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UNIVERSITY OF CALIFORNIA PUBLICATIONS
ZOOLOGY
WILLIAM EMERSON RITTER
EDITOR
VOLUME II
WITH 19 PLATES
BERKELEY
THE UNIVERSITY PRESS
1905-1906
ee
Ao coma lngtis ~
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CONTENTS.
Introduction. A General Statement of the Ideas and the present
No. 1.
No. 3.
No. 4.
Aims and Status of the Marine Biological Association
of San Diego, by Wm. E. Ritter. With 2 maps ............
Mhe Jidex of a Marine Biological Survey -...---—---.._.._-
iibhempAtreais TOM Wee SUTVC ye Cn = ee mee ee eae eee a we were store em
SIM Tre) etdurarits\ eI MRSS G0 eee eee ene een me
Order of Advance on the numerous Lines of Investigation
Knowledge of the Physical Conditions of the Area —.......
Instrumentalities for Prosecuting such a Survey -..............- z
IMG CS SiiyarOls van ala TICs bath 25 cteeee cs acer ee
Present Status, asi to Ways and Means —...______.--...----
ns GORI C ale NO GCS See ee ete ke eso, cee ee ee eee ee ee
Remarks on the present Condition of marine Biology in
(Ger eral) eee oe oat mee eee ec gee Ses eee ee ce
The Hydroids of the San Diego Region, by Harry Beal
BIRO TEE Yeats ofl PLL CS pee eee ee eee eee
Hern O CUT CAO Ting see eee Sees eek Dee ee a ee Pees eat ee
(Maal ke: Gre ADM perel ove eno al SU ace ene et soe
Key to Families, Genera, and Species -............ eo eee tA
IDESERIp LOM ot NSP CClCSm eee = aene eee ee een See
Salty ey er eas nn rn se esa es ohn eS ee
The Ctenophores of the San Diego Region, by Harry Beal
TR rs Cy emi ea tl nent ese ne eee en cre ee
A estereilOme Ol POCLOS eee aes Mee eee reat cee cece
TEST GUI oeat 290) 0h 6 ce Ae ee ns ee oe Rae eee See RR eee ee
The Pelagic Tunicata of the San Diego Region, excepting
the Larvacea, by Wm. E. Ritter. With figures 9-31.
DEAE iS SS ee ee Pee ee
JUTGETROXO KUO TKO A eee io ee ec tn RRs See
TOP Cab SS] OLSON OTS) «eas cea ese se ne mera eae
IDWSOWISISMOT OIE (S]OC OME | aos acess occ Sra ace co eee SEE
PAST riVa een NeON VBS OVINGyegt2 4 a) eh) A eee cee eee neec cee eae ne fee ee Hee Ree eer
ID eXsrobiaty over a) ea Coes, 6 2b, 5C6 = Nee eee Pe eee nee ee PP Per
The Pelagic Copepoda of the San Diego Region, by C. O.
Bisbee eg Walhbl GOO UIE OS exes nes coe: oe tener enceat odes nace
VETa HATED LU VOLET OI ee ees Soe oe oe ce sore ee Beebe eee Eee Ee
erm Ol Giyg sees Ze et Dee Aas es See ae eee
TICES HCC BOTS | 0270) <M eaee eee ereneae eee e
Key, to Genera =... a gral ued Mees eee eS Seas eben at 118-121;
Description of Species -..... 5 Pee se coe ee eens. 122-205 ;
BACTATTO heatie climes OL Currey Nyse sass ee ce ne ee eee cece eee se
PAGE
XIV
538-101
102-107
108-112
113
113-115
116-117
No. 5.
No. 6.
No. 7.
No. 8.
Index
PAGE
Non-Inerusting Chilostomatous Bryozoa of the West Coast
of North America, by Alice Robertson. With 2 figures
and Plates, 4-16) x. -.sccxsscecen sci cce soe eee ce eee 235-322
Tintin tn GLO 25 saeco neck ee ee 230-237
Mer marnOlO gy cease oekcc lees Ro a Sa ee eee ee 237-241
Tiist o£ (Spectes j2::cc.csces lai -ccks eee 241
Description) jof Species! 5.2 eet cen eee eee bates nate nae 242-292
Bibliogranpbyy ¢ 22.2 b sooo cck LESS ee ose caer eee ee mre oe 293-296
Deseription: of Plates: a2 .2c.cc ceases eee one en ee 297-322
Differentiation in Hydroid Colonies and the Problem of
Senescence, by Harry Beal Torrey. With 4 figures -...... 323-332
Observations See cse ee eee ee eee ere ee RR oN 323-331
General, consid eratioms) ooo ease eee 331-332
The Behavior of Corymorpha, by Harry Beal Torrey. With
PAL OUP OSs) sees eee ee peat eh ae SESS A See Pe ad oe Ek ee 333-340
Dinoflagellata of the San Diego Region, I. On Hetero-
dinium, a new Genus of the Peridinide, by Charles
Atwood ‘Kotoid:* (Plates ilo oe eect ee 341-368
Introduction and Acknowledgments ..........-....------.-2-------200--00 341
Deseription of Genus Hleterodimium —-..2 222.222 ees 341-342
Detailed Description of generic Characters .....................-.....- 342-351
Synoptic Key to Species of Heterodinium .............................- 350
Discussion of Species -......... Sh i es ee 301
Biblio praplyy. » co ewcse ss ee acasn ance ce eae nae eee 363
Description of Plates -......... DR a Ee ORs ast A oc nd 364-368
pgs tS ee ee ee ee ae ete ea Ae, COL als
Phe tollawing Senos in'G <Ro an Archa logy, Egyptian
Ae
ean Archaeology and Ethnology. nd nthropolé cal Memoirs a
_Deprtinents of Anthropology: ese $2 :
_ EDA, -ARCHAROLOGY,
voll.
