 
Cellaria, Ell. and Sol., 1786. 
Scrupocellaria Van Beneden, 1844. 
Cellularia, Johnston, 1847. 
Cauda, Busk, 1852. 
Sffii/ioi-t llaria, Hiiicks. 

Zoarium jointed. Zocecia numerous in each internode. rhom- 
boid; apertun with or without scutum; a sessile avicularium at 
the upper, outer, lateral angle: a vibraculum at the lower outer 
angle, and generally a sessible avicularium on the front surface 
of each zocecium. 


1. A'ibracular chamber on every zocecium 2 

1. A'ibracular chamber not on every zocecium 3 

2. Vibraculum slightly longer than a zocecium S. varians 

2. Vibraculum as long as three zocecia S. diegensis 

3. Vibraculum not as long as a zocecium S. calif ornica 

Vol. 2] Robertson. — Non-Incr listing Brijozoa. 259 

11. Scrupocellaria calif ornica Trask. 

PI. VIII, figs. 35, 36, 36a, 37. 
Scrupocellaria calif ornica Trask, 1857, p. 114, PI. 4, fig. 2. 
Scrupocellaria brevisetis (?) Hincks, 1882, p. 462. 

Zoarium growing in large compact tufts, somewhat coarse in 
appearance. (PL VIII, fig. 36a.) Branching dichotomous, inter- 
nodes consisting of five, seven, or nine zooecia. Zooecia slightly 
attenuated below, aperture occupying more than half the front 
(fig. 36). Two, often three spines on the upper, outer margin 
and one on the inner just above the scutum. Zooecium at the 
bifurcation (zoe,. hi.) of a branch with one short spine at its apex; 
two spines below on one side and one spine on the other just 
above the scutum. Scutum (sc.) often spine-like, sometimes 
broadened below, and always curved with a downward slope. 
Lateral avicularia generally present, when absent, an extra spine 
may develop in that place, making the third on the outer edge 
of the margin (sp.). Avicularia vary in size, but unlike those 
of the following species, the fluctuation in size lies in the greater 
or less relative proportion of the muscular part (lat. av.). Fron- 
tal avicularia (fr. av.) present on each zooecium just below the 
aperture. Vibracular cell frequently lacking, and not visible 
from the front surface except on the zooecium at the bifurcation 
of a branch (v. c). Vibracula, when present, found only on 
the zooecia of the lower part of the colony where root fibres are 
given off (r.). This is usually true except of the zooecium at 
the bifurcation of a branch where vibracular cells are generally 
present irrespective of the formation of a root fibre. Vibrac il- 
ium shorter than a zooecium; vibracular chamber much like a lat- 
eral avicularium in form (fig. 35, v. ch.). The groove, lying in 
the part corresponding to the lower mandible of an avicularium, 
extending transversely across the dorsal surface of a zooecium 
(gr.), the short stout vibraculum much like the elongated upper 
mandible of an avicularium. Ocecia sparingly developed, found 
only on internodes at the extremity of the branches, three or 
four in an internode, smooth (fig. 37, oe.). Rootlets abundant 
on the lower internodes of a colony where vibracular cells abound 
with their short vibracula (fig. 36, r.). 

260 University of California Publications. [Zoology 

This species first figured and described by Trask ('57) from 
San Francisco Bay is probably the S. brevisctis, Hincks ('82) 
from Queen Charlotte Islands, of which only a short description 
without figure is given. Trask found neither vibracula nor 
vibracular chambers, but they are abundant, as I have observed, 
on the lower part of the colony, while almost entirely lacking on 
the upper portion. 

S. califomica is distributed at various points along the coast 
of California; it grows luxuriantly at Dillons Beach and at 
many places on San Francisco Bay, and occurs in small quantity 
below Point Concepcion. 

12, Scrupocellaria varians Eincks. 

PL VIM. Bgs. 38, 39; PL XVI. fig. 95. 
Scrupocellaria varians Eincks, 1882, p. 4<n. PL XIX. figs. 1-lc. 

Zoarium forming bushy tufts 12 25 nun. in height. (PI. 
XVI, fig. 95.) Branching dichotomous, internodes consist- 
ing of a variable Dumber of zocecia, those in the Lower part of 
the colony containing usually five or seven zocecia, those at a 
higher level, nine or eleven. Joints yellow. Zocecia biserial, 
alternate, slightly narrower below; aperture oval, occupying 
more than half the front ; scutum sometimes a mere spine, some- 
times bifid or trifid, usually inclined downward (PI. VIII. fig. 
38 : two spines on the upper margin, one of them just above the 
scutum, the other opposite. Both lateral and frontal avicularia 
developed on each zocecium. Lateral avicularia of different pro- 
portions, the greater number being much elongated, extending 
upward and outward, with a long, thin, pointed, curved beak 
(c. ar.) ; frontal avicularia just below or slightly to one side of 
the aperture; sessile, raised, with mandible directed transversely. 
Vibracular chamber triangular, the apex visible from the front 
(fig. 39, v. ch.) : vibraculum (v.) longer than a zocecium, and 
the groove into which it falls, extending transversely across the 
zocecium {gr.). Ocecia (fig. 38, oe.), smooth, developed on each 
zocecium of the terminal internodes. Rootlets springing from the 
base of the vibracular chamber (fig. 39, r.). 

Vol. 2] Robertson. — Non-Incr listing Bryozoa. 261 

Perhaps the most striking feature of this species is the pecu- 
liar size and prominence of many of the lateral avicularia. This 
is brought about by the great extension of the beak and is not due 
to increase of the muscular portion (mns.) ; thus, in fig. 39 (c. av.) 
the mandible (man.) is seen to be longer than the muscular 
part, and to be more than half as long as the whole appendage. 
In the shape of its vibracular chamber and in the unusually 
large size of its lateral avicularia, this species resembles S. eali- 
fornica, but as has been said, the increase of size of the avicularia 
in the two cases is due to increase of different parts of the organ. 
In other respects, S. varians and S. californica are very unlike, 
e.g., in habit, in shape of scuta, and more especially in abund- 
ance of the vibracular chambers and length of the vibracula. 

Although this species has been obtained in small quantities on 
the California coast as far south as San Pedro and La Jolla, it 
is characteristically a more northern form. It was first described 
by Hincks from Queen Charlotte Islands, and it is abundant 
in Puget Sound, both on Channel Rocks, and at San Juan. 
Material obtained in the south was dredged at 32 fathoms, while 
in Puget Sound this species is a shore form. 

13. Scrupocellaria diegensis, sp. nov. 

PI. IX, figs. 41, 42, 43, 44; PI. XVI, fig. 96. 

Zoarium forming a coarse bushy mass often 50 mm. in 
height (PI. XVI, fig. 96). Branching dichotomous, internodes 
consisting of a variable number of zooecia, nine ; thirteen, 
seventeen or more, especially in the terminal internodes. Joints 
somewhat inconspicuous, often occurring as high on the zooecium 
as the lower margin of the aperture. Zooecia biserial, the two 
rows inclined at an angle so that the internode is keeled, and 
in cross section is triangular; rather short and broad, slightly 
narrowed below (PL IX, fig. 41) ; aperture oval, with a wide 
margin, and occupying more than half the front surface. Scu- 
tum (sc.) in older zooecia large, covering more than half the 
aperture, thickened on the inner surface and raised on a ped- 
uncle. Spines three, four, or five, the one immediately above 
the scutum, and frequently the one opposite, bifid (bi. sp.). In 

262 University of California Publications. [Zoology 

well developed zocecia there are three spines on the outer mar- 
pin, the lower of these frequently bifid, the second often quite 
long, and the third, or uppermost one shorter and sometimes 
missing; two spines on the inner margin, the lower generally 
bifid, the upper frequently lacking or broken. Lateral avicuJaria 
of moderate size, usually found on each zocecium. A series of 
sessile, frontal avicularia (fr. a v.) extends between the zooeeia, 
each avicularium situated usually at the base of the spine on 
the inner margin nearest the scutum. These frequently stand 
out prominently, the beak opening upward, so that the zocecia 
frequently have the appearance of being flanked on each side 
by an avicularium, the one on the inner side being slightly 
smaller than that on the outer. The zooeciun) at the bifurcation 
of a branch usually bears a raised avicularium of extraordinary 
size, with elongated beak directed obliquely, sometimes to the 
right, sometimes to tin 1 left (hi. av.). Vibracular chambi r dorsal. 
Large, its length equalling half that of the zocecium on which it 
is placed (fip. 42. v. ch.) ; vibraculum long, two and a half times 
as long as the individual zooeeia (fig. 41. v.). The rootlet arises 
from near the base of the vibracular chamber, toward the outer 
side (figs. 42 and 43. r.) The rootlets are developed only on 
the lower zocecia of a colony: in the upper xoovia the place of 
the rootlet is marked by a pore (fig. 42, p.) Ooecia numerous, 
large, reaching to the margin of the aperture of the zocecium 
next above, and sometimes covering it (fig. 41, oe.) ; somewhat 
flattened and marked by numerous pores. 

This species grows in large bunches, and being very calcar- 
eous, and having many long vibracula. spines and other appen- 
dages which catch and hold debris, it is remarkably coarse and 
dirty in appearance and rough to the touch. It strongly re- 
sembles Cdberea, and the vibracular cells while not as large as 
those of that genus, yet show affinities to them in size, and in 
the length and slope of the groove. This species likewise shows 
certain resemblances to S. cervicornis, Busk ('52), but its habit 
is very different and it does not possess the peculiar scutum for 
which the latter is named. 

8. diegensis is extremely abundant at San Diego on the rocks 
at Ballast Point, on floats at Coronado, and wherever it can get 

Vol. 2] Robertson. — Non-Incrusting Bryozoa. 263 

a foothold. It seems to be almost the most abundant bryozoan 
of the region ; also found at many points in the vicinity of San 
Pedro. Less abundant northward, being found only in small 
quantities in San Francisco Bay and not obtained north of this 


Caberea Lamouroux. 

Cdberea Lamouroux, 1816. 
Flustra, (part), Johnston, 1847. 
Cellularia (part), Johnston, 1847. 
Caberea, Busk, 1852. 
Cauda, d'Orbigny, 1850. 
Caberea, Smitt, 1867. 
Caberea, Hincks, 1880. 

Zoarium not articulated. Zocecia in two or more series, sub- 
quadrangular, or ovate, with a very large aperture. Sessile, 
frontal avicularia on the side and front of zocecia; lateral avicu- 
laria minute. Vibracular cells very large, placed in two rows, 
stretching obliquely downwards across the back of the zocecia, 
which they almost cover, to the median line, notched above and 
traversed through a great portion of their length by a shallow 
groove. Vibracula usually toothed on one side. 

This genus is easily distinguished from others of this family 
by the peculiar shape and size of the vibracular cell as shown 
in fig. 46. It is not common in our collections, being repre- 
sented by only a small part of a colony of one species. 

14. Caberea ellisi Fleming. 

PL VIII, fig. 40; PI. IX, figs. 45, 46. 
Flustra ellisii Fleming, Mem. Wern. Soe. ii, p. 251, PL 17, figs. 1-3. 
Flustra setacea Johnston, 1847, ed. II, p. 346. 
Cellularia hoolceri, Busk, 1852, pt. I, p. 37, PL XXXVIII, fig. 2. 
Caberea ellisii, Smitt, 1867, pp. 287 and 327, PL XVII, figs. 55, 56. 
Caberea ellisii, Hincks, 1880, p. 59, PL VIII, figs. 6-8. 

Zoarium fan-shaped, dichotomously branched ; branches thick, 
widening upward (PL VIII, fig. 40). Zocecia in two or four rows, 
short, subquadrangular (PI. IX, fig. 45) ; aperture elliptical, 
occupying nearly the whole of the front, with a broad minutely 
granular margin, sloping outwards ; marginal zocecia with two 
stout spines above on the outer side, and one on the inner ; 
intermediate zocecia with one spine on each side. Lateral avicu- 

264 University of California Publications. [Zoology 

laria (lot: av.) small and inconspicuous, placed a little below 
the top of the zocecium, with a rounded mandible: frontal avicu- 
lana (fr. av.) raised, two below the aperture, placed one on 
each side, or sometimes only one; mandible rounded, directed 
downwards. Vib racula very long, serrate (v.). Ocecia flattened, 
frequently with a depressed, smooth, semicircular space in front 
from which fine striae radiate to the margin. Rootlets arise 
from the side of the vibracnlar chamber, the two on opposite 
sides uniting in the median dorsal line with each other and with 
tho^e below, forming a thick keel-like cable along the dorsal sur- 
face of the branches (fig. 46, r.). At the base of the colony 
the threads become free and attach themselves independently to 
particles of sand or to whatever forms the substratum. 

A small quantity of C. ellisi was dredged in 20 fathoms at 
Juneau. It is reported also from Queen Charlotte Islands. 

Bicellariidae Busk. 

Bit i llariada Busk, 1852. 
Bit > llarit ce Smitt, 1867. 
Bict llariidce, II incks, L880. 

Zoarium erect, with or withoul articulated stem or peduncle, 
or composed of a number of zooecia connected by tubular pro- 
cesses. Zoo rin rather loosely united in one. two, or more series, 
or disjunct, boat-shaped or sub-tubular; aperturt occupying a 
Large proportion of the front. Avicularia, when present, capi- 
tate, pedunculated, and articulated. Ocecia either pedunculated 

Or Sessile. 

The diagnosis of the family Bicellariida as given by II incks 
is here revised in order to include the genus Stirparia Goldstein. 
This genus is fairly abundant on this coast, and unlike the other 
members of the family possesses a jointed stem of varying length 
surmounted by delicate feathery tufts of zooecia. 


1. Colony erect - 

1. Colony creeping 5 

2. Colony unstalked 3 

2. Colony stalked 4 

3. Zocecia uniserial Corynoporella 

3. Zocecia bi- or multiserial Bugnla 

4. Zocecia biserial Stirparia 

5. Zooecia uni- or multiserial, with tubular connections Beania 

Vol. 2] Robertson. — Non-Incr listing Bryozoa. 265 

Bugula Oken. 

Cellularia Pallas, 1766. 
Bugula Oken, 1815. 
Cellularia, Johnston, 1847. 
Avicella Van Beneden, 1848. 
Avicella Van Beneden, 1849. 
Ornithopora d'Orbigny, 1851. 
Omithoporina d'Orbigny, 1851. 
Bugula, Busk, 1852. 
Bugula, Hincks, 1880. 

Zoarium erect, branched. Zocecia boat-shaped, or sub-quad- 
rangular, elongate, united in two or more series ; aperture occu- 
pying a large proportion, sometimes the whole of the front, not 
turned upward or oblique. Avicularia in the form of a bird's 
head, and jointed, usually one to each zocecium. Omcia sessile 
at summit of aperture, or pedunculate upon the side of the 

This genus is abundant in this region, not only in the number 
of species but also in the luxuriance of growth, and is remark- 
able for the large size which some of the species attain. Nine 
species are found, all of which are new except three; one, Bugula 
murrayana, is circumpolar, and another, B. neritina, is cosmo- 
politan in range. 


1. Zooecia biserial 2 

1. Zooecia multiserial 4 

2. Avicularia, if present, at summit of aperture B. neritina 

2. Avicularia at base of aperture B. pacinca 

2. Avicularia between summit and base of aperture 3 

3. Avicularia much elongated, half as long as zocecium, slightly more 

than half way between summit and base of aperture 

B. longirostrata 

3. Avicularia not elongated, less than half the length of a zocecium, just 

half way between base and summit of aperture B. californica 

4. Avicularia present _ 5 

4. Avicularia absent B. laxa 

5. Avicularia at base of aperture B. murrayana 

5. Avicularia between base and summit of aperture 6 

6. Avicularia less than half way from base and summit of aperture 

B. fiabellata 

6. Avicularia near base of aperture, beak much curved 

B. curvirostrata 

266 University of California Publications. [Zoology 

15. Bugula neritina (Linnaeus) Oken. 

PI. IX, fig. 47; PI. XVI, fig. 97. 
Sertularia neritina Linnfeus, 1758, ed. X, p. 38. 
Cellularia neritina, Pallas, 1766, p. 67. 

Snail-bearing Coralline Ellis, 1767, p. 40, PL 19, figs, a, A. 
Cellaria neritina, Ell. and Sol., 1786, p. 22. 
Bugula neritina, Oken, 1815. Ab. 2, p. 89. 
Acamardhis neritina, Lamouroux, 1816, p. 58, PL 3, fig. 2. 
Cellularia neritina, Johnston, 1847, p. 340, PL 60, figs. 3, 4. 
Acamarchis neritina, d'Orbigny, 1850-52, p. 324. 
Bugula neritina, Bnsk, 1852, p. 44, PL XLIII, figs. 1-6. 
Bugula neritina, Waters, 1887, p. 91, PL IV, figs. 3, 15. 

Zoarium consisting of large trashy tufts 75 to 100 mm. in 
height, brown or reddish brown, often tinged with purple. 
( PI. XVI, fig. 97.) Branching dichotomous. Zo&cia biserial, 
quadrangular, truncate above; apertux occupying more than 
two-thirds of the front; a short spine or denticle ;it the summit 
of the sides of the zocecium (PI. IX. fig. 47. den.). Ocecia very 
conspicuous and numerous (oe.) • globose, attached to the inner 
anterior angle of the zooecium by a short peduncle. Avicularia 
none. Bootlets forming a tuft at the base of the colony. 

This species is extremely abundant, especially on the southern 
coast of California. It is not only found on rocks, floats, kelp, 
etc., along the shore, but it is frequently taken by the dredge. 
and great brown masses are constantly cast on the beach by the 
waves. Its most northern limit is Monterey Bay, where it is 
obtained in small quantities. 

16. Bugula murrayana (Johnston) Busk. 

PL X, fig. 48 ; PL XVI, figs. 98, 99. 
? Eschara hispida Pallas, 1766, p. 49. 
Flustra murrayana Johnston, 1847, ed. II, p. 347, PL LXIIT, 

figs. 5, 6. 
Avicella multispina Van Beneden, 1848, Vol. 15, pt. I, p. 76, figs. 

? Ornithopora dilatata d'Orbigny, 1850-52, p. 323. 
Bugula murrayana, Busk, 1852, p. 46, PL LIX, figs. 1, 2. 
Bugula murrayana, Smitt, 1867, p. 291, PL XVIII, figs. 19-27. 
Bugula murrayana, Hincks, 1880, p. 92, PL XIV, figs. 2-9. 

Zoarium composed of bushy tufts from 25-50 mm. in height, 
sometimes consisting of masses of Flustra-like fronds 50 or 

Vol. 2] Robertson. — Non-Incr listing Bryozoa. 267 

more mm. in diameter (PL XVI, fig. 98), or of much elon- 
gated, narrow, strap-like branches (fig. 99). Zooecia multi- 
serial, the branches having from three or four, to twelve or four- 
teen rows ; alternate, oblong, slightly narrower below, truncate 
above (PI. X, fig. 48) ; aperture occupying more than half the 
front; an erect spine at each upper outer angle (sp.) ; a varying 
number of marginal spines, two to five on the inner margin, one 
to three on the outer, curving inward; occasionally in the ab- 
sence of an avicularium, a spine at the bottom of the aperture, 
curving upward (6a. sp.). Avicularia pedunculate, those on 
the marginal zooecia, when present, much larger than those on 
the intermediate zooecia ; often absent from both marginal and 
intermediate zooecia. Ocecia (oe.) large, subglobose, marked by 
radiating lines. Rootlets given off from marginal zooecia near 
base of colony. 

B. murrayana is found at Kadiak, Orca, Pribilof Islands, 
Alaska, and in Puget Sound, but does not extend farther south. 

The specimens from Orca, Prince Williams Sound, are ro- 
bust, often possessing a large number of zooecia in a series form- 
ing broad strap-like segments or branches (PL XVI, fig. 98). 
Those from Kadiak and Puget Sound are more delicate in ap- 
pearance, forming smaller colonies (fig. 99). Those dredged at 
ten fathoms in Puget Sound do not possess more than six or eight 
rows of zooecia in a series. There is great range of variation in 
the number of spines, and in the number and size of avicularia, 
among the specimens from the different localities. A consid- 
erable quantity of material from Pribilof Islands possesses the 
large marginal avicularia, while most of the material from other 
localities lacks them. 

17. Bugula calif ornica sp. nov. 

PI. X, fig. 49; PI. XVI, fig. 100. 

Zoarium consisting of a number of masses whose branches 
have a distinctly spiral arrangement about a central axis, some- 
times forming a coarse growth 75 mm. in height and from 25 
to 50 mm. in diameter in the broadest place (PL XVI, fig. 100). 

268 University of California Publications. [Zoology 

Each branch composed of many strap-like branchlets consisting 
of two series of zocecia, dichotomonsly divided to form a fan or 
frond. Zocecia narrow below, elongated, bearing three spines at 
the distal extremity, one stout and long (PI. X. fig. 49, sp.), 
extending in a direction parallel with the branch, longer than 
the other two. a continuation of the wall of the zooecium; two 
(sp.) at the top of the aperture projecting forward at right 
angles to the front surface. Aperture occupying more than two- 
thirds of the front of the zooecium. Avicularia Large, pedun- 
culate, situated near the top or not more than half the length 
of the zocecinm below the top. Oocia large, high, globose, extend- 
ing above the lower margin of the aperture of the next upper 
zocecinm. Rootlets developed on Lower zooecia, springing from 
front or ventral surface of zooecium below the aperture. 

The zocecia in this species are usually arranged biserially, 
but sometimes, especially near the point where the branch is 
about to divide it may contain for a short distance three or 
four rows of zocecia in the series (fig. 49). B. californica is 
most nearly allied to B. turbinata, but the zooecia are somewhat 
shorter and stouter; the avicularia are of much the same form 
and size, hill are placed lower down on the zooecium; and the 
spines are constantly three instead of two, and of a different 
character from those of B. turbinata. The ocecia also differ, 
those of B. californica Lacking the prominent process or peak 
described for />'. turbinata. 

Colonies of Large size occur abundantly at Dillons Beach, 
California: B. californica is also found, but of less luxuriant 
growth at Lands End, San Francisco Bay, and al Pacific Grove, 
Monterey Bay. 

IS. Bugula pacifica Robertson. 

PI. X, fig. 50; PI. XVI, fig. 101. 
Bugula purpurotincta, Kobertson, '00, p. 3i'o. 
Bugula pacifica Kobertson, '00, p. 321. 

Zoarium consisting of large bushy tufts having a somewhat 
spiral growth, often 75 mm. in height (PI. XVI, fig. 101) ; col- 
onies frequently united by a sort of cable formed by union of 

Vol. 2] Robertson. — Non-Incr listing Bryozoa. 269 

the root fibres. More delicate in appearance than the preceding' 
species, often tinted purple. Zocecia elongated, biserial, armed 
at the distal extremity with three spines, the outer one long 
(300 fi), projecting outward (PI. X, fig. 50, sp.) • the other two 
shorter (100 fi), being continuations of the margin of the aper- 
ture (sp/) • Aperture occupying almost the whole of the front 
of the zocecium. Avicularia large, pedunculate, at the base and 
to the outer side of the aperture. Ooecia remarkably small, not 
rising more than 60 or 80 fx above the zocecium, while the em- 
bryo (errib.), in many cases measuring 200 /x, extends down- 
ward into the upper part of the zooecium. Rootlets numerous, 
extending from the lowest zocecia of a colony. 