: , Hint, and J *Gilbart So Pages 690, moe tees
ae : eS Se ak
ne. Hearst Medical i in on
: = Sea, = Pytigns: Ais Sse ;
“No. ts takes and. ase bh the-Hupa, nee Pliny. Farle. Goddard.”
«Pages 88, Plates. 30, September, 1803. ~ er
“No. ie < — ee es vet tae Seite Paget 200, March, 3
we <4 2 a3 Cw ae —
_=No 3. _ “Types ‘of ae an. oa atallong b
8 = pe 22, June, 1904. ne ; :
UNIVERSITY OF CALIFORNIA PUBLICATIONS
ZOOLOGY
Vol. 2, Introduction, pp. i-xvii. April 5, 1905
A
CONTRIBUTIONS FROM THE LABORATORY
OF THE
MARINE BIOLOGICAL ASSOCIATION OF SAN DIEGO.
GENERAL STATEMENT OF THE IDEAS
AND THE PRESENT AIMS AND STATUS
OF THE MARINE BIOLOGICAL ASSOCI-
ATION OF SAN DIEGO.
BY
Wm. E. RITTER.
Director of the Station.
CONTENTS.
PAGE
Pthemideay ota. Marine brOloeiGal SUEVCY. «scess ses css + +s dces ce il
peel om ULC AasC OM CRN UTES Wi sirpelts iaf'sacre a ateratis << eiieie (enelies ree oi oe sare nisl ales ili
Some lye aalivnitvet lem SUC TMs eretortereace stn sys cia duels esis eiievaleiteceisi ea ‘avec, alate ebetsaays vii
4. Order of Advance on the numerous lines of Investigation........ vill
5. Knowledge of the Physical Conditions of the Area.............. vii
6. Instrumentalities for Prosecuting such a Survey................ 1x
fem NECeSsity, OLA SaMlarmled Stati fysc.6c ne cs iorss civics sc)4 <icis, v6 Foes cee = ae
SPETeSeninSbAbus: aS COM dys an Ge ViGAMGhis ci. 01 e<elleiiniets «ce -16 « siete « xi
QPBESTOTUCAIEINOUE! icy. «<<< ares o tue let ache afodeney ey haievel ote al sveeeioie weyarele: aah oee% xl
10. Remarks on the present condition of Marine Biology in General. xiv
ii University of California Publications. [ ZooLocy
1.—The Idea of a Marine Biological Survey.
Volume II of the University of California Publiations,
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, intensive 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 marine biology, and general biology
prosecuted by researches on marine 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 great variety of secondary
questions; but the sum total of the phenomena of marine plants
and animals will never be lost sight 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 it
gives definiteness and codrdination to action, and furnishes a
commanding point of view and stimulus. It is justifiable to hold
VOL. 2] Ritter—Introduction. 11
and be guided by it, even though assurance of opportunity to
earry 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 Pacifie adjacent to the Coast of Southern California.’’
2.—The 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 leneth 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. Asa 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 oceanie 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-
cations of the sort contemplated is continuousness 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
iv University of California Publications. [ ZOOLOGY
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 be done in a vessel much smaller, and hence
much less 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 at San Diego during
1904, an entirely typical year.
TABLE OF METEOROLOGICAL CONDITIONS, 1904.
TEMPERATURE. PRECIPITATION. WIND.
3 s Be
ae Wane ey
2 LOCALITY. a a as — i AE > a
= a a a As HA ad
JANUARY.
Nantucket. 2 .se% se. 34 22 5.98 16 11,849 60
Key West: 4).2% << one. 73 82 1.42 7 7,834 31
Parallone.. <incce-. see 54 49 . 88 7 12,117 48
San Francisco ...... 56 45 1.05 5 4,292 26
San iDiego" si... eents 65 47 .04 2 4,310 27
FEBRUARY.
Nantucketa moun 32 19 3.86 15 11,386 47
ey: Wiest: -.:t52.s'ges 76 66 1.08 +4 7,106 33
Harallones .cscece see 53 49 6.13 16 10,149 50
San Francisco ...... 55 46 5.89 16 5,561 31
San Dieso\ ct. esters 61 48 1 50 6 3,802 36
AO OE Rai pn iy me a lla alga
te eel
oe
VoL. 2]
LOCALITY.
MOonrTuH.
MARCH.
Nantucket ...
Key West
Farallone
APRIL.
Nantucket ...
Key West
farallone
May.
Nantucket ...
Key West
Farallone
San Francisco
San Diego
JUNE.
Nantucket ...
Key West
Farallone
San Francisco
San Diego
JULY.
Nantucket ...
Key West
Farallone
San Francisco
San Diego
AUGUST.
Nantucket
Key West
Farallone
San Francisco
San Diego
SEPTEMBER.
Nantucket ...
Key West
Farallone
San Francisco
San Diego
Ritter —Introduction.
TEMPERATURE. PRECIPITATION.
5
4 SI a
a a = n S
3 S iS >A
Ss a a As
Pereira 39 29 2. 1d 10
Bre istets 79 70 1.94 4
Be eas 60 49 6.30 24
aie siete Sif AT 6.01 2a
Sieaetanchs 63 50 PA ANTS 10
Sd Siti 48 38 4.08 iif
Saat anton 80 (Al iL yl of
sta epets 57 50 2.29 13
Digroneraee 63 50 1.29 8
Stee 66 52 15 S
EON anio 61 50 2.39 1
See ae 82 ike 13.01 12
a cecke 55 51 ss il
Be 3 eres 66 52 30 1
eet 65 56 12 3
Naowres 65 54 2.38 12
Nor stewen 86 ath 1 740) 112
satan aue 55 51 OL il
Soe 66 52 Trace 0
Siteeeanrc 69 60 0 0
Sie ects 74 62 2.09 9
aiayaeateats 87 77 1.40 i131
omer th tia 56 52 0 0
Bronce 62 52 .02 1
eoctere F/il 62 0 0
Meesaneys 71 61 2). 29 12
Cea aie ae 88 (tat 4.24 13
ae ath 56 53 Trace 0
sane ate 62 52 .06 2
erste ts 76 66 Trace 0
eee 67 56 .78 5
Se. eitSea 87 77 3} 515) 16
atthe state No records.
BP coneats al S17 5 OT 5
76 64 Trace 0
WIND.
ment in miles.
Total move-
11,294
7,242
11,940
7,126
5,041
10,274
7,378
10,890
6,544
4,665
9,033
6,018
14,993
8,921
4,153
9,019
6,856
13,757
9,448
4,531
8,011
6,750
11,974
10,574
4,335
8.377
6,417
11,066
9,674
4,165
8,869
6,092
7,141
4,132
Max. velocity,
miles per hour
ow w
Wo - Oo
V
vi University of California Publications. | ZooLocy
4
7
3
TEMPERATURE. PRECIPITATION.
E je ee
4 = = 2's Ss
o = Shs oF sey)
= LOCALITY. = a as E : Ea e A
= = = a AS oa aq
OCTOBER.
Nantucket) eS a-ecrne. D7 47 1.01 8 11,700 48
ey. SWiesb. eves ote 82 74 fi 14 8,675 71
Hamalloner ence see ese 61 56 2.01 7 10,791 50
San Francisco ...... 68 56 Deo 7 5,506 32
San Dieco: .f see ee: 74 59 Say 3 4,171 20
NOVEMBER.
Nantucket. “tf. es. 46 35 3.29 8 11,394 60
Fieve WeSt) 2.2r¥. cancer 76 68 6.22 12 7,573 43
Harallones .e. . ssen 2 59 51 1.58 6 9,168 43
San Aran CIsGO sae. 63 es LOT 3) 3,851 22
Sans Diegourn a... . es 74 54 0 0 3,930 23
DECEMBER.
Nantucket) Js%.4.. 35 25 4.67 17 13,184 64
Ney SWiOSUi:.. ern me's 74 65 . 34 2 6,841 26
Barsalloner eee. tetercis Sil 51 2.22 10 11,431 58
San Francisco ...... 55 46 1.59 10 4,876 38
san) DIe€Z0) «25 ca). 2 66 51 2.46 tC 3,884 19
Perhaps the most important fact, from the present point of
view, exhibited by this table is that pertaining to winds. It will
be noted, for San Diego, that the maximum velocity for the year
was 36 miles an hour, in February. On the basis of the “*‘ Beaufort
9?
Seale’’ 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
‘*Centle 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
oe
RGUELLO
Pt.CONCEPTION
Santa Barbara
O Los Angeles
PrVINC ENTE de ee Hill
PrFERMIN
Santa Barbara Is.”
Begqs Rock © Santa Catalina Is.
San Nicolas as
a9s San Clemente Is.
PTLAJOLLA
cheeT 4 >?
los. 3g, 569 807
Tanner Bank Bae Gi hoa
4/1
660 324 AIG
69 Cortes Bank 2 743 344
7 : 15@. jo a o2
Bishop Rk;.. 5!
Los Cornonados
1053
Map 1.—Showing the Area to be surveyed. Modified from United States Coast and Geodetie Survey Chart.
’
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Crit 4 Garr it ah OP Pyadodnlnu Ase i ale
Vou. 2] Ritter—Introduction. vu
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 vo!-
ume contains the following contributions to a knowledge of the
fauna:
No.1.—The Hydroids of the San Diego Region, by Professor
iE Bs lorrey.
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. O. 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.
vill 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 eases 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 bibhographical 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 order of 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 got well in hand,
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 facts
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, 1S 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-
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Vou. 2] Ritter—Introduction. 1X
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, ete., 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.
Physies, 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 California Publications. | ZOOLOGY
7.—Necessity of a Salaried Staff.
Obviously, there must be codrdinated 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 ealled 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 at the Station, that he might
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 chemist could easily be
found who would be glad to examine water samples that might
be sent to his own laboratory; and geologists there would be who
under like conditions, from their geological interest, would will-
ingly report 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, largely 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
works.
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-
ee a wee
a ld
VoL. 2] Ritter.—Introduction. Xl
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 codperation 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, 1904.
The present officers of the Association are:
H. H. 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. Seripps and Miss Ellen Seripps, 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. E. 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 some capacity with
the University, have been members for longer or shorter times
on assignment to particular pieces of work, and for the most
part 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. O. Esterly, A.B.,
Assistant in Zoology; John F. Bovard, B.S., Assistant in Zool-
ogy; Margaret Henderson, B.S.; H. M. Evans; L. H. Miller,
M.S., Assistant in Zoology; Robert Williams, B.S.; and Effie J.
Rigden.
9.—Historical Note.
Our work in this area did not begin with the San Diego
Association, or even with San Diego as a base of operations.
During six weeks of the summer of 1893 a party of teachers and
students from the Department of Zoology of the University of
Califoynia, 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 at
San Pedro, put in several weeks 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
.
Vou. 2] Ritter.—Introduction. xili
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 possible. 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 oecu-
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 lke-
wise spent at San Pedro, but this year nothing was done at sea,
attention being restricted to the httoral fauna.
During both these seasons formal courses of instruction in
Zoology were given as part of the regular University Summer
Session.
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
vears 1903 and 1904 the boat house at Coronado Beach, given and
in part fitted up by the Coronado Beach Company, served as ¢
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. R. Rowland, Mr. Van Nuys, The Los Angeles
Terminal Railroad, Mrs. Margaret Fette, Mr. J. H. Shankland,
Mr. John E. Plater, and Mr. Charles M. Wright.
By far the largest givers to the station since its removal to
San Diego have been Mr. E. W. Seripps, Miramar; Miss Ellen B.
Scripps, La Jolla, and Mr. H. H. Peters, San Diego. In addition,
the following have contributed substantially: Mr. Wm. Clayton,
for the Coronado Beach Company; Mrs. F. L. Keating, Mr.