This species was formerly ( '00) somewhat tentatively iden- 
tified as B. purpnrotincta Norman, but after more extended 
study of specimens from many different localities it is unques- 
tionably a new species. As formerly suggested by the writer, it 
is given the specific name pacifica, since it seems to be charac- 
teristic of the Pacific Coast. It ranges from Pribilof Islands, 
Bering Sea, to San Francisco Bay, the climax of growth, both 
in quantity and size being reached at Yakutat, Alaska. The 
purple color of the specimens obtained at this place is partic- 
ularly noticeable. This, as I have said, resides in part in the 
tissue lining the zooecia, and in part in the degenerated poly- 
pides or "brown bodies," and is quickly lost after the material 
has been placed in alcohol. From one locality in Puget Sound 
where the species was obtained in abundance, it possessed a dis- 
tinctly greenish hue, while at many other places it is white or 
colored slightly yellow. The extreme shallowness and flatness 
of the ooecia are remarkable characters but are not peculiar 
to this species being found in at least one other species from 
this coast, Bugula longirostrata. The ovum matures in the 
upper part of the zocecium rather than in the ooecium, the em- 
bryo frequently blocking the mouth of the former. In all cases, 
where an embryo has formed, the polypide has degenerated into 
a "brown (purple) body." Unlike those species which possess 
ooecia elevated above the mouth or orifice of the zocecium, func- 
tional polypides and embryos cannot exist simultaneously in 

270 University of California Publications. [Zoology 

the same zocecium, and probably on account of the purely me- 
chanical obstruction caused by the embryo. 

The distribution of B. pacifica extends from Pribilof Islands, 
Bering Sea, to San Francisco Bay. It has been obtained from 
St. Paul, Pribilof Islands ; is very abundant at Yakutat, and is 
found in smaller quantity at Orca, Prince William Sound, 
Alaska; abundant, but colonies smaller, at Sidney, opposite 
Port Orchard Navy Yard, Puget Sound, less abundant on Chan- 
nel Rocks, Puget Sound; fine colonies obtained from rocks at 
Dillons Beach, California ; smaller quantity obtained from Lime 
Point and Fort Point, San Francisco Bay, California. 

1!). Bugula flabellata .1. V. Thompson, 

PI. X, figs. 51, 52. 
Bird's Head Coralline Bills, 1767, Cor. ed., p. 109. PI. XXXVIII, 

fig. 7. 
Cellularia avicularia (part) Pallas, 17f>(>, p. 68. 
Flustra avieularis, Johnston, 1847, p. 346, PI. L XTII, figs. 3, 4. 
Avicularia flabellata Thompson, 1847, MS. Brit. Mus. ; Gray, Brit. 

Mus. Eadiata, p. 106. 
Avieella avicularia Van Beneden, 1848, p. 75. 
Ornithoporina avicularia, d'Orbigny, 1850, p. 322. 
Bugula flabellata, Busk, 1852, p. 44, Pis. LI, LI I. 
Bugula avicularia form:; 2, B. flabellata, Smitt, 1867, pp. 290, 345. 
Bugula /lain /lata 'I lioinpson, 1868, PI. VI, fig. 9. 
Bugula flabellata, Norman, 1868, PL VI, fig. 9. 
Bugula flabellata, Smitt, 1871-72, pt. I, p. 18, PL V, figs. 48-52. 
Bugula flusi r aides, Yen-ill, 1879, p. 52. 
Bugula flabellata, Hincks, 1880, p. 80, PL XI, figs. 1-3. 
not Bugula flabellata, K<>l>ortson, 1900, p. 431. 

Zoarium from 12-25 mm. in height, consisting of a number 
of fan-shaped fronds divided into narrow branches (fig. ."ill. 
Branching dichotomous. occurring where a branch reaches a 
width of six or seven rows of zocecia. Zocecia elongated, of a hi mt 
equal length throughout, apt rture occupying the whole or almost 
the whole of the front; usually two spines placed one above the 
other at each upper angle, the upper spines stiff and flaring, 
the lower often of great length and frequently crossed in front ; 
on marginal zocecia there are generally three spines on the outer 
side and two on the inner (fig. 52). Avicularia less than half 

Vol. 2] Robertson. — Non-Incru sting Bryozoa. 271 

way from the top of each zooecium ; those on the intermediate 
zooecia smaller than those on the margin ; marginal avienlaria 
of medium size, point of mandible bent abruptly downward. 
Oacia small, globular, smooth, well elevated above the top of the 
zooecium, with a double line forming a band on the oral rim; 
opening wide. 

This species differs slightly from Hincks' description of B. 
flabellata, in the number of spines, the English species appar- 
ently having but two on each side of the zooecium. It agrees, 
however, with the figure given by Busk in which he represents 
three spines on the outer side of the marginal zooecia. In other 
respects the English and American species seem to be identical. 

Bugula flabellata has been obtained in small quantity on the 
piles in San Diego Bay. 

20. Bugula pugeti sp. no v. 

PI. X, figs. 53, 54; PI. XI, fig. 55. 
Bugula flabellata, Kobertson, 1900, p. 321. 

Zoarium composed of numerous small somewhat spiral tufts 
from 10 to 25 mm. in height (fig. 53). Zooecia multiserial, 
each branch consisting of from four to seven rows (fig. 
54); long, narrowed slightly below; aperture occupying two- 
thirds or more of the front ; three spines on the marginal zooecia, 
two on the outer side {on. sp.), one on the inner (in sp.) ; inter- 
mediate zooecia with two spines only; the middle of the anterior 
edge of each zooecium occupied by a round knob or process 
whose significance is unknown (k.). Avicularia (av.) found 
only on the marginal zooecia, large, beak somewhat curved, sit- 
uated half way from the top, or just below the middle of the 
zooecium. Ooecia none. Embryos (PI. XI, fig. 55 emb.) develop- 
ing in the anterior part of the zooecium, and no additional ooecium 
being formed. Embryos abundant in the older parts of the 
colony, and those zooecia possessing well developed embryos con- 
taining no polypide. Rootlets extending from the lower zooecia 
of the colony, forming a cluster at its base and attaching it to 
shells, rocks, or sea weed. 

272 University of California Publications. [Zoolooy 

This species resembles B. flabellata in some respects, and was 
previously so identified ('00), but it is characterized by a total 
lack of ocecia and is believed to be an undescribed species. A 
negative character is. to be sure, an unsound basis, in most 
cases, upon which to establish species, but there can be no doubt 
in this instance, that ocecia are nut developed. A well matured 
colony gives many branches in which the life cycle can be read. 
At the anterior edge of a branch the younger polypides are 
found in all stages of development. Tn the zocecia just posterior 
to these. I'nll grown polypides appear and the genital products 
are abundant. In still older zocecia, a young embryo appears 
close to the anterior edge of the orifice of each zooecium, and 
tin' polypide in such a zooecium begins to degenerate. In older 
zooria. nothing is left of the polypide but a brown body, while 
the embryos are well developed, almosl ready to hatch (PI. XI, 
fig. 55, (nib.; />. />.) ■. in still older zocecia. regenerating polypides 
(re. i>(l.) are found in zocecia containing no embryos and from 
which the larvae, presumably, have escaped. The lack of the 
ovicell and the development of the embryo entirely within the 
zooecium, are not the only characters which separate this form 
from /»'. flabellata. The shape of the avicularia, and the number 
and position of the spines are sufficient and constant specific 

This species has been obtained mainly in Puget Sound. It is 
abundant on Channel Rocks at extreme low tide, and has been 
dredged at San .Juan Island, Puget Sound. A small quantity, 
dredged at ten fathoms, was obtained at Sitka, Alaska. 

21. Bugula curvirostrata, sp. no v. 

PI. XI. figs. •",(). .17, 58. 

Zoariimi consisting of numerous strap-like, clichotomously 
branched tufts, from 35 to 75 mm. in height (fig. 56) ; the 
branches much tangled, due to the binding fibres (b. f.) given 
off from the different parts of the colony which unite adjacent 
branches or portions of a branch. Zocecia multiserial, alternate, 
varying from one or two, to four or five, or even seven or eight 
rows (fig. 57) ; long, somewhat truncate above, narrow below, 

Vol. 2] Robertson. — Non-Incr listing Bryozoa. 273 

the sides of the zocecia frequently terminating above in a minute 
denticle (d.) ; aperture occupying two-thirds of the front; below 
the terminal denticle a spine (sp.) sometimes occurs. Avicu- 
laria characteristic, occurring on the lower half of the zocecia, 
to the side of the aperture and either just below, or 
just above its basal margin ; on the intermediate zocecia, below 
the aperture ; on marginal zocecia, usually above the base of the 
aperture; marked by great relative length of beak (bk.) and by 
the distinct sharp curve at its extremity; mandible also curved 
so that at rest the "jaws" do not approximate each other except 
at the tip of the mandible (man.). As is often the case in multi- 
serial species the appendages of the marginal rows are larger 
than are those on the intermediate ones. In this case the mar- 
ginal avicularia (av.) are often gigantic in comparison with the 
intermediate ones (av/). Oozcia (fig. 58, oe.) globose, large, 
marked by faint stria?. Rootlets (fig. 56, r.) very abundant, 
more numerous in the lower part of the colony, arising from 
the dorsal side of the marginal zocecia. Those arising on zocecia 
in upper part of colony frequently attach themselves to adja- 
cent branches to one side or below. 

In habit of growth and general appearance this species re- 
sembles B. murrayana. The difference, however, in the size of 
the various units of the colony in the two species will be clearly 
seen in a comparison of figs. 57 and 58, PL XI, with fig. 48, PL X, 
drawn to the same scale. In B. curvirostrata the spines are 
small, weak, and few in number, while the large size of those of 
B. murrayana is one of its distinguishing features. The avicu- 
laria are also very different both in size and shape ; both species 
possess the curved beak, but the greater relative length of the 
beak of B. curvirostrata and its curvature which prevents its 
complete approximation with the mandible, separate this species 
from B. murrayana, 

B. curvirostrata has not been obtained north of Pacific Grove, 
but is found in considerable abundance at various localities south 
of this point. It appears to be characteristic of southern waters, 
and may perhaps occupy the place in the southern fauna taken 
in the north by B. murrayana. 

274 University of California Publications. [Zoology 

22. Bugula longirostrata, sp. now 

PL XI, figs. 59, 60. 

Zoarium consisting of delicate feathery tufts, branches some- 
what spirally arranged (fig. 59). Branching dichotomous, the 
branches long and slender. Zooecia (fig. 60, zoe.) biserial, much 
elongated, slightly narrowed below, the lateral walls rolling for- 
ward, terminating at the summit on each side of the margin in 
a minute denticle (d.). Aperture occupying more than three- 
fourths of the front of the zooecium. Avicularia (av.) extremely 
slender and long, being more than half the Length of a zooecium; 
the beak (bk.) alone being almost hall* as long as the whole 
appendage; pedunculate, arising on the outer, lateral wall of 
I he zooecium about half way from its summit; when extended in 
a direction parallel witli the zooecium, the tip of the beak of the 
avicularium extends beyond the summit. Oa'cia (oe.) small and 
shallow, just sufficiently high to inclose the ovum (o.), but cover- 
ing only a portion of the embryo (<>nb.), the major portion of 
which occupies t lie upper part of the zooecium. Rootlets arising 
from the dorsal wall of the lower zooecia. 

This species is the most delicate of all the California Bugulas. 
In the relatively large size of its avicularia it resembles the 
preceding species, B. curvirostrata. In the Bugulas generally. 
the muscular part of the avicularium exceeds the beak in size; 
in the two forms. B. curvirostrata and B. longirostrata, the rela- 
tive size of these two parts is reversed, and the beak in each is 
the longer. B. longirostrata also resembles B. pacifica in the 
shallowness of its ooecia, these being insufficient to inclose the 
embryos when fully grown. The ooecium of B- longirostrata is 
slightly larger than that of B. pacifica, but relatively to the 
length of the zooecium it is much the same in both species. 

B. longirostrata has been obtained from one locality only, in 
the submerged valley off La Jolla at a depth of 125 fathoms. 
The specimens consist of a few detached pieces, one of which 
possesed rootlets as shown in the habit sketch (fig. 59). 

Vol. 2] Robertson. — Non-Incr listing Brxjozoa. 275 

23. Bugula laxa, sp. nov. 

PI. XII, figs. 61, 62. 

Zoarium consisting of numerous fronds much divided into 
segments possessing from two to fifteen rows of zooecia in a 
series (fig 61). The fronds forming tangled masses due to the 
great number of rootlets which attach themselves to other fronds 
or to other objects with which they are growing. Zooecia adher- 
ing very loosely together ; narrowed below, having at each ante- 
rior angle a stiff prominent spine (fig. 62, st. sp.), and on each 
margin of the aperture a varying number of smaller spines 
(c. sp.) curving over and frequently meeting and overlapping 
in the middle line ; the number of spines on a side varying from 
three or four, to six or nine, according to the position of the 
zooecium, the outer, marginal zooecia usually possessing, besides 
the stiff spines at the summit, nine spines at regular intervals 
along its length ; the intermediate zooecia having a smaller num- 
ber; aperture occupying the whole of the front of the zooecium. 
Aricularia lacking. Oozcia (oe.) rounded, prominent, with 
faintly radiating striae. Rootlets (fig. 62, r.) numerous, arising 
from the right or left dorsal anterior angle of the marginal, and 
of many of the intermediate zooecia. 

This species resembles both Bugula and Flustra so greatly 
that it is difficult to decide into which genus it should be placed. 
The shape of the zooecia, the looseness of their connection with 
each other, and more especially the fact that each individual 
zooecium arises from a bud formed independently from a parent 
zooecium, and not from a common growing marginal region as 
in the Flustras, indicate a closer relationship to Bugula. Its 
habit is very characteristic. The great number of root fibres 
developed on its dorsal surface cause it to adhere somewhat 
closely to the substratum, in this case a sponge, and to form a 
tangled mass from which it is difficult to separate large pieces. 
The colony begins in a single zooecium from which either one 
or two buds arise forming branches which extend sometimes as 
long narrow strips of one or two series of zooecia, sometimes as 
broad fronds of many series. The adjacent rows of zooecia are 

276 University of California Publications. [Zoology 

so loosely connected that in the broader frond-like portions. 
lacuna? frequently occur where zooecia have apparently failed to 
form, thus producing the open and irregular appearance pecu- 
liar to this species. 

B. laxa occurs in considerable abundance on Channel Rocks. 
Puget Sound. Found in small quantity in material from Pacific 

Beania Johnston. 

Beania Johnston, 1S47, ed. II, p. 372. 
Diachoris Busk, 1852, pt. I, p. 53. 

Zoarium subcorneous or calcareous, ered or decumbent. 
Zooecia sessile or suberect, scattered, or in loosely connected 
groups, united to one another by slender tubes originating from 
the dorsal or lateral surfaces; aperturt occupying the entire 
front, the margin usually furnished with hollow spinous pro- 
cesses arching over the opening; mouth terminal. 

As here defined, Beania is intended to include the genus 
Diachoris of Busk, Bincks, Jullien, and others, following the 
usage of MacGrillivray and Waters. Bincks ('85) regards Dia- 
choris of Busk as an artificial division hul considers it to be 
more nearly related to Bugida because of its hoat -shaped ZCKEcia 
and articulated a\ iciilaria. These characters do not, however, 
distinguish Bugula alone, whereas the peculiar mode of connec- 
tion of the zooecia by tubes of varying Length forms a good diag- 
nostic character common to both Beania and Diachoris as orig- 
inally defined. 

24. Beania mirabilis Johnston. 

PI. XII, figs. 63, 64, and text figure 1. 

Beania mirabilis Johnston, 1847, p. 372, text figs. 69, 70. 

Beania mirabilis, Husk, 1852, pt. I, p. 32, PI. XXIV, figs. 4, 5. 

Beania mirabilis, Sniitt, 1867, pp. 295 and 357. 

Beania mirabilis, Hincks, 1880, p. 95, PI. IV, figs. 8-10. 

Zoarium consisting of numerous minute branching tufts, 
some of which are free, some adherent to the substratum by 
means of rootlets given off from each zocecium (fig. 63). Zooecia 

Vol. 2] Robertson. — Non-lncr listing Bryozoa. 277 

boat-shaped (fig. 64, zoe.), having two erect spines (e. sp.) above 
the month, and a variable number of marginal spines, six, seven, 
or eight on each side of the aperture; tubes (con. t.) connecting 
the zoceeia arising near the base of each zocecinm laterally or on 
the dorsal surface, i.e., the side opposite the aperture; near the 
point of origin of a tube a septum (sep.) forms, the longer por- 
tion of the tube being continuous with the zocecium of which it 
forms a sort of pedicel ; opposite branches may be given off from 
the base of each zocecium (br.). Ooeciaf Rootlets (text fig. 1, 
r. f.) consist of tubular processes of varying length given off 
from the basal surface of each zocecium and terminating in a 
disk from which finger-like processes extend anchoring the colony 
to the substratum. 

Beania mirabUis is obtained in considerable quantity growing 
over Alcyonaria and Bowerban~kia at Dead Mans Island and 
Santa Catalina Island. 

25. Beania longispinosa sp. nov. 

PI. XII, figs. 65, 66. 

Zoarium sub-erect, consisting of groups of zoceeia united by 
tubular connections (fig. 65). Zoceeia boat-shaped, slightly con- 
tracted above, alternate; aperture membranous occupying the 
whole of the front ; orifice terminal ; spines remarkable for their 
length, the two upper ones (fig. 66, st. sp.) stiff and flaring out- 
ward; seven to nine pairs (cr. sp.) on the margin, inclined in- 
ward, crossing above the aperture, their tips frequently ex- 
tending beyond the lateral edges. Connecting tubes very short, 
often hidden by the overlapping lateral walls of the zoceeia, 
usually two on each side, uniting lateral zoceeia, one at each end 
uniting with a zocecium laterally and proximally. No avicu- 
laria. Ocecia (oe.) small, situated at the summit of the zoceeia, 
marked by parallel striae extending longitudinally. Rootlets very 
long, extending from the dorsal surface of many of the zoceeia, 
sometimes 50 or more mm. in length (fig. 65, r.). 

This species seems to be related to Diachoris hyaclesi Jullien 
( '88) in the number of spines and the arrangement of the con- 

278 University of California Publications. [Zoology 

necting tubes, but the difference in both these characters are 
sufficiently distinctive to constitute another species. The number 
of connecting tubes varies somewhat, there being six, seven, and 
sometimes eight from one zooeeium. 

Beania longispinosd was obtained in several fathoms of water 
off La Jolla, California, growing upon rock. 

Stirparia Goldstein. 

Stbrparia Goldstein, 1879 or 1SS0. 
Stir/tarta, Ilincks, lss.'5. 
Bicellaria (part). Busk, 1884. 

Zoarium consisting of ered segmented stalks, bearing on their 
summits feathery tufts whose branches eonsisl of zocecia bise- 
riaily arranged. Zocecia of the normal Bicellarian or Bugulan 
type, i.e., turbinate, with apertun looking more or less upward, 
turned obliquely inward; or boat-shaped, with aperture occu- 
pying almost all of the front wall, not directed upward and 
zoiecia not turned inward; lower portion tubular. Avicularia 
and ocecia pedunculate. 

The diagnosis of this genus as given by Hincks ('83) is here 
amended in order to include the two types of zocecia found in 
the species from this coast, as well as the ocecia, of which Hincks 
makes do mention. Ooecia are abundant on all the specimens 
and are of the Bicellarian type, being pedunculate upon the inner 
side of the zooeeium near the top. 


1. Zocecia with aperture turned upward 2 

1. Zocecia with aperture not turned upward 3 

2. Aperture occupying less than half the front, and armed with five 

or six spines S. ciliata 

_'. Aperture occupying half or more of the front wall 3 

'.). Margin of zocecia armed with two, three, or four spines 

S. occidentalis 

:'.. M;irgin of zocecia armed with two spines on the upper outer angle 

S. calif ornica 

Vol. 2] Robertson. — Non-Incrusting Bryozoa. 279 

26. Stirparia ciliata, sp. nov. 

PI. XII, figs. 67, 68, 69; PI. XIII, figs. 70, 71. 

Zoarium composed of numerous flabellate tufts borne on the 
summits of a number of erect, stiff, segmented branching stalks, 
the whole attaining the height of 25 or more mm. (fig. 67). 
The segments (fig. 68, seg.) of the stalk aborted zocecia, the lat- 
eral disk at the distal end of each representing the aborted aper- 
ture (ab. ap.). Stalks branching at irregular intervals, branches 
(br.) frequently arising from the aborted aperture. Segments 
of about equal length and formed by rather deep constrictions, 
there being no distinct joint or articulation, the segments of the 
stalk passing insensibly into the normal zocecia forming the tufts 
at the distal end of the stalk ; the first zooecium of the branching 
tufts similar to the distal segment (dis. seg.) of the stalk, being 
often but little shorter, and but slightly modified in other re- 
spects from the ordinary stalk segments. This first zooecium tubu- 
lar, elongated, possessing a circular aperture occupying about 
one-third the length of the zooecium, and surrounded by spines ; 
from its anterior extremity two zocecia arise, and as growth 
proceeds the zocecia that subsequently appear in the branch ac- 
quire the typical form. Zocecia of the Bicellarian type, broadly 
ovate above, tubular below, the aperture looking upward, the 
tubular portion inclined inward (PI. XIII, fig. 70). Aperture 
occupying about half the front or less, margin thin, surrounded 
by six or seven spines (c. sp.) which tend to curve inward. Five 
or six spines on the outer edge, one on the inner just opposite 
the lowest outer spine; zocecium at bifurcation (bi. zoe.) usually 
lacking all spines except the two opposite lowest ones. Avicu- 
laria (av.) pedunculate, small, situated just below the aperture 
on the outer side, usually developed sparingly. Ocecia (PL XIII, 
fig. 71, oe.) pedunculate, globose, developed on the side of the 
aperture just above the spine on the inner margin ; found on 
every zooecium of some of the branches, especially of those at 
the tips. Rootlets springing from the lower segments of the 
stalk, proceeding downward close to the stalk, often enclosing it 
with a sheath, then becoming free at its base, spreading, often 

280 University of California Publications. [Zoology 

becoming branched, and attaching themselves to grains of sand, 
to hydroid stems, or other substrata. 

In the possession of a chitinous stalk marked by constrictions 
more or less deep but not distinctly articulated, <S Y . ciliata is re- 
lated to 8. annulata, Maplestone (79). It differs markedly from 
that species, however, in the total lack of the annulations found 
on the segments of the stalk of 8. annulata and for which it is 
named. Many evidences are afforded of the zooecial origin of 
these segments. Instances are found where a segment assumes 
the shape externally of an ordinary zooecium, i.e., it becomes 
wider at the distal end and possesses an aperture on whose mar- 
gin are two and sometimes three spines (PL XII, fig. 68, ab. ap.). 
The contents of such a segment, however, are similar to that of 
adjacent segments. The avicularia which are usually minute 
often become more aumerous and slightly larger Dear the tips of 
the branches. 

This is a rather widely distributed California species, being 
found among the material from Lands End, Fort Point, and 
Lime Point, San Francisco Bay; Pacific Grove, Mendocino City, 
and Dillons Beach, California. 

27. Stirparia occidentalis, sp. nov. 

PI. XIII, figs. 72, 73, 74. 