Henry W. Putnam, Mr. G. W. Marston, and Hon. U. 8. Grant.
10.—Remarks on the Present Status of Marine Biology in General.
Situated as our station is, on a biologically almost unknown
part of a little known ocean, our first concern, chronologically,
must be with local conditions and problems. The meagerness of
knowledge, not only of the fauna and flora, but also of the ocean-
ography of the eastern part of the North Pacific can hardly be
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 urgent need of further exploration of the Pacific
from about 150° W. Lone. 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 the dinoflagellata, the radio-
laria, and the chaetognatha, about which there is hardly a
recorded observation. Even the better studied groups, like the
fishes, the mollusks, 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
— “cela
Vou. 2] Ritter—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 Baltie 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-
eance, biologically, of the continental shelf? What of Murray’s
‘‘mud line’’?
Xvi University of California Pubications. [ ZooLocy
When viewing this whole field of knowledge, and the means
and methods of investigation, one must be 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 need. This they have met, and are
meeting, magnificently. No other instrumentality has contrib-
uted so largely to the promotion of general 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 field, and to see more clearly what methods and instruments
must be used in their prosecution.
We are in position to appreciate, for 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 segment 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. 2] Ritter.—Introduction. xvii
I verily believe the value of the experimental and statistical
methods now so largely used 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 codrdi-
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,
ete.; and we are already in possession of important clues to the
way these questions must be studied; but we must learn, through
eareful 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.
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4
UNIVERSITY OF CALIFORNIA PUBLICATIONS
ZOOLOGY
Vol. 2, No. 1, pp. 1-43. December 21, 1904
CONTRIBUTIONS FROM THE LABORATORY
OF THE
MARINE BIOLOGICAL ASSOCIATION OF SAN DIEGO.
THE HYDROIDS OF THE SAN DIEGO
REGION.
BY
HARRY BEAL TORREY.
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 hes 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,t 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-
cies, (2) the permanent name if some change has been necessi-
tated, and (3) all synonyms in papers dealing with Pacific Coast
species.
1 With the single exception of S. pedrensis.
3 University of California Publications. [| ZOOLOGY
S& Ba
;|.9o Fs : ;
# (24) 5| El es £|%
TABLE OF DISTRIBUTION. 2 § = & Fy og : | 3
s| 2 (84/4 |<|B4| é z|8
w © |e » | eS] 96 ; ~
B\eied | Ba) eran) etace ied oe
a|/ulO |alalSala| <« |ale
Gymnoblastea
Bougainvilliidae
1. Bimeria gracilis Clark................]..... ren eee =e | hep Pescara (ee eRe | el Lc
2. Bougainvillia glorietta..... ........ ARB eg Pes LA ES He et i a2)
Eudendriidae | ¥
3. Eudendrium rameum (Pallas)... ...... +/+)... JGal\ sai) INE |e
4, ramosum (Linn.)..).0000) 00... yt] el IG Ye cece tee.
be TR eek pete RR | OIE he WAS lies a leet ate neh se Wiad rei tte pee
Hydractiniidae | ;
6. Hydractinia ealiforniea .. .......... ee ee bee at essa erence ee
Pennariidae Ps | ie ici a ka
7. Corymorpha palma T.............-.... ease AE + | + |..... lanl Fal cede al oe sh Ree
8. Tubularia crocea (Ag. )........ 0... |.s.|.0-- sgl Peg ist otra ted Cer Ree oe ee ears
Calyptoblastea Pa
Haleciidae
9. Halecium annulatum T...............|'..... eee eee ste | caacculeeesea eee Pacem toy We ?
10. kofeirdi Tt. (2.0 Pa ed SS ES eRe q
iil washingtoni Nutting ..... Nhe eee Spel eosees [see ectt | cere Oe heey mee 1
Campanulariidae | |
12. Campanularia edwardsi eA Pons eed See mba pa corks dao tos cone [Seen] eee ?
rh everta Clark .. ce ae! foceses| SE | HE [eee [ea Roe | RS a Nicec, 4 eee 1
14. HGSPOVI Be =< Serncsnysec| ese ficee [tee = |. a PE ae? eee ee ‘
15. hineksr Aldor | lie cle) ete ered oe Baa ae D
16. volubilis (Linn.)..| ....)-+|+]+/+ | 1 |+]| NE |.....[...... j
17. Obelin corona s...26- ics te b Peilles). Vee od Pa bores BA [5 ee [es all te ;
18. dichotoma (Linn.) ..... .... ap. | foe aroha sess - jhe eel ,
19. geniculata (Linn.).. ...... |......]...... +\/+/+) 461+] NE L.. -f- }
B02 Oletia aK Riya meet ee | ve Seats ties [ested eso iesostte lee lc cone | oe eae ;
21, compressa (Clark) ......... Sad, eT hcl Pcp actl Borel Peer | = ee Senet eee) (ee, ,
99) henderson) 22. ea eee RA as cs od eee) Eee Ue Fle ay [ee | ’
23. WNT VEGPSLCRUUIS sco 5 aes tel| eestor Wah cee ect Poni reeset | rate sal eee ‘
24. Calycella syringa (Linn.)......... |) | + |+}+|I1G)|+/N-+B)..... |...... ?
Lafoeidae | )
25. Lafoea dumosa (Flem.) ........... j}+/+/+/+/4+]SG/4+] NE ......)...... }
Sertulariidae | i
Sertularella group :
96. Sertularella halecina T...............)..0...| 2... |... ee py eee! cae sed he cen ie Maa ’
97. Pedrensis ese see + yo. [ieee We cera egal a ;
28. tenella (Alder) |... |..... +]... os Pee, a Petes ed eta vcd [le + :
29, tricuspidata(Alder), + | + |... eee a bee se a Gd a i
30. turgida (Trask) .... |...... +/4+)+) 0... nate ery |e Ae doe en ell ee
Dynamena group 4
31. Dynamena cornicina McC, ....... Pees ey nea eg We hd Hk OR 7
32. Sertularia desmoidis T.............. ised Boel he| S1F econ onset een) cots eee se eee
33. furcata Trask .......... | otto ad ese aes Ae eee a ioe
Thuiaria group
34. Sertularia filicula E. & 8... ...... +/+/+/+/+} G@ i+] NE h....J....
Plumulariidae
35. Aglaophenia diegensis T. .. eee eae SN eee eel peda Doe eke oe eee
36. inconspicua T. Be Reel Pee + a... 3 eta oe eM Mee ne | bee
37. pluma (Linn. a eee ig. A | ea Se Wee fee ae Sl eee 8 + Yo...