Zoarium composed of flabellate tufts borne on the summit of 

a number of erect, stiff, segmented branching stalks, the whole 
an inch or a little more in height (fig. 72). The segments (seg.) 
of the stalk not so evidently aborted zooecia as are those of the 
preceding species, the lateral aborted apertures appearing only 
occasionally; the articulation (art.) of the segments distinct; 
segments differing in length, growing noticeably shorter to- 
ward the distal end of the stalk, and the transition (tr.) from 
stalk to zooecial tuft being very sudden. Zooecial tufts relatively 
short, not half as long as the main stalk, branching dichotomous. 
Zocecia biserial, alternate ; at the formation of a branch, the 
series of zooecia separated by the interpolation of two new zocecia 
on the inner side so that no one zooecium can be said to be at 

Vol. 2] Robertson. — Non-Incr listing Bryozoa. 281 

the bifurcation (fig. 74, hi.), i.e., the branches bifurcate between 
two series of zooecia forming the proximal branch and not upon 
a single zocecium. Zooecia (zee.) of the Bugulan type, elongated, 
turned outward, but aperture not turned distinctly upward; 
aperture elongated, occupying more than half the front surface, 
the lateral margins raised and terminating at the upper angles 
in long curving spines (c. sp.). Two or three spines (sp.) on 
the upper, outer margin, and one on the inner (sp.). Avicularia 
(av.) pedunculate, situated below and to one side of the aper- 
ture; present usually on every zocecium. Oa'cia (oe.) peduncu- 
late, to one side of the summit of the zocecium. Rootlets arising 
from the lower segments of the stalk, proceeding downward 
close to it, and at its base spreading freely and attaching them- 
selves to the substratum, becoming entangled with those of other 
colonies and forming a sort of cable whose strands are further 
bound together by organisms parasitic upon them. 

These two species 8. ciliata and S. occidentalis resemble each 
other in habit of growth, but the zocecial portion of the latter 
is relatively shorter and the stalks relatively longer than are 
those of 8. ciliata. This is so evident that a macroscopic exam- 
ination is sufficient to separate them. The greater length of the 
stalks of S. occidentalis is due probably to the greater length of 
many of the segments and not to a greater number of segments. 
Thus a segment of the lower part of a stalk shown in PI. XIV, 
fig. 76, is twice the length and thickness of those of the upper 
portion of the same stalk represented in PI. XIII, fig. 73. 

This species is found in several localities on the coast of Cali- 
fornia and Puget Sound. 

28. Stirparia calif ornica sp. no v. 

PL XIII, fig. 75; PI. XIV, figs. 77, 78, 79, 80. 

Zoarium consisting of tall stalks surmounted by large, feath- 
ery, branching tufts of zooecia (PL XIII, fig. 75) ; the stalk, 
both absolutely and relatively, longer in this species than in 
the two preceding, being two and a half or three times as long 
as the zocecial tufts. Segments of the stalk stout, some very long, 

282 University of California Publications. [Zoology 

others short, showing- a kind of alternation sometimes, and grow- 
ing decidedly shorter toward the distal end: zooecial origin of 
stalk segment not always apparent, and transition into ordinary 
zocecia, abrupt. Single stalks broad at point of attachment to 
substratum, sending out ribs (PL XIV, fig. 77, rb.) of chitinous 
material which are connected by a flat web-like disk of cuticle. 
In older stalks, branching occurs a short distance above the base. 
Colonies solitary, no instance found of runners or stolon-like 
processes uniting several. Zocecia resembling those of Bugvla, 
especially of B. /><i<i/i<a ; elongated, only slightly narrower below 
than above: apertun occupying nearly the whole of the front; 
zocecia not turned outward nor is the aperture upturned (PI. 
XIV, fig. 80). Lateral margins of aperture raised, especially ;ii 
the summit of the zocecia where each angle terminates in a long 
jointed spin* ; from the distal margin of a zocecium a second 
longer spine (sp.) appears, ma king two, less frequently three, on 
the outer margin, and one on the inner. Avicularia (av.) small, 
often minute, pedunculate, on the lower part of each zocecium 
just below the aperture. Ooecia {oe.) globose, pedunculate, on 
the inner side at the summit of each zocecium. Rootlets spring- 
ing from lower segments of the branches of the stalk, passing 
downward close to stalk and attaching themselves to basal disk. 
(PI. XIII, tig. 75, r.) 

I>nt two species of Stirparia have been hitherto described, S. 
glabra, Bincks ('83), and 8. anwulata, Maplestone ('79). both 
from Australian waters. To these, three others from the collec- 
tions on this coast are here added. In the three West American 
species, two types of zocecia are discernible, and two types of stalk 
segments. In all accounts hitherto given of this genus the Bicel- 
larian form of the zocecium is considered characteristic, so much 
so that Busk ( '84) retains S. glabra in the genus Bicellaria. In 
the three species here described, there is shown a transition from 
the Bicellarian type to the Bugulan. Thus, S. ciliata closely 
resembles Bicellaria ciliata in shape and ornamentation of zoce- 
cium, but in S. occidentalis there is a decided approach to the 
Bugulan type, and in S. californica we find the boat-shaped 
zocecium characteristic of Bugula. In all, the avicularia and 

Vol. 2] Robertson. — Non-Incr listing Bryozoa. 283 

ooeeia are much alike both as to size and position, and resemble 
those of Bicellaria. The stalk of these three species shows varia- 
tion tending- toward greater specialization. Attention has already 
been called to the fact that, as Hincks ('83) observed, the seg- 
ments of the stalk are probably aborted zooecia. Their zocecial 
nature is most plainly shown in S. ciliata, in which as shown in 
PI. XII, fig. 68, the distal segment (dis. seg.) of the stalk is not 
unlike a normal zocecium in shape, and in the possession of a 
definite aperture round whose margin seven spines may be 
counted. This zooecium-like segment, however, does not contain 
a polypide, but instead, a strand of tissue similar to that found 
in other segments of the same stalk. In the remaining species, 
8. occidentalis and S. californica, the zocecial character of the 
segments of the stalk is much less apparent, the aperture on 
each being inconspicuous and the articulation of the stalk being 
much more definite, showing a higher grade of specialization 
than is found in 8. ciliata. With the specialization of the seg- 
ments of the stalk there is correlated greater size, both in length 
and thickness, not only of the stalk as a whole but of the indi- 
vidual segments as well. 

Attention has already been called to the difference in size 
between the lower and upper segments of the stalk of S. occi- 
dentalis (PI. XIII, fig. 73; PL XIV, fig. 76). A similar differ- 
ence exists in the segments of the stalk of 8. californica, fig. 78 
representing in outline one of the longer, fig. 79, one of the 
shorter segments, all the figures being drawn to the same magni- 
fication. Furthermore, a distinction is made between S. califor- 
nica and any other members of the genus in the possession of a 
foot disk by which the colony adheres to the substratum. Fig. 77 
represents the mode of attachment of a small colony consisting 
of a single segmented stalk. Where the stalk approaches the 
substratum it spreads out in a thin membranous disk, strength- 
ened at intervals by chitinous ribs (rb.) which form in the ecto- 
cyst of the lower portion of the stalk and extend to the circum- 
ference of the disk. A colony may consist of a single stalk 
arising from such a disk and surmounted by a zocecial tuft; or 
as in PI. XIII, fig. 75, the stalk which forms the foot disk may 

284 University of California Publications. [Zoology 

divide into a number of secondary stalks (br.) each surmounted 
by its zocecial tuft and sending down rootlets which attach them- 
selves to the base. The colonies of S. California) grow in isolated 
groups, thus differing from 8. ciliata and 8. occidentalis, which 
form tangled masses of stems springing from many interwoven 

8. ciliata and 8. occidentalis are shore forms, being' found 
between tides growing on sea weed, other bryozoa, etc. 8. cali- 
fomica, on the other hand, is a deep water form, being dredged 
at a depth of 125 fathoms in the submerged valley off La Jolla. 

Corynoporella Bincks. 

Cori/in'iHin Ha llincks, ISsS. 

Zoarium forming delicate white tufts, consisting of zocecia 
disposed in a single series, and facing one way. dichotomously 
branched. Zocecia more or less clavate, each originating from the 
dorsal surface of the one beneath it. immediately below the sum- 
mit; elongated, attenuated, tubular; aperturt occupying a half 
or less of the front surface. Avicularia articulated, attached to 
the side of the aperture. Ocscia globose. 

29. Corynoporella spinosa sp. nov. 

PI. XXV, figs. 61, 82, 83. 

Zoarium erect, much branched, each internode consisting of 

a single zocecium (fig. 81); hr<in<ins (fig. Si', hr.) arising on 
the dorsal surface near the summit, singly or in pairs. Zocecia 
sub-clavate, narrowed below, widened above, the distal margin 
truncate or only slightly rounded; aperturt membranous, occu- 
pying a third or more often a half of the front surface; sur- 
rounded h\ a thin raised margin on the distal edge of which 
are three minute spines or denticles, two lateral and one median, 
the latter being frequently absent. Avicularia (au.) large, 
rounded, pedunculated, situated half way between the summit 
and the base on the side of the aperture ; mandible flat and 
rather broad and short, terminating in a point below the ab- 
ruptly bent beak. Oa ] ci<i (fig. 83, oe.) globose, prominent, as 

Vol. 2] Robertson. — Non-Incrusting Bryozoa. 285 

wide as the zocecial aperture, with line's radiating from the front 
rim ; few in number. Rootlets (fig. 82, r.) springing from a disk 
on the side of most of the zooecia, just below where the branches 

But one other species of this genus is described, viz.. C. tenuis, 
Hincks ('88) from the St. Lawrence. As Hincks remarks, "the 
zocecia bear a strong resemblance to those of the genus Brettia, 
but the articulated avicularium is a link connecting it with 
Bugula." The species C. spinosa has a decided resemblance to 
Bugula in that its zocecia approximate the boat shape typical for 
that genus. It seems also to bear a resemblance to Bugulella 
Verrill ('79), but in Corynoporella the method of branching as; 
well as the mode of union of the zoreeia are distinctly different 
from that in this genus. 

Obtained from material from Alaska growing on a crab. 
Exact locality unknown. 

Notamiidae Hincks. 

GemellariadcB (part) Busk, 1852. 
Notamiidce Hincks, 1880. 

Zocecia in pairs, each pair arising by two tubular prolonga- 
tions, one from each member of the pair next below it; at each 
bifurcation a new series of zocecia intercalated into the branches. 

Synnotum Hincks. 

Noiamia, Waters, 1885. 
Synnotum Hincks, 1S86. 
Synnotum, Waters, 1896. 

Zoarium consisting of erect, slender, bifurcating shoots, at- 
tached by a mass of tubular fibres given off from the base of 
the primary zocecium. (Fig. 93.) Zocecia in pairs joined back 
to back, elongated, expanding from the base upward, the front 
surface occupied by a membranous aperture: sessile avicularia 
and an articulated avicularium between the zocecia in each pair 
;it the summit. Ocecia? 

286 University of California Publications. [Zoology 

30. Synnotum aviculare (Pieper) Hincks. 

PI. XIV, figs. 84, 85. 
Gemellaria avicularis Pieper, 1881, Vol. IX, p. 43, PL II, fig. 5-6. 
Notamia avicularis, "Waters, 1885, Ser. 2, Vol. 5, p. 6. 
Synnotum aviculare, Hincks, 1886, Ser. 5, Vol. 17, p. 257. 
Synnotum aviculare, Waters, 1896, p. 14, PL I, fig. G, 7. 

Zoarium minute, slender, of delicate texture, glossy, simple 
in habit, consisting of long branches which bifurcate at rather 
distant intervals (fig. 84). Zocecia attenuated below, expanded 
above (fig. 85) ; aperture occupying more than two-thirds of the 
front, narrowing below, the margin thin and unarmed: lateral 
aviculari-a {lot. av.) small, sessile placed at the top of the zoce- 
eium on the inner side, developed on every pair of zou-cia. widen- 
ing from the base upward with a minute beak; median avicularia 
(me. av.) pedunculate, capitate, placed at the top of each pair 
of zocecia in the median plane, subglobular, smooth, the beak 
\t-vy slightly produced, with a sharp spike-like extremity. 

The sessile avicularia in the California specimens are not 
alternate' as Hincks ( 'So') describes for those from the Adriatic, 
but appear rather on every pair. The capitate avicularia are fre- 
quently Lacking in our colonies but this may be due to mutilation. 
The mull, Is (fig. So, >■.) frequently arise quite high up on a 
branch from the fronl surface on the line midway between the 
top and base of a pair of zocecia, sometimes even upon the aper- 
ture of a zoo'cium. 

S. aviculare occurs with other minute forms of bryozoa at 
Dead Mans Island. San Pedro, and Ballast Point. San Diego, 

Cellariidae Hincks. 

Escharidce (part) Johnston, 1847. 
Salicornariadce Busk, 1852. 
Ceilariece Smitt, 1867. 
Cellariida? Hincks, 1880. 

Zoarium erect, calcareous, articulated: branching dichoto- 
mous or non-diehotomous. Zocecia usually rhomboidal or hexag- 
onal, disposed in series around an imaginary axis so as to form 
cylindrical shoots. 

Vol. 2] Robertson. — Non-Incrusting Bryozoa. 287 

Cellaria Lamouroux. 

Cellaria Lamouroux, 1812. 
Salicornaria Cuvier, 1817. 
Cellaria (part), Johnston, 1847. 
Salicornaria, Johnston, 1847. 
Cellaria (part) d'Orbigny, 1850. 
Salicornaria, Busk, 1852. 
Cellaria, Hincks, 1880. 

Zoarium jointed at intervals, internodes connected by horny 
tubes. Zoozcia alternate, depressed in front, and surrounded by 
a raised border. Avicularia immersed, irregularly distributed, 
situated above a zocecium, or in place of one. Oozcia immersed. 


1. Branching dichotomous; avicularia with triangular mandible directed 
downward C. borealis 

1. Branching not dichotomous 2 

2. Internodes small, avicularia large, twice the size of an ordinary 

zocecium, mandible rounded, directed upward C. mandibulata 

2. Internodes long, few, avicularia small, mandible rounded, directed 
upward or outward C. diffusa 

31. Cellaria borealis (Busk) Smitt. 

PI. XIV, fig. 86; PI. XVI, fig. 102. 
Salicornaria borealis Busk, 1855, p. 254, PI. I, figs. 1, 2, 3. 
Cellaria borealis, Smitt, 1867, pp. 383 and 361, PI. XX, fig. 17. 

Zoarium growing in luxuriant masses 75-100 mm. in height 
(PI. XVI, fig. 102), branching dichotomous, except at the ex- 
tremities where the internodes are frequently tipped with three 
or four very small terminal internodes. Internodes of varying 
length, about a centimeter on an average, club-shaped; joints 
not black ; young actively growing portions of the colony bright 
flesh pink. Zoozcia (PI. XIV, fig. 86, zoe.) elongate, very little 
wider toward the middle than at the two ends; alternate, sur- 
rounded by a raised crenulate border, the rows of zonecia sepa- 
rated by sinuous lines; front wall calcareous, and except upon 
the operculum which is large and semicircular on the distal 
edge, is marked by rather coarse punctures; orifice at summit 
of zooecium. Avicularia (av.) above the zocecia distributed at 

'288 University of California Publications. [Zoology 

irregular intervals; mandible triangular, pointing downward. 
Oceciaf Rootlets not very numerous, springing from the lower 
internodes of a colony. 

This is a handsome, well marked species, originally figured 
and described by Busk from the west Greenland shore. Tt is 
abundant on the rocks at Orca, Prince Williams Sound. Tt 
seems to be a strictly northern species not being reported smith 
of Queen Charlotte Islands. 

32. Cellaria mandibulata Bincks. 

I I. XV, figs. 87, 88, 89; II. XVI, fig. 103. 
Cellaria mandibulata Eineks, Issl'. p. 462. 
Cellaria mandibulata Ilin.ks. 1884, \>. 203, PL IX. fig. 7. 

Zoarium forming nmsses of delicate branches often growing 
from 75-90 mm. in height (PI. XVI, fig. 103). luh, nodes 
slender, short, the longesl from »i 1<> 8 mm. in Length. Branch- 
ing irregular, branches usually arising near the t<>|> of an inter- 
node, Imt often from the middle, or from any point on its sur- 
face; when arising from the top of ;m internode, two branches 
are seldom opposite, the one usually being higher than the other. 
.1 hi hIs black. Zocecia of various shapes depending apparently 
upon age. In young stages, they are slender, narrowed almost 
to a point above and below, widest about the middle (PI. XV, 
I'm'. 87 I -. in older stages they are somewhat wider, truncate above 
and below | fig. 88) ; in the oldest stage, zoiecia broader and 
shorter relatively (fig. 89); alternate, in a line from one end of 
the internode to the other, surrounded by a broad more or less 
crenulate border (fig. 88, cr. bor.). In young stages, this border 
follows the length of the internode in sinuous lines which ap- 
proach each other but do not meet above and below the zooecia 
(fig. 87) ; in older stages, the sinuous border becomes united 
above and below the zooeeia, and the erenulations are less marked 
(fig. 89). Zoo?cial wall convex, covered with minute punctures, 
which in the oldest stage seem to cover the border with one 
continuous slightly punctate mantle (fig. 89). Operculum (op.) 
semicircular, with straight lower margin, a denticle at each lower 

Vol. 2] Robertson. — Non-Incr listing Bryozoa. 289 

corner, and two others opposite at the corners of the upper mar- 
gin. Avicularia (av.) characteristic, large, few in number, only 
one or two in an internode, of simple form, almost exactly like 
extra large zooecia ; much broader than ordinary zooecia, with a 
large semicircular chitinous mandible extending upward, whose 
upper margin is considerably elevated above the surface of the 
internode; above the avicularium is a small orifice resembling 
in shape and position an immature ocecium (or.) Ooccia not 
found in the youngest zocecial stage described above. In older 
stages the orifice of the ooecium (fig. 88, oe.) appears circular, 
or somewhat oblong, occupying the space on each zocecium above 
the operculum; in the fully developed stage (fig. 89, oe.), the 
ocecia resemble minute avicularia, with a straight lower margin, 
semicircular above and slightly raised above the surface of the 
zocecium. Numerous fine brown rootlets spring from the lower 
zooecia of the colony. 

Without doubt this is the species described by Hincks from 
Queen Charlotte Islands. It is readily distinguished from other 
Cellaria of this region by its large dark avicularia which may 
be seen easily with a lens of low power. The method of branch- 
ing, which Hincks seems to think is probably not characteristic 
of the species, is certainly typical both of this and another spe- 
cies, C. diffusa, less commonly found here. C. mandibulata has 
a wide distribution but is most common in southern waters, 
being frequently dredged in the vicinity of both San Pedro and 
San Diego. 

33. Cellaria diffusa sp. no v. 

PI. XV, fig. 90; PI. XVI, fig. 104. 

Zoarium consisting of a relatively small number of rather 
long cylindrical internodes connected by dark chitinous joints; 
internodes varying in length, the longest attaining a length of 
35 or 40 mm., the whole colony often 75 mm. high, and having 
a straggling diffuse appearance due to its method of branching. 
Branching irregular, branches arising at any point on an inter- 
node, always in the middle of a zocecium; two branches, some- 
times three or four arising at the distal end of the internode 

290 University of California Publications. [Zoology 

(PI. XVI, fig. 104). Joints black. Internodes club-shaped, stout. 
Zoo3cia depressed, narrower below and above than in the middle, 
truncate at each end ; front wall calcareous, covered with minute 
protuberances. (PI. XV, hi:. 90.) Operculum {op.) some dis- 
tance below the top of the zooecium, lower edge slightly curved, 
bearing a denticle id.) near each corner, upper edge semicir- 
cular. Oaecial orifia {oe. or. I situated just above the operculum 
and resembling it in shape; a broad mucro (m.), a continuation 
of the calcareous zooecial wall in the middle of its lower edue. 
Avicularium {av.) in place of a zocecium, almost square. Root- 
lets springing from the lower zooecia of the proximal internode, 
passing down dose to the wall of the internode. then spreading 
out disk-like around the l»;ise of the colony. 

In its habil of growth this species resembles C. australis 
Hincks ('84). The stems are not divided into internodes of 
definite and equal Length by a regular dichotOmous branching 
as is usual in typical Cellaria, but consist of rather stout. Long, 
club-shaped cylinders giving off branches at any point appar- 
ently, though as in C. ausiralis and G. mandibulata which it 
reseiiililes in habit of growth and method of branching, always 
from the middle of a zooecium. The zooecia resemble those of 
C. australis in shape, hut the position and shape of opercula 
and oiecia are different. Both have avicularia of the same gen- 
eral type, and of a character similar to C. fistulosa. 

Fine colonies have been obtained at S;m Juan Island, and in 
Port Orchard Channel. Pugel Sound. It has also been dredged 
both at San Pedro and San Dieuo. 

Flustridae Smitt. 

Escharidce (part), Johnston, 1847. 
Flustridce (part), d'Orbigny, 1850. 
Flustrada (part), Busk, 1852. 
Flustridce Smitt, 1867. 
Flu.strUhr, Hincks, 1880. 

Zoariwm corneous and flexible, expanded, foliaceous, erect 
or sub-erect. Zoacia contiguous, multiserial. Avicularia usually 
of a simple type. 

Vol. 2] Robertson. — N on-Incr listing Bryozoa. 291 

Flustra Liunaens. 

Flustra sp., Linn., 1758. 
Eschara (part) Pallas, 1766. 
Flustra, Smitt, 1867. 
Flustra, Busk, 1852. 
Carbasea Busk, 1852. 
Flustra, Hincks, 1880. 

Zoarium erect or sub-erect. Zocecia disposed in a single or 
double layer, more or less quadrangular or linguiform, with a 
raised margin, the aperture occupying the whole or a consid- 
erable portion of the front, and closed in by a membranous 
covering. Oc&cia immersed or raised. 

34. Flustra lichenoides Robertson. 

PI. XV, figs. 91, 92; PI. XVI, fig. 105. 
Flustra lichenoides Eobertson, 1900, p. 322, PI. XX, figs. 7, 7a, 8. 

Zoarium unilaminar, consisting of broad foliaceous green or 
brownish green fronds. (PL XVI, fig. 105.) Zocecia in alter- 
nate rows, slightly arched above, narrowed below, the distal 
margin raised and armed at each corner with an erect stiff 
spine (fig. 91 e. sp.), often slightly curved; below on each side 
of the aperture two other spines (c. sp.), flattened and curving 
inward, often meeting in the middle above the aperture ; aperture 
occupying the whole front. Ocecia (oe.) rather low, but raised 
above the surface of the zocecia ; not as high as the erect spine 
at its base. Avicularia? Rootlets (fig. 92 r. f.) arising from 
the upper corner of most of the zocecia by means of which the 
fronds are attached to the substratum, and above which the 
colony does not rise to any height but spreads out in convoluted 

Considerable variation occurs in the development of the 
spines. Specimens are found in which no spines are visible 
except those at the corners ; others which have but one pair of 
lateral spines, or in which the spines are unevenly developed on 
the two sides. Variation also occurs in the root fibres. These 
frequently anastomose and form a net work of fibres below the 
lowest frond. Root fibres from overlapping fronds often attach 

292 University of California Publications. [Zoology 

themselves to the margin of the zocecia of a lower frond. The 
Laminae are sometimes united back to back, but are easily sep- 
arable, their union being effected by means of short fibres. 

This species occurs sparingly on the Alaska coast; in Puget 
Sound in considerable abundance; to some extent on the coast 
of California as far south as San Francisco. 

35. Flustra membranaceo-truncata Smitt. 

PL XV, figs. 93, 94. 

Flustra membranaceo-trmcata Smitt, 1867, p. 3;"8, PI. XX, 
figs. 1-5. 

Zoarium c posed of a number of ereet, nnilaminate fronds 

(fig. 93). ZooRcia irregularly quadrangular, truncate above and 
below, often narrowed below (fig. 94) ; armed with a delicate 
spine (sp.) at the distal corners; apertun occupying the whole 
of the front; operculum (op.) semicircular. Avicularia (ar.) 
sparingly developed, in place of a zooecium; mandible semicir- 
cular, directed upward. Otiecia (oe.) small, immersed, no1 quite 
as wide as the zooecium against which each projects. 

This appears to be the species which Smitt ('67) describes 
and figures, ami which he finds in Arctic Seas growing on ascid- 
ians. sertularians, etc., either creeping or erect. The material 
in this collection consists of a few small erect fronds obtained at 
Pribilof Islands, Beriim Sea. 

I'n in rsity of California. 

Berkeley, February l~>, 1905. 

"Vol. 2] Robertson. — Non-Incrusting Bryozoa. 293 


Allman, J. G. 

1S56. A Monograph on British Fresh-Water Polyzoa. Eay Society, Lon- 
don. Pp. 1-120, Pis. I-XI. 

Busk, G. 

1849. Observations on Anguinaria spathulata. Trans. Micr. Soc. Vol. 
II, pp. 123-126. 

Busk, G. 

1852. Catalogue of the Marine Polyzoa. Part I, Cheilostomata (part). 
London. Pp. 1-54, Pis. I-LXVIII. 

Busk, G. 

1855. Zoophytology. Quart. Journ. Micr. Science, Vol. Ill,- pp. 253-256 
and 320-322, Pis. I-IV. 

Busk, G. 

1881. List of Polyzoa collected by Capt. H. W. Feilden in the North- 
Polar Expedition, &c. Journ. Linn. Soc, Vol. XV, pp. 231- 
241, PI. I. 

Busk, G. 

1884. Eeport on the Polysoa collected by H. M. S. "Challenger," during 
the years 1873-76. The Cheilostomata, in voyage of H. M. S. 
"Challenger," Zoology X, pp. 4-216, Pis. 36. 

Conrad, T. A. 

1855. Note on the Miocene and Post-Pliocene deposits of California, with 
Descriptions of two new Fossil Corals. Proc. Acad. Nat. ScL, 
Philadelphia, p. 441. 

Coppin, J. 

1848. Description of a new genus of British Marine Zoophytes belonging 
to the family Eucratiada?. Ann. Nat. Hist., (2), Vol. IT, 
pp. 273-274, PL X. 

Ehrenberg, C. G. 

1831. Symbols Physica?. 

Ellis, J. 

1767. Versuch einer Natur-Geschichte der Corall-Arten. Niirnberg. Pp. 
168, Pis. I-XLVI. 

Ellis, J. 

1767. An Account of the Actinia sociata, &c. Phil. Trans., Vol. LVII, 
pp. 428-436, PL I. 

Ellis and Solander. 

1786. A Natural History of the Zoophytes. Pp. 208, Pis. 63. 

294 University of California Publications. [Zoology 

Farre, A. 

1837. On the Minute Structure of some Polypi, &c. Phil. Trans., pp 
387-426, Pis. 8. 

Fewkes, J. W. 

1889. New Invertebrata from the Coast of California. Bull. Essex 
Institute, XXI, pp. 99-146, Pis. I- VII. 

Gabb and Horn. 

1862. Monograph of the Fossil Polyzoa of the Secondary and Tertiary 
Formations of North America. Journ. Acad. Nat. Sci., Phila., 
n. s., Vol. V, pt. II. pp. 111-179. Pis. 19-21. 

Goldstein, J. R. Y. 

1879. New sptrirs of Polyzoa. Quart. Journ. .Micr. Soc, Victoria. Vol. I. 

Harmer, S. F. 

1896. Polyzoa. The Cambridge Natural History. Vol. II. pp. 465-533. 

Hassall, A. H. 

1841. Catalogue of Irish Zoophytes. Ann. Nat. Hist., Vol. VII. pp. 
363-373, Pis. VI X. 
Hincks, Th. 

1880. A History of the British Marino Polyzoa. 2 vols. Vol. I, pp. 

CXLI, 601. wood cuts B6j Vol. II, Pis. 83. 
Hincks, Th. 

1882. Polyzoa of Queen Charlotte Islands. Ann. Nat. Hist., (5) Vol. X. 
pp. 248-256. 
Hincks, Th. 

1N83. Contributions toward a General History of the Marine Polyzoa. 
Ann. Nat. Hist. (5), Vol. XI, pp. 193-202, Pis. 2. 
Hincks, Th. 

1884. Polyzoa of Queen Charlotte Islands. Ann. Nat. Hist. (5), Vol. 
13, pp. 203-215, PI. I. 
Hincks, Th. 

lsS4. Contributions toward a General History of the Marine Polyzoa. 
Ann. Nat. Hist., (5), Vol. XIII. pp. 356-369. Pis. 2. 
Hincks, Th. 

18S5. Contributions toward a General History of the Marine Polyzoa. 
Ann. Nat. Hist., (6), Vol. XV, pp. 244-257, Pis. VII-IX. 
Hincks, Th. 

1888. The Polyzoa of the St. Lawrence: A Study of Arctic Forms. Ann. 

Xat. Hist. (6) Vol. I, pp. 214-227, Pis. XIV ami XV. 
Jelly, E. C. 

1889. A Synonymic Catalogue of the Recent Marine Bryozoa including 

Fossil Synonyms. London. Pp. 322. 

Vol. 2] Robertson. — Non-Incr listing Bryozoa. 295 

Johnston, G. 

1847. British Zoophytes. 2 vols. Vol. I, pp. 251-406; Vol. II, Pis. 

Jullien, J. 

1888. Mission Scientifique du Cap Horn. T. VI. Zoologie, Bryozoaires. 
Pp. 92, Pis. 15. 

Lamouroux, J. V. 

1812. Extrait d'un memoire sur la classification des polypiers coralli- 
genes. Bull, des sci. pour la Soc. Philomatique. 

Lamouroux, J. V. 

1816. Hist. Nat. des Polypiers coralligenes flexibles. 

Linnaeus, L. 

1758. Systema naturae. Ed. 10. Pp. 1384. 

MacGillivray, P. H. 

1887. Catalogue of Marine Polyzoa of Australia. Tr. Roy. Soc. Vict. 
Vol. 22, pp. 187-224. 

Maplestone, C. M. 

1879. On Victorian Polyzoa. Quart. Journ. Micr. Sci., Victoria. Vol. 
I, p. 19. 

Nitsche, H. 

1869. Beitrage zur Kenntniss d. Bryozoens. Zeitschr. wiss. Zool., Vol. 
XX, pp. 1-36, Pis. 3. 

Norman, A. M. 

1868. Notes on rare British Polyzoa, with descriptions of new Species. 
Quart. Journ. Mic. Sci., n. s., Vol. VIII, pp. 212-222, Pis. 3. 

Oken, L. 

1815. Lehrbuch der Naturgeschichte. Abt. 2. 

Orbigny, A. d' 

1850. Paleontologie Fran§aise-Terrains eretaces. Vol. V, Bryozoaires. 
Text, pp. 1191; atlas, Pis. 800. 

Pallas, P. S. 

1766. Elenehus Zoophytorum. Pp. 451. 

Pieper, F. W. 

1881. Eine neue Bryozoe der Adria, Gemellaria (?) avicularis. Jahresber. 
Westfalischen Prov.-Vereins, Vol. IX, pp. 43-48, PI. I. 

Robertson, A. 

1899. Studies in the Pacific Coast Entoprocts. Proc. Cal. Acad. Sci., 
(3), Vol. II, pp. 323-348, PI. I. 

296 University of California Publications. [Zoology 

Robertson, A. 

1900. Papers from the Harriman Alaska Expedition. The Bryozoa. 
Proc. Wash. Acad. Sci., Vol. II, pp. 315-335, Pis. XIX-XXI. 

Savigny, J. C. 

1811. Iconographie des Zoophytes de l'Egypte, from the 'Description de 
Smitt, F. 

1867. Kritisk Forteckning bfver Skandinaviens Hafs-Bryozoen, ofv. K. 
Vet. Akad. Forh., pp. u7;i-4Ui». Pis. 5. 
Smitt, F. 

1871-72. Florida n Bryozoa. Kgl. Svenska Vet. -Akad. Eandlingar., Vol. 
X. pp. 20, Pis. 5. 
Trask, J. 

1857. On some new Microscopic Organisms. Proc. Cal. Acad. Sci., Vol. 
Ill, pp. 110-115, Pis. 2. 

Van Beneden, P. J. 

1848. but les Polypiers bryozoaires de la Mer du Nord. Bull. 

Acad. Royal Beige., Vol. XV. pp. 67-82, PI. 1. 

Van Beneden, P. J. 

1849. Recherches sur les Bryozoaires de la mer du Nord (suite), et 

projel 'rune classification des animauz de ce groupe. Bull. 
Acad. Royal Beige., Vol. XVI. pt. 2, pp. 644-658, Pis. 2. 

Verrill, A. E. 

L879. Notice of recenl Additions to the Marine Fauna of the Eastern 
Coasl of North America. Amer. Journ. s.-i., ser. 3, Vol. 
XVIII, pp. 52-54. 
Waters, A. W. 

1879. Bryozoa of the Hay of Naples. Ann. Nat. Hist. (5) Vol. Ml. 
pp. 28-43, Pis. 2; pp. L14-126, Pis. 4; pp. 192-202, Pis. 3; 
pp. 267-281, I 'Is. l 1 . 

Waters, A. W. 

L885. On the Use of the Avicularian Mandible. Journ. Roy. Micr. Soe. 
7ol. 5, pp. 77-1 779, PI. XIV. 

Waters, A. W. 

L887. Bryozoa from New South Wales. North Australia, &c« Part I. 
A.m. Nat. Hist. (5) XX. pp. 81-95; PI. IV. Pari 11. 1. e. pp. 
181-20::. Hart III. I. .-. pp. 253-265, HI. VII. 

Waters, A. W. 

1889. Bryozoa from Now South Wales. Ann. Xat. Hist. (6) Vol. IV. 
L889, pp. 1-24, Pis. I-III. 
Waters, A. W. 

1896. Notes on Bryozoa from Rapallo and other Mediterranean locali- 
ties — chiefly CellulariidcB. J. Linn. See., Lond. XXVI, pp. 
1-21, Pis. I and 11, and 3 fig. in text. 

Vol. 2] 

Robertson. — Non-Incrusting Bryozoa. 



a. — area. 

ab. ap. — aborted aperture. 
ad. — adnate part of zooecium. 
ap. — aperture. 

b. sp. — basal spine. 

b. f. — binding fibre. 
bi. — bifurcation. 

bi. av. — avicularium at bifurcating 

bi. sc. — bifid scutum. 
bi. sp. — bifid spine. 
bl". — beak. 
br. — branch. 

c. av. — curved avicularium. 
cr. sp. — crossed spine. 

c. sp. — curved spine. 
car. val. — cardiac valve. 
con. tu. — connecting tube. 
cr. bor. — crenulate border. 
cr. br. — creeping branch. 

d. — denticle. 

dis. zee. — distal zooecium. 

e. br. — erect branch.^ 

e. sp. — erect spine. 

/'/-. av. — frontal avicularium. 

fr. zee. — frontal zooecium. 

fst. zee. — first zooecium. 

giz. — gizzard. 

gr. — groove. 

i. — intestine. 

in. sp. — inner spine. 

int. — internode. 

,;'. — joint. 

~k. — knob. 

kit. av. — lateral avicularium. 

lat. zee. — lateral zooecium. 

in. — mouth. 

man. — mandible. 

me. av. — median avicularium. 

mem. ap. — membranous aperture. 

m us. — muscle. 

o. — ovum. 

oe. — ocecium. 

oe. or. — ocecial orifice. 

03S. — oesophagus. 

op. — operculum. 

or. — orifice. 

ov. sp. — outer spine. 

ov. — ovary. 

p. — pore. 

par. mils. — parieto-vaginal muscle. 

pd. — polypide. 

ped. av. — pedunculate avicularium. 

ph. — pharynx. 

pr. br. — primary branch. 

py. val. — pyloric valve. 

r. — rootlet. 

rb. — rib. — root chamber. 

r. fb. — root fibre. 

re. in us. — retractor muscle. 

re. sh. mus. — retractile sheath muscle. 

re. pd. — regenerated polypide. 

S. It. — snake's head. 

sc. — scutum. 

sec. br. — secondary branch. 

seg. — segment. 

sep. — septum. 

sh. mus. — sheath muscle. 

sm. av. — small lateral avicularium. 

sp. — spine. 

st. — stalk. 

ter. br. — tertiary branch. 

tr. — transition from segment of stalk 
to zooecium. 

tu. — tubular part of the zooecium. 

n. r. — upward extending rootlet. 

V. — vibraculum. 

v. c. — vibracular cell. 

v. ch. — vibracular chamber. 

zee. — zooecium. 

zee. bi. — zooecium at bifurcation. 

zee. br. — zocecial branch. 

zee. or. — zooecial orifice. 


Fig. 1. — Aetea anguina Linnaeus. Eabit sketch, about natural size. 

Fig. 2. — A. anguina. Portion of a colony enlarged showing variations 
in height of the erect, tubular part (tu.) of the zooecia. Adherent 
(ad.) portion attached to the substratum like a stolon is part of the zocecium 
which rises at its anterior end into the ringed tubular portion (tu.). The 
polypide (/"/.) can he seen inside several of the zocecia; when fully ex- 
panded it protrudes through the operculum (op.) at the summit of the tube. 
The zooecia are separated from each other by a septum (sep.) at the anterior 
end of each. X 30. 

Pig. 3. — A. anguina. Pari of a zocecium in which the polypide (pd.) 

is in a state of retraction; upper part of tubular portion broken 
away. Polypide retracted by cunt raid inn of retractor muscle fibres (nuts.) 
attached to the wall of the adherent portion (ad.) of the znnviuin. In the 
adherent portion the n\ a ry (OV.) is visible. X 70. 

Pig. 4. — .!. anguina. A single zocecium much enlarged showing position 
of ocecium (oe.) just outside the aperture ("/»•)- below the operculum on 
the ventral side of the zocecium. Within the ocecium is an ovum in early 
cleavage Btage I < nth.). X 70. 

Pig. ."). — Aetea truncata Landsborough. Portion of a colony enlarged to 
show variations in height of the erect, tubular portions ('«•), also length 
and slenderness of the adherent portions ("</.) of the zooecia. X 30. 

Fig. 6. — A. truncata. Habit sketch, about natural size. 


Univ. Cal.Publ.Zddl.Vdl.II. 

Roberts lin] Plate IV 

7fi t"V 




Fig. 7. — Eucratea chelata Linnaeus. Babit sketch, about natural size 

Pig. 8. — K. chelata. Enlarged to show zooecia and method of branching 
[e, in.) just below the aperture (op.)» :|1 "' ''"' creeping branches (cr. br.) 
given off from the sides of the zooecia. X 50. 

Pig. 9. /•.'. (Inlaid. Showing ooecium (oe.) on a small aborted zocecium. 
r Hincks.) 

Pig. L0. Gerr.ellaria loricata Linnaeus. sketch, natural size. 

Fig. 11. G. Icricata. Portion of a branch enlarged to slmw arrange 
incut of zooecia and method of branching. • 30. 

Pig. L2. Menipea ternata Ellis and Solander. Eabil Bketch of sped 
men fr< m Pugel Sound. 

Pig. 13. — M. ternata. Habit sketch of specimen from California. 

Pig. 1!. .1/. ternata. Enlarged portion of colony from Pugel Sound 
showing zooecia in groups of three, except the ocecial internodes (oe.). Lat 
era! ai icularia I av. | well developed. ■ 30. 

Pig. 15.- .U. ternata. Dorsal view of the zocecium at the bifurcation of 
a branch (bi. br.) showing the adjoining zooecia {lat. see.) and root chamber 
(i-.ih.) with upward extending rootle! (r.). 30. 

Pig. 16. M. ternata. A single zooecium to shew the position of the two 
kinds of root fibres, those anchoring the colony (r.), arising in a Bimple 
rod disk (r. d. ) on the fronl wall of the zooecium; those extending upward 
(a. >•.). arising from a rather large chamber (r. ch.) projecting from the 
zooecia! wall jusl above the lateral avicularium. ■ 30. 

Univ. Cal.Publ.Zddl.Vql.II, 

Rdbertsdn] Plate V 




fhcto-luhlg ar 


Fig. 17. M. ternata. Enlarged portion of colony from California. 
Much like the preceding except that the lateral avicularia are often lacking-. 

Fig. 18. — M. gracilis Busk. Habit sketch, natural size. 

Pig. 19- .1/. gracilis. Portion of a branch enlarged to same Bcale aa 

figs. 14 and 17, showing the greater length i E the zooecia, the large tlabellate 
scutum, the relatively small size of the avicularia (OV.), and the shape 
and position of the two sorts of root chambers (r. d. ami r. ch.) similar to 
thi se of .1/. ternata. X 30. 

Fig. 20.- .1/. gracilis. Portion of an ocecial internode showing develop- 
meiit of avicularia (av.) on the front of the zooecium close to the ooecia 
(oc). X 30. 

Fig. 21. .1/. gracilis. Dorsal view of the zooecium at the bifurcation of 
a branch (hi. see.), showing the lateral adjoining zooecia (lat. see.), and 
rod chamber (r. ch.) with upward extending rootlel («. v.). X 30. 

Fig. 22.- .1/. occidentalis Trask. Habil sketch of a single branch to 

show method of branching. Somewhat diagrammatic 

1 i^ r . -'A. — .1/. occidentalis. A few internodes enlarged to show- size of 
avicularia (at;.), position and size of scutum (sc), the nature of the jointed 

spines (sp.), and the crossed spines (c. Sp.). Drawn to the same scale as 
Bgs. 11. 17. and 19. X 30. 

fig. l'4. — M. occidentalis. An ooecial (oe.) internode consisting of five 
zooecia. X 30. 

Fig. 25.— M. occidentalis. Lateral view of a zooecium, showing a. root 
chamlier (r. ch.) from which there extends a downward extending rootlet 
( r. ) . X 30. 


Univ. Cal.Publ.Zdql.Vdl.II. 

Roberts dn] Plate VI 




Fig. 26. — M. occidentalis catalinensis, subsp. dov. Portion of a branch 
showing finger-like processes on scutum (sc), and bifid spines (c. sp.). X 50. 

Pig. 27. — .1/. occidentalis catalinensis. Lateral view of a zocecium show- 
ing runt chamber (r ch.) and downward extending root fibre (>'.). 

Fig. 28. Menipca cncta Robertson. Habit sketch, natural size. 

Fig. 29. — M. (i<rin. Enlarged portion of a colony from Kadiak show- 
ing lateral avicularia (lat. av.) } frontal avicularia (fr. av.) } and Bpines 
(sp.). X 30. 

Pig. 30. — .1/. erecta. Enlarged portion of a colony from Sitka showing 
absence of lateral avicularia (lat. av.) ; showing also rootlet (r.) springing 
directly from zoceeial wall. X 30. 

Pig. 31. — .1/. erecta. Enlarged portion of colony from San Juan showing 
absence of spines. X 30. 

Fig. 32. — Menipea pribilofi, sp. nov. Habit sketch, natural size. 

Fig. 33. — M. pribilofi. Ooecial internode (oe.). X 30. 


Univ. Cal.Publ.Zddl.Vql.II 

Rdbertsdn] Plate VII 




Fig. 34. — M. pribilofi. Portion of a branch enlarged Bhowing number 
and length of jointed spines (sp.), size of lateral avicularia (Int. av.) } 
frontal avicularia (fr. av.), and root chambers in the two positions (r. ch. 
an. I r. d.). X 30. 

Fig. 35. Scrupocellaria californica Trask. Dorsal view of part of two 
zocecia to show vibracular chamber (v. ch.), short vibraculum (v.), groove 
of vibraculum (gr.) } and lateral avicularium belon (av.). X 50. 

Pig. 36. — S. californica. Portion of a branch enlarged showing varia 

tiiia in size of lateral avicularia (int. av.), the vibracular cell (v. c), on 
zooecium at bifurcation (zoe. hi.), visible from the front, while on other 
zooecia only the short vibraculum (v.) is seen from the front, and only on 

the lower zocecia which possess root fibres (r.). • 50. 

Pig. 36a. — S. californica. Habit sketch. 

Pig. 37. — S. californica. Three zooecia with ooecia (oe.). X 50. 

Pig. 38. — Scrupocellaria varians Bincks. Portion of a branch enlarged 

to show bifid and trifid scutum (SC.), lateral avicularia varying in size from 

very small (s. av.) to the large curved variety (c. av.) ; also the vibracular 

cell visible from the front (r. ch.), and the Long vibraculum (v.) present on 
each zoecium. X 20. 

Pig. 39. — S. varians. Dorsal view of a portion of two zocecia to shew 
position of vibracular chamber (v. ch.) with its long vibraculum (v.) and 
its groove (gr.), with the root disk (r. d.) and root fibre (jr. /'.) on its 
basal margin; also lateral avicularium with its much elongated curved beak 
(c. av.). X 20. 

Pier, do.- Caberea ellisi Fleming, sketch. 


Univ. Cal.Publ.Zddl.Vdl.II. 

Rdbertsdn] Plate VIII. 




Fig. 41. — S. diegensis sp. aov. Portion of a branch enlarged showing 
form of zooecia (~oc.) with large scutum (sc), spines on tin- upper margin 
of the aperture (ap.) one or more of which are bifid (hi. sp.) ; also lateral 
avieularium (lat. av.) and frontal avicularium on the zooecium at bifurcation 
(bi. av.). On those zooecia not possessing ouvia the vibracular ehaml er 
(v. ch.) plainly visible frith its long vibraculum (v.). X 5D. 

Fig. 42. — S. diegensis. Dorsal view of a few zooecia to show vibracular 
chamber (v. ch.) sustaining the long vibraculum (v.) at its summit, and 
showing the groove (gr.) extending in the direction of the length of the 
zooecia. At the base of the vibracular chan 1 er and to one side of the groove 
is the pi re (p.) from which the rool fibre extends (r.). X •";<>. 

Pig. 43.— &. diegensis. A vibracular chamber enlarged to slow detail. 

Fig. 44.-- S. diegensis. 'The avicularium on zooecium at bifurcation en 
larged to show its large muscular portion (mus.) and the obliquely directed 
mandible (man.). 

Pig. 45. — Caberea ellisi Fleming. A few zocecia enlarged. Vibracular 
chamber visible from the front (v. ch.); the minute lateral avieularia (Int. 
<n-.)\ serrated vibraculum (v.). X 50. 

Pig. 4(i. — C. ellisi. Dorsal view of a few zooecia to show vibracular 

(v. ill.) chambers ea ling obliquely across the hack of each zooecium, 

the groove '///•.). and the long vibraculum (v.); also the rootlets (r.) aris- 
ing from each vibracular chamber and proceeding downward through the 
middle of the branch. Rootlets from one side only shown. X 50. 

Fig. 47. — Bugula neritina (Linn.) Oken. Portion of a branch enlarged 
showing zocecia (zee.) with large aperture (ap.). margins of which termi- 
i ate in denticles (d.) ; also pedunculated ocecium (of.). X 30. 



[Rdhertsdn] Plate IX 




Pig. 4.8.— Bugula murrayana Johnston. Portion of a branch enlarged 
showing zocecia (ece.) with margins of aperture armed with variable number 
of spines (sp.) ; avicularium (<ir.) or spine (sp.) a1 base of aperture. Mar- 
vie:.! avicularia lacking on this specimen. - 30. 

Fig. 49. — B. calij ornica sp. doy. Portion of a branch enlarged to show 
spines terminating the upper margins of the aperture (sp.),long miter spine 
ion. sp.), and position of avicularia (av.). 50. 

Fig. 50. — B. pacifica sp. cov. Portion of a branch enlarged to show 
zooecia (see.), spines (sp. and sp.'), and the shallow ooecia (oe.); also the 
embryo (emb.) in the upper part of the zooecium. ■ 50. 

Pig. 51.— Bugula flabellata Thompson. Habit sketch, natural size. 

Pig. 52.- Bugula flabellata. Portion of a colony enlarged to show ar- 
rangement and branching of zooecia, the spines (sp.), avicularia (av.), and 
ocecia (oe.). ■ 30. 

Fig. 53. Bugula pugeti sp. uov. Habit sketch. 

Pig. 54. /•'. pugeti. Zooecia with lorg aperture (op.), marginal avicu- 
. the three spines on the marginal zooecia (out. sp.), and the 
process on the distal margin (A.). X 30. 