38. struthionides (M. ) seta Eee pe eater cat eg El)
39. Diplocheilus allmani ..................| .... eed cecil STEAL ene a ee ese
40. Plumularia alicia T. as el fac go) el ems ee ean [eg
41. megalocephala Aeelecaliog: sf auld |: Socata oS. elt Se Zaye
42, plumularioides(C.) ons 5 =e A al eee lice ge A oe eo dle
43 setacea (Ellis) seen la +)4+/+)4 |... cn (eee ame a tate, le Pav
Totals .......... Kosts Wed ants stem | 7 {11/20/42/14] 8 |13/ 8 |1]e
Vou. 2] Torrey.—Hydroids of the San Diego Region.
KEY TO FAMILIES, GENERA AND SPECIES.
. No true hydrothecae or gonangia .......... .......... ........ Gymnoblastea
1. True hydrotheeae and gonangia present.........._..... Calyptoblastea
2. Tentacles in proximal and distal sets; hydranth abruptly set off
OTHE SUC Litem meres Ar cee tv eel Ei ee a Pennariidae
Pealentaclessimuonercirclet, filufomm eee sc ue es
3. Solitary nutritive polyp, rooted in sand; perisare rudimentary .........
sogicna Cucest eeepc ee Be ST ne ES eee aa Corymorpha
3. Several anaes polyps from common stolon; perisare well
developedssconophores! feds ee eee Tubularia
4. Not more than 30 proximal tentacles; doneohores without tentacles
Ee REE re PRA 5 CoE INE ly RAR SO ee A Deen, C. palma (p. 9)
5. About 25 proximal tentacles; gonophores with 6-10 flattened pro-
OSS OS UME Maree Pest Stas rene SOS NRE Re See TN ene T. crocea (p. 10)
6. Colony encrusting; hydranths clavate; spiral zooids; spines on
hydrorhiza _..... get eee ae Be EN Nea Brae Hydractiniidae, Hydractinia
Gr © ol Omiyanon Chia aerate. rat 6 te ae ee Sp AP Oe DLAONE LE Sent os
7. Sterile hydranths with 6-10 tentacles; blastystyles sai knoblike
Gembaclesy ete ete ea pte ea eee NE ote le ok H. californica (p. 19)
8. Proboscis trumpet sisal. lonychiaivansm 1aVONe THOUSNUEO WAC es ene ee
Weal rn erent Need Ss. ba AS i wy Ae Eudendriidae, Eudendrium
8. Proboscis conical; hydranth fusiform .............. .......... Bougainvilliidae
9, Stem and principle branches polysiphonie............... E. rameum (p. 8)
9. Stem polysiphonie at base only; branches simple...
shy jal otek eng eee R, LE ee ee OE LN ER ne ad . E. ramosum (p. 8)
ORR ONOPHOTES xe ClSPOROSACS =e i ane eee eee eee Bimeria
10. Gonophores liberated as medusae, with four pairs of tentacles...
ee fe mg te ero ie NL Gi Nur eB ine ton LU ee as EN Bougainvillia
11. Stems simple, only partially annulated; hydranth with 10-12
LOMUACIE Sentence ence ey tne Marra eelomte Lie rr eke B. gracilis (p. 6)
12. Stem and branches without annulae, often twined about each other..
serie ot Ta aR oo oer eae EN aE IE EE B. glorietta (p. 7)
Sp eNemat OplOTesy Sos emit eye ey ets ee ee ee ee en eee
13. Nematophores present; hydrothecae sessile, on one side of stem or
Joram cle szomiliya emer st ted eo wen ays pear ieeeeee ee Plumulariidae
14. Hydrothecae with definite basal septa; gonangia not aggregated
14. Hydrothecae without definite basal septa, tubular; smooth margined;
Cone Cdayame rer ate den hers eee eee ele A yy — latoeidae
15. Hydrothecae much wider than deep ......... .. Use ESS ety see Haleciidae
15. Hydrothecae campanulate, usually deeper than wide, never adnate
TO POU GEN CURA ce sae casey tine eo a Egle Campanulariidae
15. Hydrotheeae sessile, adnate or immersed, in 2 to 6 rows. Sertulariidae
16, Gonangia with sporosaes ..............--.--....- ESO ee Halecium
(PMOLOUI NOt ASCIC]OG te eenem ree = eth nie ety ee oe Re meee e A Deere
17. Stem fascicled; ultimate branches gontealnres H. washingtoni Ge. 11)
. Annulated; hydrathecae about half as deep as broad, margin everted
Re es de ke CM h ee ee ein eee et eh LS eee H. annulatum (p. 10)
~]
10
University of California Publications. [ ZOOLOGY
. Not annulated; hydrotheeae shallow, margin not everted...