Univ. Cal.Publ.Zddl.Vdl.II. 

Robertson] . 




Fig. 55. — B. pugeti. A few zocecia of lower part of a branch enlarged 
to show tlic embryo (emb.) in the upper part of the zoceeium, and the 
brown body (6. b.) in each zoceeium containing an embryo; a regenerating 
polypide (re. pd.) in one zoceeium. ■ :i<». 

Fig. ."<;. Bugula curvirostrata sp. uov. Babit sketch, aboul natural size. 

Fig. 57. — B. curvirostrata. Portion of a branch enlarged to slmu zocecia 
and the (two sorta of avicularia (av. and av'.). X 30. 

Pig. 58. />'. curvirostrata. Portion of a branch Bhowing three zocecia 
with oo eia (oe.). X 30. 

Pig. 59. — Bugula longirostrata sp. nov. Jlaliit sketch, aboul aatural size. 

Pig. 60- /.'. longirostrata. Portion of a branch showing attenuation of 
zocecia, and avicularia (av.), and the shallowness of the ocecia (oe.) which 
only partially cover the embryos, (emb.). X 50. 

[ Roberts on] Plate XI 



Fig. 61. — Bugvla laxa sp. aov. Eabit sketch. 

Pig. (>:.'. /»'. laxa. I'm! inn of :i branch enlarged Bhowing form of zoce- 
cia, marginal spines (sp.) and ooecia (oe.). ■ 50. 

Pig. 03. — Beaiiia mirabilis Johnston. Eabil sketch. 

Pig. (il. /»'. mirabilis. A tew zocecia enlarged in show mode of connec 
tinii (con. /.), the erecl and decumbenl portions, the former with margins 
armed with spines. X 50. 

Pig. 65. Beania longispinosa sp. aov. Eabil sketch. 

Fig. 60. — B. longispinosa. Portion of n colony enlarged to show char- 
acter of zooecia, spines, ami the ooecia (oe.). X 30. 

Pig. 67. Stirparia ciliata sp. qov. Eabil sketch. 

Pig. 68.- 8. ciliata. Portion of the stalk enlarged showing seymeniatiou 
(seg.), also the beginning of tlie zoceciaJ branches (see. br.). X 50. 

Pig. 69.- S. ciliata. Lower segmenl of stalk. ■ :,n. 


Univ. Cal.Publ.Zddl.Vdl.II. 

Roberts dn Plate XII 



Fig. 70.- S. ciliata. Portion of zooecial branch enlarged to show shape 
and arrangement of the zooecia (see.), the curved marginal spines (c. sp.), 
and t he minute avicularia. >. 50. 

Fig. 71.— S. ciliata. A few zooecia enlarged to show pedunculate ooecia 

• ). X 50. 

Pig. 72. Stirparia occidental™ sp. aov. Eabil Bketch, natural size. 

Fig. 73.— S. occidentaUs. Upper portion of tin' stalk and a few of tin' 
lower zooecia, showing a specialization <>t' the segments (sep.) of the 
their articulation (art.), and the Budden transition into zooecia where the 
zooecial tufts begin (fgt. zee.). ■ 50. 

Pig. 74. —S. occidentals. Portion of a branch to show character of 
zooecia (see.), of avicularia (av.). spii-es > x/<. ). ami ooecia (oe.). > 50. 

Pig. 75. Stirparia califomica sp. cov. Habil sketch, natural size. 


Univ. Cal.Publ.Zddl.Vdl.II. 

Roberts qn Plate XIII 




Fig. 76. — Stirparia Occident alis, sp. nov. One of the lower segments of 
the stalk drawn to the same scale as fig. 73. PI. XIII, to show difference 
in length i f segments in fche upper and lower parts of the stalk, x 50. 

Pig. 77.- Stirparia californica sp. nov. Portion of the stalk enlarged to 
show mode of attachment of single stalk to substratum by means of a foot 
disk. ■ 50. 

Pig. 78. 8. californica. One of the longer segments of the stalk en- 
larged to the same scale as fig. 76. X 50. 

Fig. 79. — 8. californica. One of the shorter segments of the stalk en- 
larged to tin' Bame scale as the preceding figure. X 50. 

Pig. 80. — S. californica. Portion of a branch enlarged showing Bugulan 
character of the zooecia, position of aviculari and the Bicellarian 

like ooecia (oe.). X 50. 

Pig. 81.- Corynoporella spinosa, sp. nov. Babil sketch. 

Pig, 82. C. spinosa. A t'.-w zooecia enlarged showing method of branch- 
ing, position of avicularia (<"■.>. of spines (sp.) } and rootlets (r.). '■' 50. 

Pig. 83. C. spinosa. A single zooecium with ocecium (oe.). X 50. 

Pig. 84. Synnotum avicvluvi Pieper. Babil sketch. 

Fig. 85. — S. avicvlare. Portion of branch enlarged showing character 
nf zocecia, root fibres (r.), of sessile (*. av.), ami pedunculate avicularia 
(ped. or.), x 50. 

Fig. 86.- Cellaria borealis Busk. A few zooecia enlarged in show I 
arrangement, the avicularia (av.) just above the operculum (op.) sur- 
rounded by the crenulate border (cr. bor.) similar to that separating the 
lines of zooecia. X 50. 



Rdbertsdn] Plate XIV 


Pig. 87. — Cellaria mandibulata Eincks. Two zocecia, young stage show- 
ing lateral crenulated border (cr. bcl.). 

Fig. 88. — C. mandibulata. Several zocecia at an older stage, showing 
operculum (op.) with its four denticles (</.). the immersed ocecia above, 
their orifices (oe. or.) only visible, and an avicularium with its Large man- 
dible (man.). The crenulated border (cr. bor.) has formed above and be 
Low t!i" zocecia, thus surrounding them. X 50. 

Pig. 89. — C. mandibulata. Thret al a still older stage to si 

Increase of calcareous wall and final form of ocecia] orifice (oe. or.) re- 
sembling a small avicularium. X 50. 

I [g. 90. Cellaria diffusa sp. nov. A few zocecia (see.) to show shape, 
ocecia! orifice (oe. or.) with the lower margin developed into a mucro 
and the avicularia (av.). X 50. 

Fig. 91. — Flustra lichenoides Robertson. A few zocecia enlarged to show 
erect spines i'. sp.), operculum (op.), and ocecia (oe.). X 50. 

Pig. 92. -F. lichenoides. The dorsal surface of a few zocecia showing 
n i I fibres springing from the distal angle of each zocecium. 

Pig. 93. — Flustra membranaceo-truncata Smitt. Eabil sketch. 

Fig. 94. — F. membranaceo-truncata. A few zocecia showing arrange- 
ment, immersed ocecia (oe.), ami avicularium of simple structure in place of 
zooecium (av.). X 50. 


Univ. Cal.Publ.Zddl.Vdl.II. 

Robertson] Plate XV 



Photograj lis to show habit of growth. 

Pig. 95. Scrupocellaria " inns Hincks. 

Pig. 96. Scrupocellaria < Robertson. 

Pig. 97. Bugula neritina (Linnseus) Oken. 

Pig. 98. Bugula murrayana Johnston. Specimen from Orca, Alaska. 

Pig. 99. Bugula murrayana Johnston. Specimen from Pugel Sound. 

Pig. LOO. Bugula californica Robertson. 

Pig. 101.- Bugula pacifica Roberts 

Pig. L02. d llaria borealis i Busk i Him 

Fijj. 103. (ilium mandibulata Eiincks. 

Fig. 104.- Cellaria diffusa Robertson. 

Pig. IQ5.—Flustra lichenoides Robertson. 

Univ. Cal.Publ.Zddl.Vdl.U. 

Rdbertsdn Plate X/l 


Vol. 2, Nos. 6 and 7, pp. 323-340 December 13, 1905 











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An examination of Pacific Coast hydroids during the past 
few years (vid. Torrey, :02, :04) has convinced me that many 
if not all species change their structural type with age. Stems 
straight proximally may become sinuous distally. Branches 
which alternate during the early stages of colonial development 
may later originate in pairs. Length and annulation of hydro- 
thecal pedicels, size, proportions and . ornamentation of hydro- 
thecae may similarly vary with the distance from the base of 
stem or branch. One hydroid in particular exhibited marked 
changes in structure with the growth of the colony and appeared 
to be especially favorable material for the investigation of cer- 
tain questions of differentiation. 

This species was described some months ago as Clytia bakeri 1 
(Torrey, :04, p. 16). The stems, usually unb ranched, rise from 
a creeping hydrorhiza to the height of 20 or 30 mm. For a 
few millimeters (2 to 5) each stem is free of polyps and is com- 
pletely annulated. The annuli, short at the base, increase grad- 
ually in length until they become, at the first hydranth pedicel. 

1 The Hydroids of the San Diego Region. Univ. Cal. Publ. Zoology. II, 
No. 1, 1904. 

324 University of California Publications. [Zcologt 

two to four times as long as broad. They ultimately grade into 
the internodal segments of the stem, each giving rise at its distal 
end 1" one hydrothecal pedicel. The pedicels themselves vary in 
length and annulation from the base of the stem toward the tip. 
Tin- first pedicel may exhibit 4 to 7 annuli. The last hydro- 
theca may possess no pedicel at all. or one of l>ut a single an- 
quIus. The number to a pedicel varies with much general uni- 
formity from one of these extremes to the other. 

Prom base to tip, then, each stem presents two main serial 
changes. (1) Below the hydranth region, the cauline annuli 
grow progressively longer. (2) Within the hydranth region, the 
number of annuli to a pedicel progressively decreases. Their 
is a strong suggestion in these facts thai they depend on changes 
in the internal physiological conditions of growth. Since polyps 
are readily regenerated where stems are sectioned, a method a1 
once presents itself for determining the differentiation a1 a given 
level of the stem at a given time. 

The question to answer which the investigation was first un- 
dertaken was: Will the stem .-it a given level tend to regenerate 
the structural type which it originally produced there: or will 
it produce instead, a structural type characteristic of another 
region .' For example, will a cu1 through the portion of the stem 
where each hydrothecal pedicel has five annuli initiate the re- 
generation of a pedicel with five or with one annul ns. the number 
characteristic of the <list;il region? 

The facts obtained from the experiments fall into three cate- 
gories, according as the regeneration followed ilia cut through 
the distal portion of the stem. (2) a cut through the middle por- 
tion, lot heteromorphically from tin- cut basal end. 

In all. 16 operations were performed. Of these. 7 were unsuc- 
cessful. Among the remaining 9, the cut in 5 had been made 
through the distal region, 3 through the middle region, and 6 
had regenerated heteromorphic stalks. The small number of 
cases involved is offset by the clearness of the results. 

The record of experiments in the first category is as follows: 

I. — July 2(i. No. l'. Stem with 24 hydranths and terminal 
hud. Enumerating from base to tip, 

Vol. 2] Torrey. — Hydroid Differentiation and Senescence. 325 

Pedicels 1, 2 with 3 annuli 
Pedicels 3-8 with 4 annuli 
Pedicels 9-11 with 3 annuli 
Pedicels 13-17 with 1 annulus 
Pedicel IS with annuli 
Pedicel 23 with annuli 
Pedicel 24 with 1 annulus 

10 a.m. Cut off terminal bud, 24, 23, and part of 22. 

July 28, 10 a.m. New hydranth No. 22 regenerated within old 
cup, on a short stalk from which hydrotheca is separated by a 
single constriction and is therefore sessile. 

Result : regeneration is according to the local, not the basal 
structural type. 

II. — July 26, No. 3. Stem with 17 hydranths and terminal 


Pedicel 1 with 3 annuli 

Pedicel 2 with 3 annuli 

Pedicel 3 with 4 annuli 

Pedicel 7 with 1 annulus 

Pedicel 9 with 1 annulus 

Pedicel 11 with annuli 

Pedicel 13 with annuli 

Pedicel 15 with annuli 

Pedicel 17 with not even nodal constriction 

12 m. Removed terminal bud, 17, 16, and part of 15. 

July 27, 9 :15 a.m. No. 16 has regenerated as a bud on short 
stalk ; no constriction whatever between latter and hydrotheca 
(fig. 1). Though there are two cut surfaces, but one bud has 
been formed, in place of the originally proximal individual. 
No. 16 developed later into a hydranth, the skeleton remaining 
in the same condition except for a slight general thickening. 

Result : regeneration is according to the local, not the basal 
structural type. 

III. — August 5, No. 4. Stem with 15 hydranths and terminal 


Pedicels 1-4 with 2 annuli 
Pedicels 5-11 with 1 annulus 
Pedicel 12 with annuli 
Pedicel 13 with 3 annuli 
Pedicels 14-15 with 1 annulus 
Pedicel 16 with annuli 

326 University of California Publications. [Zoology 

8 :45 a.m. Removed terminal bud, 15, 14, and 13. 

August 7, 9:45 a.m. Xos. 13 and 14 have regenerated, each 
with a single annulus in the pedicel. 

August 8, 2 p.m. Stem segment with bud of 15 has ap- 
peared; single annulus below hydrotheca. 

Result : all the regenerated parts accord with the structural 
type characteristic of the distal region (fig. 2). 

IV. — August 5, No. 5. Si cm with 17 hydranths. 

Pedicel 1 with 8 annuli 
Pedicel 2 with 6 annuli 

Pedicel 3 with 4 annuli 
Pedicel 4 with ."> .-111111111 
Pedicel ."» with - annuli 
Pedicel 6 with 4 annuli 
Pedicel 7 with 3 annuli 
Pedicels 8-9 with 2 annuli 
Pedicels 10-11 with :'> annuli 
1'edicels 12 L5 with - annuli 
Pedicel 16 with 1 annulus 
Pedicel 17 with <• annuli 

8:50 a.m. Removed 17 and 16. 

August 11. No. 16 has regenerated. Pedicel with 1 (pos- 
sibly 2) annulus and 16-18 tentacles. 

Result: this stem was somewhat irregular with regard to the 
number of annuli in successive pedicels. The regenerated ped- 
icel, however, corresponds with the local structural type. 

V. — August 5. No. 6. Stem with 13 hydranths and terminal 

Pedicel 1 with t> annuli 

Pedicel 2 \\ ith 4 annuli 

Pedicel 3 with 5 annuli 

1'edicel 4 with 4 annuli 

Pedicel 5 with 3 annuli 

Pedicel 6 with 4 annuli 

Pedicel 7 with 3 annuli 

Pedicel 8 with 1 annulus 

Pedicel !* with 3 annuli 

Pedicel 10 with 2 annuli 
Pedicels 11-14 with 1 annulus 

8:55 a.m. Removed terminal bud. 13 and 12. 

Vol. 2] Torrcy. — Hydroid Differentiation and Senescence. 327 

August 8. Nos. 12 and 13 regenerated, each with 1 annulus in 

Result : regeneration according to the local, not the basal 
structural type. 

Further experiments of this sort are unnecessary to show con- 

Figs. 1, 2. Normal regeneration from distal wound. 
Fig. 3. Normal regeneration from middle region of stem. 
Fig. 4. Heteromorphic regeneration from base of stem. 

The arrow indicates in each case the level of the cut. 

328 University of California Publications. [Zoology 

clusively that, under the conditions of the experiments 1 regen- 
eration from cuts through the distal region of the stem produces 
structures of the local, i.e., distal type. 

Experiments belonging to the second category, cuts made 
through the middle region of the stem. 

I. — August 5, No. 8. Stem with 17 hydranths and terminal 


1 with 5 annulj 


i 2-5 with 4 annulj 


(I with annuli 


7 with 3 annuli 


8 with 5 annulj 


9 with 2 annulj 


lu with 4 annuli 


11 with 1 annulus 


1l' with i' annuli 


13 with 1 annulus 


11 with 3 annuli 


15 with 1 annulus 


Hi with - annuli 


17 with 1 annulus 


18 with 1 annulus 

10 a.m. Cut stem lie! ween 6 and 7. 

August 8, 3:30 p.m. No. 7 lias regenerated without annuli 
immediately below the hydrotheea. 

Result: regeneration according to the distal, not the local 

II. — August 5, No. 9. Stem with 17 hydranths. 
Pedicel 1 with 5 annuli 
Pedicels 2-3 with 4 annuli 
Pedicel 4 with 7 annuli 
Pedicels 5-6 with 4 annuli 
Pedicels 7-9 with 3 annuli 
Pedicels 10-11 with 2 annuli 
Pedicel 12 with 3 annuli 
Pedicel 13-17 with 1 annulus 

11 a.m. Cut stem between 4 and 5. 

1 The stems under observation were removed from the hydrorhiza and 
rested on the bottom of flat glass dishes containing about 300 c.c. of sea- 
water. None of the colonies took food during the course of the experiment 
beyond what they could get from the original supply of water, which was 
not changed. There is no reason to suppose that the results depend upon 
the conditions to which the stems were subjected, for several observations 
of regeneration in nature indicated that results are essentially similar in 
the two cases. 

Vol. 2] Torrey. — Hydroid Differentiation and Senescence. 329 

August 8, 9:45 a.m. Hydrotheca of No. 5 with 1 anuulus 
immediately below it, supported on a segment of stem with an 
annulus next the cut (fig. 3). 

Result: regeneration according to the distal, not the local 

III.— August 5, No. 10. Stem with 20 hydranths and ter- 
minal bud. 

Pedicel 1 with 5 annuli 

Pedicel 2 with annuli 

Pedicel 3 with annuli 

Pedicel 4 with 7 annuli 

Pedicel 5 with 5 annuli 

Pedicel 6 with 7 annuli 

Pedicels 7-9 with 3 annuli 

Pedicels 10-21 with 1 annulus 

11 :10 a.m. Cut stem between 6 and 7. 

August 7, 10:30 a.m. No. 7 has regenerated in a manner 
practically identical with that of the preceding experiment. 

Result: regeneration according to the distal, not the local 

The three cases in the second category seem to show that the 
mid regions of the stems no longer possess the capacity to pro- 
duce the type of structure originally arising there, but that they 
do produce a type of structure peculiar to the latest formed 
portion of the stem. During their growth, the stems appear to 
have changed their character throughout their length. 

The facts belonging to the third category concern the cases 
of heteromorphosis from the proximal cut end of the stem. 

I. — August 5, No. 4 (see above). 

August 8, 10 a.m. Heteromorphic stem. 

August 11. Heteromorphic stem with terminal hydranth. 
Immediately below the hydrotheca are 3 annuli. The rest of 
the stem is similar in all respects to the heteromorphic stem in 
fig. 4. 

II. — August 5, No. 8 (see above). 

August 11. Heteromorphic stem similar in all respects to 
preceding (I), with the exception that development had stopped 
before a hydranth had appeared. 

330 University of California Publications. [Zoology 

III. — August 5, No. 7. Stem with 11 hydranths and termi- 
nal bud. 

Pedicels 1-2 with 6 annuli 
Pedicel 3 with 2 annuli 
Pedicel 4 with 5 annuli 
Pedicel 5 with 3 annuli 
Pedicel 6 with 1 annulus 
Pedicel 7 with 2 annuli 
Pedicels 8-11 with 1 annulus 

August 8, 3:30 p.m. Heteromorphic stem with three annular 
l>as;i] segments, a long intermediate segment and one annulus 
immediately below the terminal bud. 

IV. — August 5, No. 10 (see above). 

August 8. 4:30 p.m. ETeteromorphic stem differing but 
slightly from preceding till), surmounted by a hydrotheca 
(fig. 4). 

Latter small, and diaphragm no1 apparent. Eydranth at- 
tached to wall by a number <>t amoeboid strands which are com- 
mon along the stem but are seldom met with in hydrotheca. 

V. — August .">, No. 9 (see above). 

Augusl S. 4:lo p. in. Beteromorphic stem with 3 basal an- 
nuli. a long stem segment and a p Iy defined annulus imme- 
diately below the hydrotheca. Shoulder process on side of long 
stem segment earrying growing stalk of normal type. 

VI. — Augusl 5, No. 5 (see above). 

August 11. Heteromorphic shoo! with terminal hydrotheca. 
Three basal annuli and a very long non-annulated stem segment 
which passes directly into the hydrotheca, without an interven- 
ing constriction or annulus. 

Disregarding for a moment the stalk, the number of annuli 
occurring immediately below the hydrotheca is typical of the 
latest formed (distal) region of the normal stem in 4 of the 
above 6 eases. In the other 2 cases, the number is larger than 
the typical distal number on the same stem but is smaller than 
the proximal number on one of the stems (II). It exceeds the 
proximal number on the other stem which, however, is excep- 
tionally small (I). The tendency to develop according to the 

Vol. 2] Torrey. — Hydroid Differentiation and Senescence. 331 

distal type is therefore conspicuous so far as the pedicels are 

With respect to the stalks, it may be said that they begin 
always with one or two short segments characteristic of the 
earliest formed portion of the normal stem. These segments 
may even be shorter than the parent segment next the wound. 
It Mould seem that in this particular the regenerating stem de- 
velops according to the embryonic type. But the duration of 
this type of development is so short, lasting through the forma- 
tion of one or two segments only, that it closely resembles what 
has already been seen in the figures of regular regenerations 
in the lower regions of the hydranth-bearing zone, viz., that 
the new structures are almost invariably initiated by the for- 
mation of an annulus or part of one. 

General considerations. In seeking an explanation which 
shall simplify as well as summarize the results presented in the 
foregoing paragraphs, I think we must pass by any hypothesis 
which rests solely upon a basis of morphological determinants. 
That regeneration at a given level may not reproduce the struc- 
tural type characteristic of that level, while it does reproduce 
the type characteristic of a later level of the stem is a fact that 
is hardly simplified by the assumption of a residual germ plasm. 

So, too, does it seem improbable that the structural type is 
the result of a functional balance between an organism possess- 
ing an unmodified regenerative capacity and the conditions sur- 
rounding it. This view would necessitate a change in the envi- 
ronment between the time a polyp first appears and the time it 
is regenerated in a somewhat different form. But no such 
change is evident. The polyps half way up a stem were sub- 
jected during their development to external conditions essen- 
tially identical with those which surrounded the developing indi- 
viduals distal to yet differing from them. 

The facts, however, appear to give strong support to the 
view that the stem, instead of retaining unmodified its regener- 
ative capacity, actually loses with age its ability to produce 
structures which formerly characterized it ; and that this is 
owing to a modification of conditions within the organism, which 

332 University of California Publications. [Zoology 

govern its behavior without being necessarily a part of it. These 
conditions are probably chemical in nature, intimately con- 
cerned with the metabolism. A destruction or addition of sub- 
stance or substances in the course of the development is readily 
conceivable as the efficient cause of the structural modification. 
The relation between internal and external conditions is under 

The resemblance of the phenomena of colonial differentiation 
in C. bakeri to the phenomena of senescence is so strong as to 
suggest a similar interpretation for both. 

The experiments will be continued. 

September 1, 1905. 


Vol. 2, No. 7, pp. 333-340, Figs. 1-5 December 13, 1905 







Iii a former paper ( :04)/ it was shown that Corymorpha pos- 
sesses an unusually wide range of activities for a hydroid. It 
responds to mechanical and thermal stimuli, to chemical stimuli 
that produce their effect as mechanical irritants, not as odorous 
substances, and to gravity. In the movements of the stem, it 
resembles such naked forms as Hydra, Claim and Hydractinia. 
The stem is everywhere sensitive to mechanical stimuli, bending 
from side to side or shortening under their influence. Such re- 
actions are due to the contractions of longitudinal muscle fibers 
which are situated in the usual fashion deep in the ectoderm. 
A lengthening of the stem may be caused wholly or in part by 
the circular endodermal musculature. 