es das cease haat ee Sear kw ran ee eee H; kofoidi (ex 1)
19. Opereulum of numerous small triangular pieces; unbranched;
hydrotheeae tubular .......... een a Calycella, C. syringa (p. 20)
MSO) ING ovyo xe @ ua a nae ay enc a wae ceo Fee eee
20, Gonophores SGSsiles SPOTOSRCB co aeere eee ee Caapenuiena
202 SG ONO MORES ETE C: W012 GS ie hee eee ee nay eee cern ee eee
21. Stems branched, non-fascicled; nedattniees with 12-14 marginal
IHX 1 Motes a a Le RED Tale Pete Sa ag Na eo WALT aD io! C. edwardsi (p. 11)
Dy Stems wmbranched sf. co. eee ce eee eeereee
22. Hydrotheeal margin with 12-15 crenations; gonangium compressed,
paris AVORbMEO: 2c ees Poca es a C. everta (p. 12)
22. Hydrotheea with 10-12 square- siege teeth............ C. hincksi (p. 13)
99
. Hydrotheea deep, with 11-12 short, sharp teeth....C. hesperia (p. 12)
. Hydrotheea small, tubular, with 9 low blunt teeth
bas Ps PP rey ee ete cee _C. volubilis (p. 13)
23. Medusae ieorated with ab dleastulortentacles! 225. Obelia
23. Medusae liberated with 4 tentacles... 2.0.0.0... 1c cee eee Clyttia
24) Eiydrothecal marcin Su 0 th eects. cec ccs ca cneaas cone acereseree ALP hp
24. Hydrothecal margin bidentate........... .............. Rote corona ate: 14)
25. Branching irregular; stem non-geniculate....... a. dichotoma (p. 15)
25. Usually unbranched, except for pedicels; latter on shoulder pro-
}. Stems simple .........
. Branched; hydrotheea delicate, with about 14 carinate teeth
. Hydrothecae in two rows ......
2. Hydrothee ss margin dentate :
. Hydrothecal margin entire, everted, hydrotheca tubular; branches
. Three marginal teeth -.................- ... :
3. Four marginal teeth; hydrothecae and. Hone paauienee
cesses of geniculate stem... ......................O. geniculata (p. 15)
. Stems and branches polysiphonic; colonies large and _ bushy,
hydrothecal margin with 12-15 teeth ........ C. universitatis (p. 19)
.C. hendersoni (p. 18)
. Unbranched; hydrotheas Te with thick wall, crenate margin;
gonangium compressed, wide aperture .. ...... C. compressa (p. 17)
. No branches except hydranth pedicels; latter usually with but
single annulus; hydrothecal margin smooth C. bakeri (p. 16)
, “Eiydmothecne Biratg hit. 2c. cc.c is cxes tosh eres eke eerie ene Lafoea
29. Hydrothecae sessile; stem creeping or erect and polysiphonic
SE c Nee als nGImosa: (pees)
. Hydrotheecae alternate, one to an internode .. . Seneniavetia group
. Hydrotheeae opposite, one pair to an rien ane re Dynamena group
. Hydrothecae subopposite to alternate, more than two to an internode
A Sch ee OE AR tice hG re Se Ete tens oe D DUWIATIAl group
and gonangia rising within hydrothecae.........S. halecina (p. 26)
eM Pete ach Le. Ee A TL er ee ee fee Satenellang (p-e28)
. Hydrothecae corrugated; internodes long, slender; gonangia
SpimOdOse 4.5 8S, face an te ces wee PEO MEM SISH(D AS AM)
20
21
23
99
oo
29
34
- *
ee
Vou. 2] Torrey—Hydroids of the San Diego Region.
34.
3o4.
Hydrothecae smooth; internodes moderate; gonangia with high
narrow transverse corrugations ..................... S. tricuspidata (p. 28)
Hydrothecae smooth or roughened; internodes stout, geniculate;
gonangia with distal spines or annulae or both..S. turgida (p. 29)
. Margin of hydrotheca without teeth... .... S. desmoides (p. 30)
. Margin of hydrotheea with two teeth 0... pee ee a ee
. Teeth prominent; mouth of hydrotheea large... S. furcata (p. 31)
. Teeth moderate, mouth of hydrotheea small ...... D. cornicina (p. 30)
. Aperture of hydrotheca round; hydrotheca abruptly narrowed
GUISE Sa eeree ean ORR ck, A ey Mice a tees IEA Oy Se S. filicula (p. 32)
fy Nemalophoresmixedslyacl DLO dal ase eee ene ee nee em en
: Nematophores fixed by a marrow base 2... cc coc eecceseeee eee ceceees
. Stem simple; hydrotheca with dentate margin, no anterior intra-
theeal ridge; corbulae without hydrothecae at bases of leaflets...
Aglaophenia
. Stem simple; hydrotheca with anterior intrathecal ridge; single
sapracalyeine sarcostyle; no corbulae........ 2. oe... Diplocheilus
. Hydrotheea with 11 irregular teeth ............... A. struthionides (p. 35)
pEivdrothecanwitheONteetihe:£ 8. cn) eed Se eee eee ane en
41. Median tooth recurved................ .... Een ee rm, eee te oe ee
“lee Median tooth notmecurvedss ss ee A. pluma (p. 34)
42. Hydrocladial nodes well marked; mesial nematophore reaches mouth
Otmhiy Cobh Cae tees 5, tose OL. en Se de ee, A. inconspicua (p. 34)
42. Hydrocladial nodes weak; mesial nematophore does not reach mouth
ObghiyGro te Cae es ose yr ere. Soe da AA eager A. diegensis (p. 33)
43. Margin of hydrotheca smooth; mesial nematophore very short and
| ONROYEN Fk ot ras Ae ee dae OE POLE Rs Pre Ped D. allmani (p. 36)
44. Hydrocladia pinnately disposed on erect stem without thorny pro-
cesses or nematophorous branchlets; no corbulae ........ Plumularia
45. Hydrocladial internodes thecate and non-theeate ...... ne Rey ai eae nee
45. Hydrocladial internodes theeate only ..... .. P. plumularioides (p. 38)
NOG AILSE pluminamS Vers Cpe tet a ete eee tale Ae ae
. Nodal septa alternately transverse and oblique; thecate internodes
twice as long as non-theeate —...0..... oe eeeeee cece Pealician(on 3)
. Hydrocladia borne on short processes of stem, always alternate .......