The long proximal tentacles, about twenty-four in number, 
may respond singly to direct mechanical stimulation or to a 
stimulus applied either to another tentacle or a distant portion 
of the stem. In all cases of effective stimulation, direct or indi- 
rect, they bend invariably toward, never away from the mouth. 
The reaction is rapid ; the recovery, which ends in the resumption 
of the expanded condition, is slow. 

The initial reaction of the distal tentacles to all effective 

stimuli is, on the other hand, always away from the mouth ; 

1 Biological Studies on Corymorpha. I. C. palma and Environment. 
Jour. Exp. Zoology. I, No. 3, 1904. 

334 University of California Publications. [Zoology 

after Avhich, strong irregular movements toward and away from 
the mouth appear. The proboscis, which carries the distal ten- 
tacles to the number of about fifty in a crown around its summit, 
may react to mechanical stimuli, directly or indirectly applied, 
by bending in the direction of the stimulus. 

These reactions of tentacles and proboscis constitute an effi- 
cient prehensile mechanism. A small object which happens to 
stimulate a proximal tentacle on its oral side is at once swept 
toward the mouth. The distal tentacles, which may be carried 
toward il by the bending of the proboscis, then move outward. 
meeting and transporting it by a subsequent inward movement 
to the mouth. 

All the motions thus far described arc due to muscular con- 
tractions. Locomotion is produced, as in Hydra, by the activi- 
ties of amoeboid cells at the base of the stem. And the circu- 

],-iti »f fluids i;i the coenosarcal canals is accomplished by the 

usual ciliary action, supplemented by the expansions and con- 
tractions of the proboscis and stem. 

Aside from these types of motion, the stem exhibits a marked 
geotropism assuming when at rest a vertical position. This 
orientation does not appear to be dependent in any way upon 
muscular activity. The behavior not only of the stem as a whole 
lint of pieces of one-eighth or one-tenth its length from the base 
or various other regions, indicates that the stem is everywhere 
sensitive to the stimulus of gravity and furnished with an effi- 
cient mechanism Eor bringing about a response. This mechanism 
is expressed, 1 believe, in the large, highly vacuolated cells of 
the endodermic axis which forms the core of the stem. Changes 
of orientation, according to this view, are produced by relative 
changes in the turgidity of such cells on opposites sides of the 
stem. Whether the orientation is to be reckoned as a contrac- 
tion phenomenon, though this is probable, cannot be said defi- 
nitely at this time. It is indeed a fact that the axial cells may 
not only decrease in size, as when the stem is shortened, but also 
increase in size, as when the stem is lengthened without loss in 

Vol. 2] Torrey. — The Behavior of Corymorpha. 335 

Two conclusions which were formerly held, later experiments 
have shown to have been founded on data which were mislead- 
ing owing to the conditions of experimentation. The first was 
that a change in the polarity of a region is accompanied by a 
change in the reactions of the axial cells in this region. The 
second was that, regardless of the point at which it is supported, 
whether proximally or distally, the stem would orient itself ver- 
tically, distal end uppermost. These results were obtained on 
individuals which had been kept in the laboratory during the 
warm days of summer for a week or ten days, and did not be- 
have with the constancy or the precision which characterize the 
actions of the individuals observed last March. The latter were 
used for experimentation immediately upon their capture, and 
were kept under conditions which permitted a vigorous, healthy 

With regard to the first conclusion, it may be said that while 
such a change of polarity as heteromorphosis of the proximal 
end of a stem segment would be accompanied by an upturning 
of this end, the result would not be achieved by a change in the 
reactions of the axial cells in this region. The essential factor 
lies -rather in the relation of the region in question to the sub- 
stratum. In studying the regeneration of Corymorplia, the ob- 
servation was frequently made that from pieces resting on the 
floor of the aquarium, cut from the distal half of a stem, U- 
shaped figures would be formed, fastened to the floor by the 
loop, the two arms extending vertically upward, each crowned 
with sets of developing tentacles. By the side of these hetero- 
morphic pieces were many others fastened to the substrate by 
one end, which possessed an incipient holdfast, the other end 
developing tentacles. In both cases, the behavior of the axial 
cells was constant ; the cells nearer the center of the earth were 
relatively larger than those on the other side of the stem. The 
proximal ends of the pieces which developed holdfasts remained 
lowermost merely because they were adhesive and clung to the 
substrate, which the distal ends could not do. 

This fact appeared distinctly in the results of my recent 
experiments bearing directly upon the second conclusion stated 

336 University of California Publications. [Zoology 

above. I had found previously that when stems relieved of 
hydranths and the weight of sand grains adhering to the hold- 
fasts were supported at their distal ends, they would assume 
the normal vertical orientation, proximal end down. Last March, 
the experiment was repeated many times, with special precau- 
tions against errors. Fresh animals were used, and their distal 
ends were fastened by a single loop to a rigid thread. The sup- 
port thus afforded was secure and ample for various movements. 
In every case, without regard for the amount of holdfast pres- 
ent, the unsupported proximal end travelled upward and came 
to rest only when the stem had reached a position approximately 
vertical, bu1 upsidt down. 

Comparing this result with the movements of stems attached 
by the proximal end. either resting on or hanging from the sub- 
strate, il is obvious that no reversal of behavior in the axial cells 
need be assumed. The variable fa. -lor is connected with the point 
of support. 

By the foregoing observations, Corymorpha seems to be re- 
moved from the category of animals for the elucidation of whose 
behavior none of the familiar mechanical explanations of geo- 
tropism seem to apply. Its geotropic reactions appear now to 
accord with the theory which Davenport formulated with refer- 
ence to the geotropism of free-swimming organisms. There is 
a difference between the resistance encountered by the stem as it 
moves upward friction plus weight) and the resistance it en- 
counters when it moves downward (friction less weight). This 
difference is expressed in the stem by a tension on its upper 
side when it is inclined in any degree from the vertical. There 
is no tension on the Lower side of the stem unless it be hanging 
downward at some angle; in which case i1 never equals the ten- 
sion mi the upper side except when the stem is vertical, its posi- 
tion of rest. 

The evidence formerly presented in favor of the view that 
the axial endoderm cells, and not muscles, govern the geo- 
tropic orientation of the stem, has been strengthened to some 
extent by a further experiment. In my original experiments, 
cuts were made at frequent intervals half way through the stem 

Vol. 2] 

Torre ij. — The Behavior of Corymorpha. 


on one side, or alternately on both sides. The stem was then 
laid on the floor of the aquarium, a cut side uppermost. In one 
to two hours, the stem had assumed an erect position. The cuts 
were intended to destroy the effectiveness of the muscles of the 
stem, leaving the column of axial endoderm cells intact. The 
muscles as a whole were weakened by the cuts. And the slowness 
of the reaction, coupled with the fact that the orientation was 
accomplished before the wounds closed, strengthened the view 
that muscles were not concerned in the reaction. 

Other facts supporting the same conclusion have since been 
obtained. A piece was removed from a stem by two cuts which 

formed with each other an obtuse angle (fig. 1). On the cut 
faces, the axial endoderm was exposed. The rounded surface 
representing the original stem wall was still covered with ecto- 
derm, provided with longitudinal muscles. Figs. 2 and 3 were 
drawn from this piece when it was in contraction and expansion 
respectively. It will be noticed that in the contracted condition 
(fig. 2), the course of the longitudinal muscle fibers is no longer 


University of California Publications. [Zoology 

straight ; the angle made by the cut surfaces is greater than it 
was at first ; there is a tension factor on the side of the stem 
opposite the layer of muscles which tends to draw the remotest 
edges of the muscle layer toward each other against the pull 
of the muscles themselves. In the expanded condition (fig. 3), 
though the piece lengthens, it bends strongly toward the cut 
surfaces. The latter now form an angle greater than 200 de- 
grees. At the upper end of the figure, the tip of the piece has 
curled sharply over toward the wound. The tension is still 
more apparent here than in fig. 2. On the assumption that the 
axial endodenn cells have lessened their volume on the side to- 
ward the wound relatively to the volume of the axial cells on 
the side away from the wound, the configurations shown in figs. 
2 and 3 are intelligible. The axial cells themselves did not push 
out between the edges of the wound but gave every indication of 
being under rest raint. 

When long. thin, oblique slices are cut from a stem as in 
fig. 4. it follows inevitably that the narrow wedge-shaped ends 
curl toward the cut surface, as in fig. 5. The relatively de- 
creased turgidity of the axial cells next the wound appears to 
account for this condition also. 

In the light of these facts, it becomes clear that the axial 
endoderm cells under certain conditions are capable, by chang- 

Vol. 2] Torre y. — The Behavior of Corymorpha. 339 

ing their turgidity, of producing movements in the stem com- 
parable with those which are due to stimulation by gravity, 
though the evidence does not demonstrate that gravity itself 
affects them, directly or indirectly. But they are active, not 
passive elements, which increases the probability that they may 
be concerned in the geotropic response. 

If it be true, as seems probable, that the axial cells do govern 
the geotropic response, Corymorpha stands alone among the met- 
azoa in possessing a tropic mechanism distinct from the body 
musculature. In a recent paper, Holmes ( :05) has shown that 
Ranatra exhibits uncommonly clear cut reactions to light. Here 
the tropic mechanism involves the same sense organ, the same 
musculature and to some extent at least the same nervous ele- 
ments which serve in responses to other classes of stimuli, in- 
ternal and external. Though Ranatra is an unusually favorable 
object for the analysis of the tropic mechanism, the confusion 
of various factors operating over the same lines renders an ade- 
quate analysis impossible at present. The presence of a geo- 
tropic mechanism distinct from the musculature in Corymorpha 
simplifies the problem. And I am led to believe, further, that 
the simply organized nervous system of the hydroid need play 
no part in the geotropic reaction. The axial endoderm cells 
probably change their volume under a given tension which is 
applied directly to them. Each cell may be considered a unit, 
not necessarily dependent in its actions upon its neighbors, act- 
ing with them only in so far as they may be subjected simulta- 
neously to similar stimulation. 

The response of Corymorpha to gravity is strikingly similar 
to the negative geotropism of the caulicles of plant seedlings, 
which is unquestionably dependent upon volume changes in the 
stem cells. The latter are again strikingly similar in structure 
to the axial cells of Corymorpha. There appears to be in the 
hydroid mechanism no more opportunity for a pleasure-pain 
type of reaction than among the plants; and there is no sign 
of it in the geotropic movements of the stem. Neither are there 
signs of trials and errors. The movement of the stem is very 
gradual, very definite, very direct. I know of no animal which 

340 University of California Publications. [Zoology 

more closely approximates the plant in structure and tropic re- 
sponse. If the behavior of the one be explicable on the basis of 
direct reactions to stimuli, of the reflex type, I do not see how 
the behavior of the other can be excluded from a similar inter- 


Vol. 2, No. 8, pp. 341-368, Pis. 17-19 January 6, 1906 









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Vol. 2, No. 8, pp. 341-368, Pis. 17-19 January 6, 1906 







The investigations of the plankton of the Pacific at the San 
Diego Station during the past three years has brought to light 
a number of species belonging to the family Peridinidae which 
do not conform to any known genus. They all present in com- 
mon a number of characters which call for the establishment of 
a new genus for the reception of the several species represented. 

To this genus also belong seven species described by Murray 
and Whitting ( '99) from the tropical Atlantic as members of 
the genus Peridinum to wit: Peridinium Jiindmarchii, P. milnt ri, 
P. blackmani, P. leiorhynchum, P. trirostre, P. tripos, and P. 
doma. Gonyaulax triacantha Jorg. is also to be referred to this 
genus. All of these species except P. trirostre and Gonyaulax 
triacantha have been seen by me and a fuller discussion of their 
structure will appear elsewhere. In the following pages the brief 
discussion of each is based upon Murray and Whitting 's figures. 

I am indebted to Miss E. J. Rigden, assistant in the summer 
of 1904 at the San Diego Marine Biological Station, for some of 
the sketches utilized in the plates accompanying this paper and 
also for the skill and thoroughness of her examination of the 

342 University of California Publications. [Zoology 

plankton which brought to Light the most of the species here dis- 
cussed. The types and cotypes of the species here described for 
the first time are in the collections of the University of California. 


gen. now 

The form of the theca resembles that of Peridinium in the 
presence of a median or somewhat postmedian girdle which en- 
circles the theca at its greatest diameter and in two antapical 
horns, always directed posteriorly or nearly so. The posterior 
ma ruin or list of the transverse furrow is suppressed or feebly 
developed in comparison with the anterior one. especially at its 
distal end, while the anterior one. as it' in compensation, is often 
excessively developed. The suture lines are demonstrated with 
great difficulty and some uncertainty in most of the species. 
The plates are as follows: three apicals, one left intercalary, six 
premedians, seven postmedians, one furrow plate, and three (?) 
antapicals, as shown in the accompanying text figures. 

On the ventral face about midway between the apical and 
flagellar pores is a small pit or pore-like area in the mid ventral 
suture. An actual opening in this area has not been demon- 
strated. In the suppression of the posterior border of the girdle, 
in the number and arrangement of the plates and in the presence 
of the ventral pit on the epitheca Heterodinium differs from 
Peridinium, although in form and general appearance species of 
the two genera strongly resemble each other. 

In some species and possibly generally in the genus there is 
a decided asymmetry to the theca brought about by a torsion of 
the body on the main axis in clockwise direction, looking from 
the posterior toward the anterior end. This is especially notice- 
able in the scoop-shaped forms such as //. scrippsi. 


The following is a more detailed description of the characters 
found in the genus. The theca is expanded in the equatorial 
region, with more or less well marked dorso-ventral flattening 
and ventral excavation near the flagellar pore. It is spheroidal. 

vol. 2] Kofoid. — On Heterodinium. 343 

ellipsoidal, elongated, rotund, flattened, or even scoop-shaped. 
The length always exceeds either diameter, and the transdiam- 
eter at the girdle equals and more often exceeds the dorso- 
ventral one. The greatest transdiameter is usually at the girdle 
but in some species the epitheca or hypotheca may exhibit a 
slightly greater diameter. The greatest dorso-ventral diameter 
is at the left of the flagellar pore. 

The epitheca is usually not contracted to an apical horn 
though in some species a short horn is present, and in others 
the elongated epitheca tapers gradually from the girdle to the 
apical pore with more or less concavity of the lateral margins. 
The anterior end of the epitheca is more often broadly rounded, 
being dome-shaped in the rotund species and like the end of an 
ellipsoid or even scoop-shaped in the flattened species. In some 
instances, as in H. blackmani, the epitheca is rotund at the girdle 
but flattened distally. The altitude of the epitheca is usually 
less than the transdiameter and exceeds it in only a few cases as 
in //. blackmani and II. liindmarclii. 

The ventral face of the epitheca is flattened and somewhat 
excavated, slightly in rotund species, more deeply and exten- 
sively even to the lateral margins in the dorso-ventrally com- 
pressed forms. The mid ventral face is marked by the slightly 
sinuous suture ridge which runs from the flagellar pore to the 
apex and. bears midway a pit or pore-like area, a characteristic 
structure in the genus. This varies greatly in distinctness and 
in the breadth of the widened smooth suture ridge in which it is 
placed. The apical pore is inclined, even as much as 10° in some 
species, to the right and is usually well defined though rarely 

The hypotheca is usually subequal to the epitheca, being- 
longer in H. milneri and //. spJtaeroideum, and shorter in H. 
blackmani. The posterior end may or may not show a bifurca- 
tion into antapical horns. It may be broadly rounded, dome- 
shaped and without any antapical differentiations as in H. 
sphaeroideum and //. doma, with mere spinules with or without 
lists as in H. milneri and H. murrayi, with slight median bifur- 
cation as in H. ivhittingae or with typical elongated antapicals 

344 University of California Publications. [Zoology 

as in H. blackmani. In the form of hypotheea Hcterodinium 
thus exhibits a development parallel to that found in the allied 
genus Peridinium. The ventral face of the hypotheea is chan- 
neled by the longitudinal furrow plate which in many species 
extends anteriorly so that it indents the epitheca above the 
flagellar pore. 

The girdle is usually submedian in position, though preme- 
dian in some species as for example in //. milneri, or postmedian 
in others as in //. scrippsi. The girdle is much more oblique in 
the flattened than in the rotund forms. In //. whittingae its 
plane is inclined ventro-posteriorly at an angle of 45° to the 
axis. The girdle in all species thus far observed forms a de- 
scending right spiral with a displacement accelerated distally 
and amounting to 1-3 times the width of the furrow. The must 
characteristic feature of the girdle is its incompleteness distally 
and the absence or slight development of the posterior ridge. 
The furrow is bounded anteriorly by a heavy overhanging ridge 
which in species thus far observed is no1 a ribbed fin or list but 
a heavy projection of the thecal wall. The posterior border is 
formed by a less salient ridge which becomes less prominent dis- 
tally and often diverges more widely from the anterior ridge 
towards its distal end . The feature of a more or less deficient 
posterior margin of the girdle is a constant character in all 
species thus far observed save the imperfectly known II. sphae- 
roidi mn which has, however, the ventral plates at least (the 
dorsal ones are not known) of the genus. 

The transverse furrow is indented in the thecal wall and the 
flagellar pore is found at its proximal end. 

The thecal wall is made up of discrete plates, which, however, 
are much less easily separated and much less clearly denned than 
they are in other genera of the family, as for example in Peridi- 
nium. The sutures are marked by flattened ridges or bands or 
smooth tracts in which the cleavage line may be traced in some 
eases in young individuals. The suture bands often have a 
secondary reticulation of minute polygons on their surface and 
are best seen on a deep focus. They are differentiated on the 
inner as well as the outer thecal surface. The epitheca consists 

Vol. 2] 

Kofoid. — On Heterodinium. 


of ten plates (figs. A and B) arranged as follows: three apicals 
about the apical pore, of which one (1) is a wide dorsal plate 
covering the dorsal half of the anterior end, and the other two 
(2, 3) are ventral and are separated from each other by the mid- 
ventral suture which runs from the apical to the flagellar pore. 
Between the dorsal and the two ventral plates are lateral sutures 
which in subgenera Platydinium and Eiiheterodinium become 
very heavy and much more prominent than any other sutures 

Fig. A. — Ventral view of Heterodinium scrippsi showing thecal plates; 
apicals, 1-3; premedians, 5-10; left intercalary, 4; postmedians, 11- 
17; antapicals, 18-20. X 450. 

Fig. B. — Dorsal view of same. X 450. 

of the theca. In some cases as in H. scrippsi and H. blackmani 
these lateral ridges are doubled, suggesting a narrow compressed 
plate, but in the more rotund species they are not doubled and 
there is no suggestion of the presence of such a plate. I there- 
fore conclude that they are merely doubled margins and are not 
to be regarded as the margins of degenerate plates. 

Anterior to the girdle is a premedian series (figs. A, B, 5-10) 
of 6 plates of which two are ventral (5, 10), two dorsal (7, 8), 

346 University of California Publications. [Zoology 

and one each right (9) and left (6). They are not as a rule 
symmetrically placed because of the considerable irregularity in 
their size and especially because of the disturbing effect of the 
left intercalary plate (4) -which is found between premedians 
5, (i. 7 (or 6 and 7) and apicals 1 and 3. This intercalary plate 
is often small and in compressed species is not easily found. It 
is shown clearly in H. blackmani, but is merely suggested or not 
shown at all in the rest of Murray and Whitting's ('99) species. 
Its demonstration in all species carefully examined by me leads 
me to expect it in others especially since they usually show the 
tilting of the apical pore to the right, the slight shoulder on the 
left marginal outline and the asymmetrical arrangemenl of tbe 
premedians which attend its presence in species in which it has 
been demonstrated. It is greatly reduced in size in flattened 
species such as //. whittingae, and it is certainly possible that 
this plate may be entirely suppressed in some of the flattened 
species of the subgenus Platydinium, though no conclusive evi- 
dence to that effect is at hand. 

The ventral face of the epitheca of the flattened species is 
formed by the two ventral-apical plates (2, 3) and the two 
ventral-premedians (5, 10) bu1 in the more rotund forms the 
lateral-premedians (6, 9) are partially exposed in a ventral view. 
The left intercalary in the flattened species is dorsal in position. 
but in the rotund forms as //. murrayi and //. doma it appears 
on the left shoulder in the ventral view, and may be shifted 
dorsalwards so that it does not touch the ventral premedian 5. 

The girdle plate has the form of a trough-like bund as in 
Peridiniwm and appears to be variously subdivided by reticular 
ridges some of which may be suture lines. 

The plates of the hypothecs are less clearly defined than 
those of the epitheca. especially on the ventral face. There are 
seven postmedians (11-17) adjacent to the girdle, 3 dorsal 
(13-15) and 4 ventral (11. 12, 16, 17) one of which (16) appears 
to extend to tbe tip of the right antapical without subdivision 
into anterior postmedian and posterior antapical moieties, ex- 
cept possibly in //. sphaeroidewm. The separation of the adja- 
cent postmedian (17) from the ventral median antapical is 

Vol. 2] Kofoid. — On Heterodinium. 347 

often feebly expressed. The antapieal series includes 3 plates, 
a single dorsal (18), a median ventral (19), and a left ventral 
(20). The distal end of the posterior list of the girdle usually 
descends on the suture between the right ventral and right latero- 
ventral postmedians and continues to the tip of the right antap- 
ieal horn. 

The boundaries of the plates on the right ventral face of the 
hypotheca are subject to much obscurity and considerable varia- 
tion in location. The relations of the midventral plate (19) to 
the adjacent postmedian (17) and to the longitudinal furrow 
are subject to considerable variation in connection with the vary- 
ing degrees of development of the posterior list of the girdle. 

The longitudinal furrow is subject to considerable variation 
in length. It is relatively short in some species as in H. rigdenae, 
H. sphaeroideum, and H. hindmarchi where it is less than two- 
thirds the length of the hypotheca. In other forms as II. tri- 
rostre and H. murrayi it reaches the antapieal border. It is 
without high membranous lists as a rule though one appears in 
II. murrayi. 

The thecal wall is thin and hyaline and universally, except 
in apparently young individuals, reticulate with more or less 
irregular polygons formed by thickened ridges on the outer sur- 
face. In some species, as in II. scrippsi, and perhaps in certain 
stages of growth of other species these polygonal boundaries be- 
come so prominent as to obscure suture lines, as may be seen in 
Murray and Whitting's ( '99) figures of H. trirostre, H. murrayi, 
and 77. hindmarchi. 

These reticulations are often quite regular as in H. blackmail i 
and along the lateral margins of the epitheca of //. scrippsi, or 
very irregular as on the ventral and dorsal faces of H. scrippsi 
and in H. trirostre; they may be relatively large as in H. murrayi 
or small as in H. sphaeroideum , with very heavy ridges as in 
H. trirostre or but faintly outlined as in H. milneri, or forming 
but a delicate network as in H. blackmani. In young individuals 
they may be entirely lacking and the presumption is that in gen- 
eral, individuals with partially or feebly developed reticulations 
have not as yet reached the stage of completed formation of the 

348 University of California Publications. [Zoology 

theca. The reticulations are found also on the girdle plate and 
on the girdle lists and along suture lines there are frequently 
rows of smaller polygons. In but a few cases as in H. murrayi 
and //. doma are enclosing ridges so thickened by the filling in 
of the angles as to leave a subcircular central area. Each reticu- 
lation lias typically one centrally located pore. In some species 
with coarse reticulations there are several pores in a single area, 
and frequently in all species there are minor irregularities in 
the number and position of the pores. Small polygons fre- 
quently lack the pores. The reticulation is evidently formed on 
the outer surface of the theeal wall by plasma which is extruded 
ili 101 iuIi tlie pores, for the polygons bear ;i definite relation to 
the arrangements and distribution of the pores. 