P. setacea (p. 39)
. Hydrocladia borne on long processes of stem, very slender, opposite
Cis taillivgere teeta te hee Bs ...P. megalocephala (p. 37)
36
40
43
41
42
45
46
47
6 University of California Publications. | ZooLocy
GYMNOBLASTEA.
Fam. BOuGAINVILLIDAE.
Gen. Bimeria, Wright, 1859.
1. Bimeria gracilis Clark.
Bimeria gracilis, Clark, 1876a, p. 252, pl. 38, fig. 3.
Thophosome. Stems rising from creeping hydrorhiza to height of
¥0 to 30 mm., with numerous short branches. Hydranths borne alternately
on latter, on moderate pedicels; 9-11 rather stout tentacles. Stem usually
smooth, occasionally wrinkled. Pedicels with 2 to 5 more or less indistinct
annulae at the base. Perisare opaque, extending to bases of tentacles.
Gonosome. Sporosacs ovate, borne on branches singly or in pairs,
Pedicel short and smooth; spadix branched.
Fig. 1.—Bimeria gracilis. Gonophores.
Distribution. Dredged near the mouth of San Diego Bay, in
3 fathoms, July, 1903; La Jolla, at low water, July, 1903. San
Diego (Clark).
There can be little doubt that this is Clark’s species. The
perisare was covered with minute adhering particles. Male
sporosaes are slightly smaller than female.
Vou. 2] Torrey.—Hydroids of the San Diego Region. 7
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 em. Stems, or stem and branches fre-
quently twine about each other. Perisare smooth, without annulae, ocea-
sionally wavy, unusually adhesive, covered with particles of dirt and diat-
oms, reaching bases of tentacles. Terminal hydranths largest, with 20
to 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.
\
\\
Ny
NY
/
Fig. 2.—Bougainvillia glorietta. Fig. 3.—B. glorietta.
Hydranth from below. Hydranth with tentacles
partly retracted; edge of
perisare 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 perisare
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 perisare, July 5, 1903.
8 University of California Publications. | ZooLocy
EUDENDRIIDAE.
Gen. Eudendrium, Ehrenberg, 1834.
3. Eudendrium rameum (Pallas).
Tubularia ramea, Pallas, 1766, p. 83.
Eudendrium rameum, Johnston, 1847, p. 45, pl. 5, figs. 1, 2.
Eudendrium rameum, Torrey, 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; perisare rigid, occasionally marked with nearly obsolete annu-
lations on the smaller branches. Hydranths with about twenty tentacles,
frequently atrophied in the male after the production of gonophores.’’
Gonosome. Male sporosaes 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, Cal., on float at surface
(Dee., 1901; no gonosome). Mediterranean, Norway, Gt. Britain
(Allman). Jan Mayen (Markt.) Helgoland (Hartlaub). Green-
land (Levinsen). Northern Asia (Thompson).
4. Eudendrium .ramosum (Linn.).
Tubularia ramosa, Linnaeus, 1767, p. 1302.
Eudendrium ramosum, Ehrenberg, 1834, 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; perisare 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; Pacifie
Grove, Cal. Gt. Britain (Allman). Jan Mayen, Adria, Rovigno
(Markt.). Helgoland (Hartlaub). Greenland (Bétencourt).
Vou. 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 (771), with a few shght 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
perisare is sparsely and wavily annulated. Female blastostyles
with tentacles.
Fam. HybDRACTINIIDAER.
Gen. Hydractinia, Van Beneden, 1844.
6. Hydractinia californica, n.sp.
=
Trophosome. Sterile hydranths 2 to 2.5 mm. long in extension, with
6 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 knob-like clusters of nematocysts representing tentacles; 1 to
1.3 mm. long.
Colors. Perisare deep brown, fleshy parts white.
Distribution. Off San Diego, in 50 fathoms. Covering the
shells of Dentaliwm polygonum inhabited by hermit erab.
This species is very closely allied with H. echinata (Flem.) of
Europe and H. 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 perisare proximally
10 University of California Publications. | ZooLoGy
for 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
em.; 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, and a manubrium at least
twice as long as the bell, without a mouth; tentacles wanting; velum may
be present or absent.
Distribution. San Diego and San Pedro, Cal., throughout the
year, between tides, on sand flats. Eggs laid in May, June, July.
Gen. Tubularia, Linnaeus, 1767.
8. Tubularia crocea (Ag.).
Parypha crocea, L. Agassiz, 1862, III, p. 249, pls. 23, 23a, figs. 1-7.
Parypha microcephala, A. Agassiz, 1865, p. 195.
Tubularia crocea (Ag.), Allman, 1871, p. 416.
Tubularia elegans, Clark, 1876a, p. 253, pl. 38, fig. 2.
Tubularia crocea, Torrey, 1902, p. 43, pl. 2, Figs. 22, 23.
Trophosome. Colony usually a bushy mass of stems, tangled below,
which may be 10 em. long and may occasionally branch. Hydranths with
not more than 25 proximal tentacles.
Gonosome. Gonophores borne in pendulous clusters on peduncles
arising between proximal and distal tentacles; with 6 to 10 flattened pro-
cesses, varying in size, more prominent in female, sometimes hardly visible
in male.
Distribution. San Diego Bay, San Pedro Harbor, and San
Francisco Bay, Cal. Eastern United States (Agassiz).
CALYPTOBLASTEA.
Fam. HALECIIDAE.
Gen. Halecium, Oken, 1816.
9. Halecium annulatum.
Halecium annulatum, Torrey, 1902, p. 49, pl. 3, figs. 30, 31.
Trophosome. Stems rising from a creeping hydrorhiza to a height
of 7 mm.; the longer have 2 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.
Vou. 2] Torrey —Hydroids of the San Diego Region. ital
Distribution. Coronado Is., Mexico (July, 1903), and Coro-
nado, Cal. (July, 1901). Growing on seaweed.