The protoplasmic contents of the theca are usually hyaline 
and colorless, and often only partially lill the interior of the 
theca. The nucleus has the usual ellipsoidal form with beaded 
chromatin reticulum and lies near the center of the protoplasmic 
mass no1 far from the flagellar pore. It is small and is found 
with difficulty. No instances of diffuse reddish coloration often 
seen in Peridinium and Pyrophacus have been noted as yet in 
Heterodinium. Chromatophores are entirely absent in some in- 
stances, in others they are massed in spheroidal chromospheres 
of pale greenish yellow or deep cadmium orange color. In some 
cases the chromatophores are peripheral in location and of va- 
rious forms. Vacuoles and pusules of varying form and distribu- 
tion have been observed in the cell contents. 

The dimensions of observed species are like those of Peridi- 
nium. The Largest species thus iiw recorded appears to be H. 
blackmani which has a length of 225/x. and transverse and dorso- 
ventral diameters of 135//. and 16(hx respectively; the smallest 
appears to be H. spJtaeroich ion with a length of only 42/x, and 
transdiameter of 39/x. 

The distribution of this genus from species thus far pub- 
lished appears to be limited, in the main, to warmer seas as shown 
in the following- table : 


Temperature C. 

7°-30° N. 


34°-39° N. 


19°-39° N. 


26° N. 


19°-39° N. 


14°-28° N. 


29°-31° N. 


San Diego 

32.7° N. 


Vol. 2] Kofoid. — On Heterodinium. 349 

H. blackmani 
H. doma 
H. hindmarchi 
H. trirostre 
H. leiorhynchum 
H. murrayi 
H. milneri 
H. sphaeroideum 
H. rigdenae 
H. scrippsi 
H. whittingae 
H. inaequale ] 

H. triacantha 55°-81° N. 

Their vertical distribution is not known. At San Diego no 
individuals have been taken in the many surface catches of the 
tow nets made during the past few years. They have been found 
only in the vertical catches in from 165 to 40 fathoms to the 
surface. Murray and Whitting's ('99) species were all appar- 
ently from plankton collected by filtering water from ship's 
pumps and therefore taken some 2-3 fathoms below the surface. 
The absence of chromatophores or their aggregation in chromo- 
spheres observed in individuals taken at San Diego is suggestive 
of occurrence in deep water with diminished light. The extreme 
hyalinity of some species is also indicative of a deeper habitat. 
The excessive development of the reticulum on the theca, and 
the asymmetry are evidently adaptations for flotation, on the 
one hand by increase of friction surface, which is at least doubled 
in the more rugose forms, and on the other by giving a spiral 
course to any passive descent of the organism due to gravity, and 
thus prolonging its existence in the upper strata. 

This is a genus of somewhat aberrant structure and is repre- 
sented by relatively very few individuals in comparison with 
those of Ceratium and Peridinium. I regard it as a degenerate 
form unable to maintain itself at the surface and for some reason 
deficient in reproductive vigor. In comparison with the number 
of individuals observed the number of species is large. The 
known species are all well defined and observations on different 
individuals do not indicate as yet any noticeable intergradation. 

350 University of California Publications. [Zoolooy 

The nearest allies of this genus are plainly in the family 
Peridinidae, though it shows no marked structural affinities to 
any particular genus. The form cycle found in its species re- 
sembles that of Peridinium but its thecal plates are entirely 
different. The midventral diamond-shaped plate of the epitheca 
so characteristic of Peridinium is entirely lacking in Heterodi- 
iiiiim, unless indeed the slight midventral expansion on the 
suture line be taken to represent a degenerate 1 midventral which 
seems improbable. The excessive development of reticulations 
on the surface of the theca approaches that in Protoceratium 
but this genus appears to lack the midventral pit on the epitheca, 
and has ;i narrow transverse furrow which is complete distaUy. 
Its plates (see Scbutt ('96)) are not known and it may prove 
to have a closer relationship to Heterodinium when these are 
definitely determined. 

The midventral pit on the epitheca of Ileterodinium resembles 
the so-called ••pore" iii Poroceratium gravidum (Gourret) but 
hears a different relation to the thecal plates and is probably not 
a homologous structure. In Poroceratium the "pore" lies near 
the middorso-ventra] Line in the middle of the dorsal and ventral 
apical plates, whereas in Heterodinium it lies in the suture be- 
tween the two vent ral apicals. 

The generic distinctness of Heterodinium is thus beyond ques- 
tion and it belongs with Ceratium, Peridinium ami Protoceratium 
in the sub-family Ceratiinae. 


Sphaerodinium sabgen. nov. 

Body spheroidal, antapical horns not present or only slightly 
developed as spines. Epitheca rotund without stout lateral 


1. With no antapical horns or spines 2 

1. With antapical spines 3 

'_'. Outline smooth, sutures faint, reticulations minute..i?. sphaeroideum 

'2. Outline subangular, sutures prominent, reticulations coarse. Z7. doma 

3. No apical horn, thecal markings faint H. milneri 

3. Short apical horn, thecal reticulation prominent H. murrayi 

3. Long apical horn H. triacantha 

VoL - 2 1 Kofoid. — On Heterodinium. 351 

Euheterodinium subgen. nov. 

Epitheca dorso-ventrally compressed, with straight, convex 

or concave sloping lateral margins which are usually thickened 

and have doubled ridges between the lateral margins of the 

apical plates. Antapical horns well developed. Girdle not very 


1. Epitheca with strongly convex sides, apex broadly rounded 

H. inaequale 

1. Epitheca with straight or concave sides, apex not broadly rounded ... '2 

2. Epitheca low, its altitude about one-half the transdiameter 

H. rigdenae 

2. Epitheca high, tapering, nearly equal to the transdiameter 3 

3. Left antapical bifurcated H. trirostre 

3. Left antapical not bifurcated 4 

4. Eeticulations very coarse, scantily developed H. leiorhynchum 

4. Eeticulations subregular, very delicate H. blaclma?ii 

4. Reticulations medium sized, very heavy H. hindmarclii 

Platydinium subgen. no v. 

Epitheca dorso-ventrally compressed and hollowed out ven- 

trally, scoop-shaped. Lateral margins convex, not contracted to 

an apical horn. Girdle very oblique dorso-ventrally. Antapical 

horns present. 

1. Antapical horns divergent H. scrippsi 

1. Antapical horns convergent H. whittingae 

Heterodinium sphaeroideum sp. no v. 

PI. 3, fig. 15. 

A minute symmetrical species of spheroidal form without 
apical or antapical horns. The body is spheroidal or broadly 
ellipsoidal, the length 1.1 transdiameters. Dorso-ventral diam- 
eter equal to transdiameter. Epitheca a low dome, its altitude 
0.4 transdiameters. Hypotheca exceeding the epitheca, elon- 
gated hemispherical, its altitude 0.6 transdiameters, with broadly 
rounded symmetrical antapex. Girdle premedian, transverse 
furrow indented, posterodexiotropic with very slight displace- 
ment scarcely 0.2 its width, its anterior and posterior lists equal 

352 University of California Publications. [Zoology 

and the latter not deflected distally, both formed by sharp pro- 
jecting ridges of the thecal wall. Longitudinal furrow short, 
its length less than 0.5 distance to the postmargin, broad and 
shallow, its distal two-thirds enlarged. 

Thecal plates imperfectly known. Ventral plates of typical 
number and arrangement except that the right ventral post- 
median (16) is not continued to the postmargin but appears to 
be divided into post median and antapical moieties. Suture lines 
faint, bordered by smooth structureless zones. No prominent 
lateral ridges. No lists or spines. Thecal wall minutely and 
faintly reticulate with subregular polygons with centrally located 
pores. Polygons relatively very numerous. 

Plasma dens.', heavily vacuolated, chromatophores irregular, 
peripherally located, greenish yellow; nucleus near flagellar pore. 

Dimensions- length, 42/*; transdiameter, 39/*; width of fur- 
row, 4-5/*; diameter of polygons, 2-3/x. 

Taken once in vertical haul from 165 fathoms off San Diego 

ill .1 line. 

Although this organism does no1 have the deficienl girdle 

found in other species of the genus its thecal plates, in so far 
as they are known, are those of Heterodinium. 

Heterodinium doma (Murr. et Whitt.). 

Peridinium doma Murray and Whitting ('99), p. 327, PL 30, fig. 3. 

Plainly belongs to Heterodinium because of the clearly shown 
ventral pit in the central expansion of the median ventral suture. 
The plates are only partially shown but in the one view (ventral) 
given they conform to Heterodinium so far as shown. The girdle 
and furrows are also typical. 

The species is characterized by the spheroidal form, sub- 
median girdle, broadly rounded apex, entire absence of antap- 
icals, median reticulations of subregular polygons and somewhat 
salient suture ridges. 

Reported from the warm temperate Atlantic between 34°- 
39° N. 

VoL - 2 1 K of oid. — On Heterodinium. 353 

Heterodinium milneri (Murr.. et Whitt). 

Peridinium Milneri Murray and Whitting ('99), p. 327, PI. 29, figs. 3a, b. 

The characteristic Heterodinium structures are not clearly- 
shown in Murray and Whitting 's figures. There is only a sug- 
gestion of the midventral suture of the epitheca and a markedly 
deficient posterior list of the transverse furrow. The ventral pit 
is lacking and the plates are incompletely shown. 

The species is characterized by its spheroidal rotund body, 
premedian girdle with wide displacement and considerable over- 
lap of the ends of the transverse furrow, wide zones along suture 
lines free from reticulations, and coarsely reticulated plates. It 
is closely related to H. murrayi. 

Reported from tropical Atlantic in 29°-31° N. 

Heterodinium murrayi nom. nov! 

Peridinium tripos Murray and Whitting ('99), p. 327, PI. 30, figs. 4a, b. 
non P. tripos (Midler), Ehrenberg ('33), p. 272= (Ceratium tripos). 

The specific name tripos must be rejected as it was previously 
introduced into the genus Peridinium by Ehrenberg 's ('33) 
transfer of Cercaria tripos of 0. F. Miiller (1786) to the genus 
Peridinium. As figured by Murray and Whitting ('99) this 
species shows almost none of the generic characters except the 
very deficient posterior list of the transverse furrow. The ven- 
tral pit is questionably figured and no trace of the plates is 
shown. The only evidence of the presence of the left intercalary 
is the shifting of the apex to the right. 

The species is characterized by its small size, rotund body, 
large and few subregular polygonal reticulations with a coarse 
mesh which hide the sutures and cover the whole theca. The 
girdle is premedian and the transverse furrow is much displaced 
and has considerable overhang. The apex is somewhat con- 
tracted and the antapicals bear two short finned spinules on the 
left and one on the right. The anterior list of the transverse 
furrow is membranous. There are no antapical horns. 

Reported from the tropical Atlantic in 14°-31° N. 

354 University of California Publications. [Zoology 

Heterodinium triacantha (Jorg). 

Gonyaulax (?) triacantha Jorgensen ('99), p. 35. 
Ceratium hypcrboreum Cleve ('00), pp. 14-15, PI. 8, fig. 5. 
Gonyaulax triacantha, Paulsen ('04), pp. 21-22, fig. 5. 

This form appears to belong to Heterodinium by reason of 
the reticulated thecal wall, the midventra] suture of the epitheca 
deflected to the left, the widened distal end of the transverse 
furrow, and the longer right antapica! spine. There is also 
some indication thai the distal end of the posterior list of the 
girdle is continued in the suture on the right side of the hy- 
potheca. None of the figures shows the ventral area or pit, or 
the thecal plates in full. In so far as they are indicated in Paul- 
sen's ( '04) figures, they conform to those of the genus Hetero- 
di ninni. There are difficulties in reconciling Cleve's ('00) figures 
with each other, and with those of Paulsen I '04) as well as with 
Jorgensen's description, probably (\\\c to the fact, that, as Paul- 
sen suggests; Cleve's figure is reversed, i.e., it is a view of the 
ventral face as viewed through the body from the dorsal face. 

This species probably belongs in the subgenus Sphai rodiniwn, 
though it does not possess a spheroidal body. It is characterized 
by the absence of antapica] horns and posl indentation, concave 
sides of the epitheca. the developed apical horn, and the three 
antapica] spines. 

Dimensions: — length, Ti' 84/i. ; transdiameter, about 50/*; 
dorsal-ventral, about 45/t. 

Reported from coasts of Norway and Iceland. 

Heterodinium inaequale sp. nov. 

PL 18, figs. 9, 10. 

This is a small subpentagonal species with rotund epitheca 
and unequal widely separated antapicals. 

The body in face view is subpentagonal, the two anterior mar- 
gins are quite convex, the left posterior nearly straight, the right 
slightly convex and the postmargin between bases of the antap- 
icals is concave. The length is 1.2 and the dorso-ventral diam- 
eter 0.75 times the transdiameter. The epitheca is low dome- 

Vol. 2] Kofoid. — On Heterodinium. 355 

shaped, compressed dorso-ventrally, its altitude (ventral) is 0.7 
transdiameter. No apical horn is differentiated and though com- 
pressed dorso-ventrally it is not thinned down to a sharp edge 
at the doubled lateral sutures. The ventral face is scarcely ex- 
cavated. The broad midventral suture runs from the longi- 
tudinal furrow to the apical pore swerving towards the left at 
the ventral pit. 

The hypotheca is rotund, its altitude (mid-dorsal) 0.6 trans- 
diameter. The antapicals are very unequal, the right is about 
one-half the length of the left and is abruptly incurved to an 
acute tip. The left is not incurved and is somewhat tapering. 
Its length is 0.3 transdiameter. The postmargin between the 
antapicals is slighly concave and is 0.4 transdiameter in length. 

The plates are typical in number, the left intercalary being 
confined to the dorsal face. The dorsal premedians are very 
low, scarcely exceeding the girdle in width. The dorsal post- 
medians on the other hand are unusually long, and the posterior 
angle of the right one projects slightly beyond the margin. 

The girdle is narrow and slightly oblique (15° postero-ven- 
trally) to the equatorial plane. The transverse furrow is postero- 
dexiotropic with a displacement of its own width. It is scarcely 
indented, the thecal wall forming a slight anterior ridge, and 
a small posterior one which fades into the right antapical suture 
distally. The longitudinal furrow is narrow and short, 0.6 dis- 
tance to postmargin. 

The thecal wall is structureless save for scattered pores in 
the two individuals thus far observed. These may both be young- 
stages and the older ones may be reticulate as are other species 
in the genus, but there is not the slightest evidence of reticula- 
tions on the thecal walls of these two individuals. The suture 
lines are light and faint. The midventral one on the epitheca 
is broad in the posterior half between the ventral pit and the 
flagellar pore, and the lateral sutures of the epitheca and hy- 
potheca are doubled and prominent. The right dorsal premedian 
suture is very oblique. No fins or lists were noted. 

The plasma is coarsely granular, chromatophores few, large, 
spheroidal, clustered near the center. 

356 University of California Publications. [Zoology 

Dimensions: — length, 116-120/x; transdiameter. 100/*; dorso- 
ventral, 75/x; furrow, 8/* in width. 

Taken in vertical hauls from 40-95 fathoms to surface off San 
Diego in May and June. 

This species is not closely related to any described species. Its 
asymmetry is noticeable but no other adaptations to flotation in 
the theca were found in the individuals examined. 

Heterodinium rigdenae sp. nov. 

PI. 18, figs. 6-8. 

A small pentagonal Peridinium-like species with a coarse 
polygonal reticulum and slighl obliquity <»t' the girdle, resem- 
bling /'. acutangulum Lemm. 

Body pentagonal in face view with straight or nearly straight 
subequal sides, broadly bifurcated posteriorly with short stout 
conical antapicals. Length 1.3 and dorso-ventral diameter 0.6 
times the transdiameter. Epitheca without apical horn, the 
sides Bloping in n straight line from the apical pore to the girdle, 
compressed dorso-vent rally and somewhat excavated in the mid- 
ventral region, lateral sutures prominenl and doubled. Hy- 
pe, t been also compressed, ibe right margin concave, the left with 
projecting angle at the junction of postmedian and antapical 
plates. Ventral face excavated. The antapical horns are short, 
0.2 transdiameter in length, which is nearly equal to the slightly 
curved margin which separates their bases. Their ends blunt 
with short terminal spinule. The girdle is inclined about 15° 
postero-ventrally from the equatorial plane. The transverse fur- 
row is deeply indented, more so towards its anterior than its 
posterior margin. It forms a descending right spiral with dis- 
placement slightly exceeding its width. Its distal end is feebly 
developed, the posterior ridge vanishing on the ventral face. 
The anterior ridge is a heavy projection of the body wall, nearly 
twice the height of the posterior. The longitudinal furrow is 
narrow and shallow, dilated posteriorly and extends from the 

Vol. 2] Kofoid. — On Heterodinium. 357 

flagellar pore little more than 0.5 of the distance to the post- 

Thecal plates of the normal type. Left intercalary confined 
to dorsal face. Suture lines heavy, deficient on right ventral 
area. Thecal wall covered with irregular polygons, mostly pen- 
tagonal, larger ones each with single central pore. Polygons 
relatively few, 41 on dorsal apical. In young individuals faint 
suture lines but no polygons are found. The ventral area is 
found as usual at the junction of the sutures on the ventral face 
of the epitheca. It is unusually large and contains the ante- 
riorly located reniform pit-like structure. 

Individuals thus far observed have been very hyaline and 
colorless or with pale greenish yellow chromatophores, reniform 
or irregular in shape. 

Dimensions :— length, 120-125/*; transdiameter, 90-92/*; 
dorso-ventral, 155/*; girdle width, 10/*; polygons, 2-10//. 

Taken in vertical hauls from 90-100 fathoms to surface in 
June off San Diego. 

This species resembles H. hindmarchi (Murr. et Whitt.) in 
the type of reticulations, but differs from it in its more robust 
form, shorter epitheca and antapicals. 

Heterodinium trirostre (Murr. et Whitt.). 

Peridinium trirostre Murray and Whitting ('99), p. 327, PI. 29, fig. 5. 

This species is shown to have the typical ventral plates of 
Heterodinium, the ventral pit and the deficient posterior list of 
the transverse furrow which is deflected posteriorly on the right 
antapical horn. 

The species is characterized by the presence of pointed an- 
tapical horns, the left showing a bifurcation into two apices. 
The epitheca is high, broadly cuneate with doubled lateral su- 
tures. The girdle is median, the transverse furrow being dis- 
placed only its own width. The reticulations are very coarse and 
heavy and exhibit more than the usual irregularity in form. 

Reported from 26° N. in the Atlantic. 

358 University of California Publications. [Zoology 

Heterodinium leiorhynchum (Murr. et Whitt.). 

Peridinium leiorhynchum Murray and Whitting ('99), pp. 326-327, PI. 29, 

figs. 2a, ft. 

This is unquestionably a Heterodinium as it shows a mid- 
ventral pit upon the epitheca. the doubled lateral sutures and 
diminishing posterior list of the transverse furrow. The plates 
are very imperfectly shown though there is a suggestion of the 
left intercalary of the epitheea in the slight shoulder on the left 
epitheca] margin, ami in the incomplete suture lines of the Lateral 

The species is characterized by the somewhat differentiated 
apical horn, the unequal, pointed, divergenl antapicals termi- 
nating in spines, the rotundity at the girdle, prominent suture 
ridges (incomplete in figure), smooth or coarsely reticulate thecal 
wall, and fenestrated lists of the girdle. 

Reported from the warm temperate Atlantic from L'<> 40 X. 

Heterodinium blackmani (Murr et. Whitt.). 

Peridinium Blackmani Murray ami Whitting ('99). pp. 327-328, PI. 29. 

figs. •'>". /'. '•. 

This superb and clearly marked species is the only one in 
which the left intercalary plate is shown by Murray and Whit- 
ting ('99). It is unquestionably a typical Heterodinium though 
these authors fail to show the ventral plates of the epitheca and 
the ventral pit characteristic of the genus. The remainder of 
the plates is almost completely shown. The prominent doubled 
Lateral sutures, the short Longitudinal furrow, and the poste- 
riorly deflected posterior list of the transverse furrow stamp this 
species as a typical Heterodinium. 

The species is characterized by the curved epitheca flaring 
to the greatly expanded equator, submedian girdle, divergent 
pointed antapicals. The suture lines are marked by prominent 
lists and the plates are reticulate with delicate subregular hex- 
agonal polygons, which exhibit a tendency to horizontal elonga- 
tion on the epitheca. 

Reported from the Caribbean Sea and tropical Atlantic from 
9°-25° N. 

Vol.2] Kofoid. — On Heterodinium. 359 

Heterodinium hindmarchi (Murr. et. Whitt.). 

Peridinium Eindmarcliii Murray and Whitting ('99), p. 326, PI. 29, 

figs, la, b. 

This is plainly a Heterodinium as it has the characteristic 
ventral pit in the central expansion of the median ventral suture 
of the epitheca, a short longitudinal furrow and the deficient 
distal posterior list of the transverse furrow. The plates are 
not shown but the presence of the left intercalary plate is sug- 
gested in the figure. 

The species is characterized by the long stout widely sepa- 
rated antapicals, elongated epitheca, and coarse reticulations. 

Reported from tropical Atlantic from Panama to 34° N. 

Heterodinium scrippsi sp. now 

PI. 17, figs. 1-5. 

A large species with short antapicals, scoop-shaped epitheca 
and coarse irregular reticulations. The body is subheptangular 
in face view, swollen at the girdle, and with shallow posterior 
bifurcation. The length is 1.5 and the dorso-ventral diameter 
0.7 times the transdiameter. The epitheca is long, and its alti- 
tude is about 0.8 transdiameters. The ventral face is flattened 
and hollowed out anteriorly and thins out laterally to the doubled 
suture lines on the angular margin between the apical plates. 
Posteriorly the epitheca flares out to meet the girdle. The lateral 
margins have rounded shoulders about two-thirds of the distance 
from the girdle to the apical pore which rises from the anterior 
end in a fully developed apical horn. A short ventral slot-like 
extension of the apical pore such as is found in Peridinium, fol- 
lows the midventral suture for a short distance. 

The hypotheca is shorter than the epitheca and in the mid- 
ventral line has scarcely 0.6 its altitude. To the tip of the long 
left antapical is 0.75 transdiameters. It is somewhat angular, 
is flattened ventrally and excavated in the midventral region 
between the antapicals. The dorsal side has considerably more 
flare toward the girdle. The antapicals are short and divergent, 
the right being more oblique than the left. The tips are acute. 

360 University of California Publications. [Zoology 

The postmargin is not set off from the inner margins of the an- 
tapicals with which it forms a fairly regular arc. The distance 
hoi ween the tips of the antapicals is 0.4 transdiameters. 

The girdle is postmedian, reniform in cross section, though 
somewhat thicker on the left side, and nearly perpendicular to 
the main axis. The transverse furrow is deeply indented with 
heavy overhanging anterior list and less strongly developed pos- 
terior one which vanishes distally on the ventral suture of the 
right antapical horn. The furrow is wide, postero-dexiotropic 
with a displacement equalling ils width. The Longitudinal fur- 
row is about 0.6 distance to postmargin in Length and is wide 
and shallow. 

The plates of the 1 heca are typical, the left intercalary being 
almost wholly confined to the dorsal face and of small size. Both 
pre- and postmedians are irregular in size and arrangement. 
There is an unusually Large ventral area in the midventral suture 
of the epitheca which is deflected 1<> the left and contains the 
reniform ventral pit. The suture lines are well developed and 
are laticed in places. Hyaline lists are round on the Lateral and 
postmargins of the hypotheca. The thecal wall including the 
girdle is coarsely and irregularly reticulate with well developed 
polygons of 3-5 sides, each with a single central jm.iv. Near 
the lateral margins of the epitheca these polygons are somewhal 
regular and often quadrangular. There are 108 in the dorsal 
apical plate. 

The plasma and indeed the whole organism is beautifully hya- 
line. There are a few suhspherical greenish chromatophores and 
an ellipsoidal nucleus near the flagellar foramen. 

Dimensions : — length, 140-155//.; transdiameter, 100-105//.; 
dorso-venlral diameter, 60/a; width of transverse furrow, 10-12/*; 
polygons 4-8//. rarely 12/t. 

Taken in vertical hauls from 95 fathoms off San Diego in 

I regard //. scrippsi as the type species of the genus. 

Vol. 2] Kofoid. — On Heterodinium. 361 

Heterodinium whittingae sp. no v. 

PI. 19, figs. 11-13. 

A large species with very oblique girdle, elliptical outline, 
and shallow rounded bifurcation. Body elliptical in face view 
with broadly rounded apical end and short incurved antapical 
horns which preserve the elliptical outline. The posterior bifur- 
cation extends but one-fourth of the distance to the girdle and is 
broadly rounded anteriorly. The body is very much compressed 
dorso-ventrally, forming a sharp edge at the lateral margins. 
The girdle is very oblique being inclined at an angle of 45° to 
the main axis in an antero-dorsal to postero-ventral direction. 

The length is 1.4 transdiameters and 3 times the distance 
between the greatest dorsal and ventral extensions which is found 
in the left half of the epitheca. The whole body is slightly 
twisted in a right spiral. 

The epitheca is very much flattened anteriorly and somewhat 
excavated on the ventral face, forming in fact a thin sheet which 
expands posteriorly as it meets the oblique girdle. The hypotheca 
is likewise flattened and excavated ventrally about the longitud- 
inal furrow. The lateral postmedian plates form a posteriorly 
projecting tooth on the left margin. 

The girdle forms a descending right spiral with slight dis- 
placement equalling its width. Its distal end is much widened, 
the posterior border becoming low and deflected posteriorly into 
the ventral suture of the right antapical horn. The longitudinal 
furrow is short and narrow. The flagellar pore is found as an 
elliptical opening at its proximal end. 

The thecal wall is thin, delicate and hyaline with light suture 
ridges except in the case of the lateral sutures between the apical s 
which are doubled and heavy, as are also the lateral sutures on 
the hypotheca. The plates are normal, the left intercalary being 
restricted to the dorsal face. The surface, including that of 
the girdle plate, is everywhere covered with a reticulum of irreg- 
ular polygons, each with a single central pore. In the several 
specimens thus far observed the reticulum has been very light 
and delicate. The polygons are relatively numerous, 97 having 
been recorded on the dorsal apical plate. 

362 I' ni r< rxity of California Publications. [Zoology 

The plasma is exceedingly hyaline and coarsely vacuolated 
and its total amount is relatively very small. The nucleus is 
minute (12/t), spheroidal and centrally located, and there is one 
pale chromosphere of similar size and form adjacent to it. This 
is a large species, 180/* in length, 140/a in transdiameter and 60^ 
in greatest dorso-ventral extension. Polygons 5-12^ in diameter. 

Taken in vertical catch from 85 fathoms to the surface off 
San Diego in Jul v. 

VoL - 2 1 Kofoid. — On Heterodinium. 363 


Cleve, P. T. 

:00. Notes on some Atlantic Plankton-organisms. Kongl. Svensk. 
Veten.-Akad. Handl. Bd. 34, No. 1, 34, 22 pages, 8 plates. 

Ehrenberg, C. G. 

'34. Dritter Beitrag zur Erkenntniss grosser Organization in der 
Kichtung des kleinsten Eaumes. Abh. Akad. Wiss. Berlin. 
1833, pp. 145-336, Taf. 1-11. 

Joergensen, E. 

'99. Protophyten und Protozoen im Plankton aus der norwegischen 
Westkiiste. Berg. Mus. Aarb. 1899. No. VI, pp. 1-112, 
I-LXXXIII. 5 plates. 

Midler, O. F. 

1786. Animalcula Infusoria fluviatilia et marina. Pp. LVI, 365, 50 Taf. 

Murray, G., and Whitting, F. G. 

'99. New Peridiniaeeae from the Atlantic. Trans. Linn. Soc, London. 
Botany (2), Vol. V, pp. 321-342, Tables I-IX, Pis. 27-33. 

Paulsen, O. 

:04. Plankton Investigations in the waters round Iceland in 1903. 
Medd. Comm. Havunderzog. Bd. I, No. 1. 41 pages, 11 fig- 
ures, 2 maps. 

Schutt, F. 

'96. Peridiniales. Engler und Prantl's Nat. Pflanzenfamilien. I 
Teil, Abth. b, pp. 1-30, 43 figs. 


PL ATK 17. 

Pig. l. Heterodinium scripp&i ventral view. • 725. <//». p., apical pore; 
/'/. p., flagellar pun'; /. /".. longitudinal furrow; y. a., ventral 

area ; o. />.. \ antral pit. 

Fig. 2. — Dorsal view of same. X 420. 

bHg. .'?. Oblique \i<'\\ of lefl aide of same. X 420. 

Fig. 4.- Diagrammatic apical \irw of same. > 420. 

Fig. 5.- Reticulations adjacenl to posterior list of transverse furrow. 
X 2725. 


UnivCalPubl.Zddl.Vcil II. 

Kdfdid] Plate XVII. 


Fig, ii. lh i> i-ikI i a i a in rigdenae, Neutral view. ■ 410. <'/'. / < . . .- 1 ( . i < - .- 1 1 pore; 
//. y., flagellar pore; I. /'.. longitudinal furrow; v. </.. ventral 
area ; u. p., ventral pit. 

Pig. 7. Dorsal view of the same. • 4 in. 

Fig. 8. — Vitw of lefl side of same. ■ 4in. 

Fig. 9. — Ventral \ i « - \\ //. inaequale. ■ 420. Abbreviations as in fig. fi. 

Fig. 10. — Oblique view of right side of same. 120. 

Uniy Cal.Publ Zddl.Vdl.1I. 

[ Kdfdid ] Plate XVII I 

PLATE 19. 

Fig. 11. Ventral view of Heterodinium whittingae. 420. ai>. p., apical 
pore; fl. p., flagellar pore; /. /'.. longitudinal furrow; v. a., 
\ entral area ; u. /-.. \ entral pit. 

Fig. 12. Dorsal \u-\\ of s:i mc ■ 420. 

Pig. 13. Vim of righl Edde of Bame. 120. 

Pig. II. Optica] section al girdle of same. ■ 420. 

Pig. !■"• Ventral view of //. sphaeroideum. 1500. 

[368 1 

Univ. Cal.Publ.ZddlVdl.II 

KnFDiD ] Plate XIX. 



Synonyms in italics, titles in boldface. 


A General Statement of the Ideas and the Present Aims and Status 

of the Marine Biological Association of San Diego i 

Abbreviations used in figures of Non-Incrusting Chilostomes 297 

Pelagic Tunica tes 108 

Abietinaria 21 

Acamarchis neritina 266 

Acartia 203 

tonsa 204 

Aetea 243 

anguina 241, 243, 244 

truncata 241, 243, 246 

Aetideus 144 

armatus 145 

Aeteidge 243 

Aetidiinse 143 

Aglaophenia diegensis 2, 5, 33 

inconspicua 2, 5, 34 

pluma 2, 5, 34 

struthionides 2, 5, 35 

Amallophora - ...:.... 163 

dubia 170 

A ,i<i a in aria 243 

spatulata 244 

A u I aria 215 

mediterranea * 216 

Area surveyed, The iii 

climatic conditions iv 

physical conditions of \ iii 

western boundary of vi 

Arietellus setosus 189 

Articulation 2-1 o 

Ascidiacea 97 

Augaptilus 187 

longicaudatus 188 

Avicella 265 

multispina - 266 

avicularia 270 

Avicularia flabellata 270 

Beania .--- 264, 276 

mirabilis 237, 238, 276 

longispinosa 277 

370 University of California Publications. [Zoology 


Behavior of Corymorpha 333 

Beroe forskali 47 

Beroida 47 

Beroidse 47 

Bibliography of 

( tcnophores 49 

Heterodinium 363 

Hydroids 40 

Xuii-IiK-rusting Chilostomes 293 

I 'elagic Copepoda 227 

Pelagic Tunieata 102 

Bict Hi i nn 27S 

Hi rt 1 1 ii riu 1 1, i 264 

Bict I In r ii iv 264 

Bicellariidaa 242, 264 

Bimeria 2, 3, 6 

gracilis 2, 3, 6 

Biology, marine xiv 

Biphora 53 

Bird 's Hi in! Corallim 270 

Bolina microptera 47 

septentrionaliB 47 



Bougainvillia 3, 7 

glorietta 2, 3, 7 

Bougainvilliidse 2, 3, 6 

BowerbanMa Lmbricata 239 

Bryozoa 237 

Bugula 264, 265 

avicularia 270 

ealifornica 265, 267 

curvirostrata 265. i, - 

flnhi Hutu 271 

flabellata 265, 270 

flustroidt s 270 

laxa 265, 275 

longirostrata 265, ^74 

murrayana 265, 266 

neritina 265, 266 

pacifica 265, 268 

pugeti 265, 271 

purpwrotincta 268 

Caberea 250, 263 

ellisi 263 

Cabereada? 249 

CcdaneHa 13 ° 

tin ditt mini n 133 

Index to Volume 2. 371 


Calanidae 122 

Calaninse 122 

Cala a aides 128 

Galanus 130, 135, 139, 142, 150 

lata 153 

mirabilis 133 

parvus 140 

Calanus 123 

comptus 129 

finmarchicus 125 

gracilis 128 

minor 126 

perspicax 125 

robustior 129 

tenuicornis 127 

valgus - 126 

Calycella syringa 2, 4, 20 

Calyptoblastea 2, 3, 10 

Campanularia edwardsi 2, 4, 11 

everta 2, 4, 12 

hesperia 2, 4, 12 

hincksi 2, 4, 13 

volubilis 2, 4, 13 

CampanulariidEe 2, 3, 11 

Cauda - 258, 265 

Candace 192 

bipinnata 195 

curta 196 

ethiopica 196 

melanopus 196 

pectinata 193 

Candacia ; 192 

aethiopica - 196 

bipinnata 195 

curta 196 

pectinata •- 193 

Candaciidae 192 

Carbasea - 291 

Catenaria 24; 

chelata - 248 

Catopia 1^1 

Cellaria 244, 247, 248, 249, 251, 258, 266 

anguina 244 

neritina 266 

ternata 251 

Cellaria borealis 287 

mandibulata - 287, 288 

diffusa 287, 289 

372 University of California Publications. [Zoology 


Cellarieas 286 

CellarudiE , 242, 286 

Cellarina 250 

gracilis — • 251 

Cellularia 244, 247, 248, 249, 250, 251, 258, 263, 265, 266 

a nil ni mi 244 

avicularia 270 

hookeri 263 

CellulariadcB 249 

Cellulariida 242, 249, 250 

< !entropages ' ■ ' 

bradyi 1 i - 

violaceus I 72 

» lentropagidae 170 

GentropaginsB '"' 

Ceratium 350 

Cetochilus 123 

h, Igolandicus 125 

longiremis 128 

minor j-26 

( Ihilostomata 243 

Clausocalaninse 14] 

Clausocalanua arcuicornia ... 142 

Clytemnesl ra 213 

rostrata 214 

tenuis -' I 

Clytia bakeri 2, 4, 16 

eompresae '-'. 4, 17 

bendersoni 2, 4, 18 

Coat of Mail Corallim 249 

Copepoda, collections made 113 

terminology of 114 

Corallina anguiformis 244 

( lorycseidse - 1 s 

( lorycseus 224 

\ enustus 225 

carina Ins 226 

( lorymorpha palma 2, 3, 9 

i lorymOrpha -, 3, 9 

behavior of 333 

geotropism 334 

locomotion 334 

prehensile meehanism 333 

reaction to mechanical stimuli 333 

regeneration 323 

Corynoporella 264, 284 

spinosa 284 

Index to Volume 2. 373 


Ctenophores of the San Diego Region 45 

Cyclopidse 20v 

Cyclosalpa 53 

bakeri 54 

Cydippida 45 

Dagysa 53 

Diachoris 276 

Diaptomus 171, 174 

abdominale 174 

Differentiation in Hydroid Colonies and the Problem of Senescence .... 323 

Dias 203 

Dinoflagellata of the San Diego Region. 

I. On Heterodinium, a new Genus of the Dinoflagellata 341 

Diplocheilus - - 2, 5, 35 

allmani 2, 5, 36 

Doliolidae 52, 82 

Doliolum 83 

life cycle 84 

nomenclature 84 

nurses 83, 84 

Doliolum denticwlatum 85, 91 

Doliolum Ehrenbergii 91 

gonozooid 91 

oozooid - 91 

phorozooid 94 

tropliozooid 94 

miilleri 95 

gonozooid - 95 

oozooid - 96 

phorozooid 97 

trophozooid 9' 

nordmanni 95 

Doliolum tritonis ■ = 85 

gonozooid 85 

oozooid 86 

phorozooid 88 

trophozooid 89 

Drephanopus 142 

Dubrieidlia 53 

Dynamena 2, 4, 30 

cornicina 2, 5, 23, 30 

Ectinosoma 210 

Ectoprocta 242 

Emma 250 

Endopodite - H4 

Epistome 240 

EscJiara 291 

874 University of California Publications. [Zoology 


hispida 266 

Esterly, C. 113 

EucalaninsB 130 

Eucalanus 130 

attenuatus 133 

erassua 134 

elongatus 131 

subtenuis I 35 

Eucalanus 142 

Euchaeta 156 

acuta 157 

media • 160 

sjiiiiosa 159 

Euchceta 150, I 17 

hessei L52 

pulehra 154 

KlH-liiitili;i' 156 

Eucharis grand] formis 47 

Euchirella 150 

amoena 155 < 

galeata 155 

iin-ssi tii-iisis 151 

pulehra 153 

rosl rata 152 

Euchirus 156 

Eucratea 247 

chelata 248 

Eucratiidaa 242, 247 

Eudendriidse 2, 3, S 

Eudendrium rameum -', 3, 8 

ramosum 2, 3, 8 


2, 9 

Euheterodinium 351 

Euplokamia calif ornensis 46 

Euterpe aeutifrons 212 

firm- His 212 

Plustra 291 

avicularis 270 

lichenoides 291 

membranaceo-truncata 292 

s< hid a 263 

Flustradoe 290 

Flustridse 242, 290 

Purea 114 

Gaidius 145 

pungens 146 

Index to Volume 2. ' 375 


Gemellaria 248 

avicularis 286 

loricata 247, 249 

loriculata 249 

Gemellariadce 285 

Gymnolaemata 242 

Generation, solitary of 

affinis 60 

bakeri 54 


confeederata-scutigera ; 81 

eylindrica 79 

democratica-mucronata _ 73 

tilesii-eostata 70 

Generation, aggregate of 

affinis 61 

bakeri 55 


confeederata-scutigera 81 

cylindrica 80 

democratica-mucronata 75 

fusiformis-runcinata 65 

tilesii-eostata 71 

zonaria-cordiformis 77 

Goniopsyllus -13 

Gonyaulax triacantha 341, 354 

Gymnoblastea 2, 3, 6 

Gymnoplea 113, 114, 117 

Haleciidse 2, 3, 10 

Halecium annulatum 2, 3, 10 

kofoidi ' 2, 4, 11 

washingtoni 2, 3, 11 

Harpacticus 210, 212 

Hemicalanus 171, 187 

H< 1 1 rochcetina 181 

Heterochceta 182 

clausii 185 

longicorvis - - — 186 

Heterodinium 341, 366 

blackmani 358 

doma 352 

hindmarcki 359 

inaequale - 354 

leiorhynchum : 358 

milneri 353 

376 University of California Publications. [Zoology 


murrayi 353 

rigdenae 356 

scrippsi 359 

sphaeroideum 351 

triacantha 354 

trirostre 357 

whittingae 361 

Bibliography of 363 

Definition of genus 341 

Detailed discussion of structure 342, 350 

Key to species of 350-351 

Description of species of 351-362 

Distribution of 349 

Flotation of 349 

Relationships of 350 

Eeterorhabdinae 181 

Eeterorhabdus 182 

spinifrons 183 

clausi 185 

longicornis 186 

papilliger I s 4 

:< ii sios - — ■ 186 

Eistory of Survey — 

A\:i](Hi xii 

donors riii riv 

fun. Is xiii 

laboratory x ii 

Sau Diego x '" 

San Pedro x '> 

Holothuria 53 

arium "6 

Bydraetinia calif ornica 2, 3, 9 

Eydractiniidae ... -> 3, 9 

Hydroids of the San Diego Region 1 

Tchthyophorba 171 

Ifionyx 192 

Jasis 53 

Keys to 

Eydroids San Diego Eegion 3 

families of Chilostomata 242 

genera of Bicellariidse .' 264 

( lellulariidse 250 

Gymnoplea 118 

Podoplea 205 

species of Aetea 243 

Bugula 2(35 

Calanus 124 

Index to Volume 2. 377 


Candacia 193 

Cellaria 287 

Eucalanus 131 

Euchseta 157 

Euchirella 151 

Eucratiidse 247 

Ileterodinium 350 

Heterorhabdus 182 

Menipea 251 

Scolecithrix 163 

Scrupocellaria 258 

Stirparia 278 

Kofoid, Charles Atwood 241 

Labidocera 199 

trispinosa 200 

Lafoea dumosa 2, 4, 20 

Lafoeidae 2, 3, 20 

La Jolla vi 

Leptocalanus filiformis 137 

Li nrLartiina 179 

Leuckartia 179 

List of species 

Heterodinium 349 

Hydroids 2 

Non-Incrusting Bryozoa ■ 241 

Pelagic Copepoda 116 

Pelagic Tunicata 52 

Lobata 46 

Lophothrix 163 

Lucieutiinee 179 

Lucicutia 179 

Marine Biological Association iii, xi, xiii 

Mecynocera clausi 137 

Menipea 240 

compacta 254 

erecta 251, 256 

gracilis 251, 253 

occidentalis 251, 254 

occidentalis catalinensis - 251, 255 

pribilofi 251, 257 

ternata 251 

Meteorological conditions of La Jolla, 

Table of iv, v, vi 

Metridia armata 177 

boeckii 178 

hibernica 177 

lucens 177 

:!i s University of California Publications. [Zoology 


Microsetella 210 

rosea 211 

Molluscoida 242 

Monoculus 123 

Neoscolecithrix 163 

Non-Incrusting Chilostomatous Bryozoa of the West Coast of North 

America 235 

\ < it, i m in 24S. 285 

ii r i riihir is 286 

I in if nl a I, i 24!» 

Notamiidse 242. 285 

Obelia corona 2, 4. 14 

dichotoma 2, 4. L5 

geniculata 2, t. L5 

Oithona oana 209 

plumifera 207 

sjiitiirnxl ris 207 

Oncseidaa 2 ID 

( )nc;i;i 215 

conifera 2 Hi 

minuta 217 

Ooecia 240 

Omithopora 21)5 

dilatata 266 

Ormthoporina _• < > r> 

il ii rill, ni, i 27<l 

Paracalaninse 138 

Paraealanus L39 

parvus L40 

Paraealanus hibi rnieus 177 

Parapontellina 202 

ParaponteUinse 202 

/'< g< n 53 

confoedi rata - - 81 

Pelagic Copepoda of the San Diego Region, The 113 

Pelagic Tunicata of the San Diego Region, excepting the Larvacea 5 1 

alternation of generation in 52 

Peridinium 341, 342, 344, 349 

I', ii,! in in in blackmani 341, 358 

dorna 341, 352 

hindmarchii 341, 359 

I I iorhynchum 341, 358 

milneri 341, 353 

tripos 341, 353 

trirostre 341, 357 

Pennariida? 2, 3, 9 

Phyllopus 190 

bidentatus 191 

Index to Volume 2. 379 


Physical conditions of Area viii 

Platydinium 351 

Pleurobrachia 45, 46 

bachei 46 

Plenrobraehiidas 45 

Pleuromma - 174 

Phunularia alicia 2, 5, 37 

megalocephala 2, 5, 38 

plumularioides 2, 5, 38 

setacea 2, 5, 39 

Plumulariidae 2, 3, 33 

Pleuromamma abdominalis 174 

ab dominate 174 

gracilis 17b' 

xiphias 176 

Podoplea 113, 205 

Polyzoa - 237 

Pontella 199 

Pontellidse 198 

Pontellinae 199 

Pontellinae - 198 

Poroceratium 350 

Protoceratium - 350 

Pterolyra *— - 53 

Pyrosoma 53, 98 

giganteum 98 

Pyrosomidse '. 53, 97 

BMncalanus 135, 189 

Eitter, Wm. E i, 51 

Ranatra 339 

Robertson, Alice 235 

Salicomaria 287 

Salicornariadce 286 

Salpa 53 

Salpa 53 

affinis 59 

Cabotti 73 

confcederata 80 

confoederata-scutigera 80 

cordiformis '. 76 

cordiformis-polycratica 76 

co rdiformis-zonaria 76 

costata 70 

costata-tilesii 70 

democratica 73 

democratica-mucronata 73 

ferruginea 81 

380 University of California Publications. [Zoology 


/' US i I'n rill is t!4 

fusiformis-runcinata <i4 

form cchinata 67 

in urn i ua I a "73 

octofora 80, 81 

polycratica 76 

run rin at a 64 

runcinata-fusiformis 64 

scutigi id 80, 81 

scutig era-con foederata 81 

tih si i 70 

tilesii-costata 70 

■.mill riii 7t! 

zonaria-cordiformis 76 

Salpida 53 

Salpigia Hassallia 246 
San Diego Region 

Ctenophoree of 15 

Dinoflagellata of ■"• 1 1 

Hydroids of 1 

Nun I acrusting Bryozoa of 235 

Pelagic < lopepoda of 1 1 3 

Pelagic Tunicata of 31 

Sapphirina 218 

angusta 221 

cliinsii 22] 

ilium i_"_' 1 

gemma 219 

iris 219 

Lornse 223 

salp<B 219 

Bcarlata 222 

Scolecithriciiue - 162 

ScoleeithrLx 163 

bradyi 165 

da n;i' 1'il 

persecans 166 

siniilis 1~<) 

subdentata 167 

Scribi Ua sc rib<l - 11 ' 

Scrviparia 247 

ehelata 248 

Scrupocellaria -250, 258 

bn risi. tis 259 

calif ornica 258, 259 

diegensis 258, 261 

scdbra 256 

varians 258, 260 

Index to Volume 2. 381 


Sertularella group 2, 4, 26 

halecina , 2, 4, 26 

pedrensis 2, 4, 27 

tenella 2, 4, 28 

tricuspidata 2, 5, 28 

turgida 2, 5, 29 

Sertularia anguina 32, 33 

desmoidis 2, 5, 30 

filicula 2, 5, 32 

furcata 2, 5, 3] 


anguina 244 

chelata 248 

loricata 249 

neritina 266 

Sertulariidae 2, 3, 21 

Snail bearing Coralline 266 

Sphaerodinium 350 

Staff, salaried members of . x, xii 

Status, present xi 

incorporation xi 

University's part xi 

assets xi 

officers : xi 

staff xii 

Stirparia 264, 278 

calif ornica : 278, 281 

ciliata 27S, 279 

occidentals 278, 280 

Synnotum 285 

aviculare 286 

Survey, marine biological ii 

aims of iii 

area iii 

history of xii 

Synthecium - 2, 22, 26 

Teniorinse 173 

Tethys .:. 53 

Thalia 53 

Thaliacea 53 

Thuiaria 2, 4, 21, 32 

Thecocladium 24, 25, 26 

Torrey, Harry Beal : 45, 323, 333 

Tubularia erocea 2, 3, 10 

Undeuclueta 147 

major 148 

minor 149 

382 University of California Publications. [Zoology 


Undma 123, 150, 163 

dance 164 

messinensis 151 

pulchra 153 

rostrata 152 

Jrochorda 53 


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