10. Halecium kofoidi.
Halecium kofoidi, Torrey, 1902, p. 49, pl. 3, figs. 32, 33.
Trophosome. Stems rising from creeping hydrorhiza, 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. Hach internode usually bears on a distal
shoulder process a sessile hydcotheca which does not reach beyond the
distal 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;
Catalina Is., 42 fathoms.
11. Halecium washingtoni Nutting.
Halecum geniculatum, Nutting, 1899, p. 744, pl. 63, fig. 1.
Halecium washingtom, Nutting, 1901, p. 789.
HTalecium nuttingi, 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, 799).
Distribution. Pt. Loma, July, 1901. Puget Sound (Nut-
ting).
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 campanulate, 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 3 fathoms. Woods
Hole, Mass., between tides (Nutting).
There is much variation in the size and proportions of the
hydrotheeae, correlated apparently with mode of growth. The
stems are usually densely clustered. Some of the larger stems,
however, grow more freely above the rest. These longer stems
bear the longest hydrothecae, which may be .92 mm. long. On
the crowded stems, the hydrothecae may be no longer than .60
mm. Short seattered stems produce the largest hydrothecae of
all. The diameter does not vary with the length, as the following
Measurements in mm. show: .625x.26; .67x.31; .68x.45;
10x .45; .72x39; .92x.45. As a rule, however, the diameter
is less than half the length. The theeae growing in the clusters
are relatively broader.
The gonosome was not present, July 15, 1903.
13. Campanularia everta Clark.
Campanularia everta, Clark, 1876a, p. 253, pl. 39, fig. 4.
Campanularia everta, Torrey, 1902, p. 51, pl. 4, figs. 35-37.
Trophosome. Pedicels of variable length, smooth, wavy or irregu-
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. Acrocysts may be present in female, which are some-
what 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. C. everta can be distinguished from
Clytia 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, unbranched, 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-
——— a
Vow. 2] Torrey.—Hydroids of the San Diego Region. 13
theeae less than half as broad as long (.5x.22 mm.; .6x.25 mm.), with
11 or 12 short, sharp marginal teeth. Hydranth with 22 to 24 tentacles.
Gonosome absent, July 13, 1903.
ee a
Fig. 4.—Campanularia hesperia. Hydrotheca and pedicel.
Distribution. la Jolla, Cal., between tides, on the tests of
ascidians.
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 Hinks, Alder, 1857, p. 127.
Campanularia Hinksii, Hincks, 1868, p. 162, pl. 24, fig. 3.
Campanularia hinksi, 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, pl. 13, fig. 7.
Campanularia volubilis, Hartlaub, 1901, p. 357.
Campanularia volubilis, Torrey, 1902, p. 54, pl. 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
14 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.
Trophosome. Colonies very low; stems simple, short, slightly flex-
uous, from a creeping stolon, with 3-6 annulae distal to each pedicel.
Hydranths long, narrow, tapering, margin with 8-10 teeth, each with two
sharp eusps, pedicel short, completely annulated, with 2-4 annulae.
Hydranth with about 24 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. Hydrotheea: .44x.20; .43x.18 mm.
yonangium: .76x.20 mm. (including pedicel).
ASA
Fig. 5.—Obelia corona. Fig. 6.—O. corona. Gonangia.
Hydrotheeae.
Distribution. San Diego Bay, on piles under 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.
Vou. 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, pl. 28, fig. 1.
Obelia dichotoma, Calkins, 1899, p. 356, pl. 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 undescribed 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 O. 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. ‘‘Gonothecae 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 (Hineks). White sea (Meresch-
kowsky). New Zealand (Hartlaub).
Gonosome present, July 1, 1903.
16 University of California Publications. [ ZooLocy
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, 1816.
20. Clytia bakeri, n. sp.
Trophosome. Stems clustered, about 20 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. Stem internodes usually 3 to 4 times as long as broad, each bearing
a pedicel on a shoulder process from distal end. Hydrotheecae small, con-
ical, without marginal teeth; pedicel consisting usually of but one annula
almost as long as broad.
Gonosome. Gonangia long, narrow, with bottlenose apertures, taper-
ing gradually to short peduncies; borne usually in pairs at the bases of
hydrothecal pedicels. Sporosacs abundant, 12 to 20.
: ,//
\ ke
| |
(I }
j
< oF Wa
Fig. 7.—Clytia bakeri. Fig. 8.—C. bakert. Fig. 9.—C. bakeri.
Proximal portion of Portionof stem indistal Proximal region of
stem, showing charac- half, with three typical stem with gonangia
teristic annulation. hydrothecae, one of and hydrotheca with
which is sessile. exceptionally long
pedicel.
Vor. 2] Torrey.—Hydroids of the San Diego Region. Az
Distribution. Pacific Beach, in the surf, attached in tufts to
the posterior region of both valves of the beach clam Donaz, 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 annulae
or internodes on the stem and the annulation of the pedicels.
Occasionally a pedicel has several annulae (fig. 9), but this is the
ease 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;
A8 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, 1876a, p. 214, pl. 8, figs. 5, 6.
Clytia compressa, Nutting, 1901, p. 170, pl. 17, figs. 3, 4.
Cytia compressa, Torrey, 1902, p. 58, pl. 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. Orca, Al. (Nutting). Shumagin Is., Al., 6-20 fathoms,
on Laminaria (Clark).
Gonosome present, May 28 and July 13, 1901.
18 University of California Publications. | ZooLoGy
22. Clytia hendersoni, n. sp.
Trophosome. Colonies branching, 3 to 5 em. 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.
~<a
=
|
Fig. 10.—Clytia hendersoni. Stem Fig. 11.—C. hendersoni. Stem
with hydrothecae. 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.
Vou. 2] Torrey—Hydroids of the San Diego Region. 19
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 denticulata, Torrey, 1902, p. 51, pl. 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-
war, occasionally 1 or 2 annulations distally. Medusae numerous, oldest
with 4 tentacles.
Dimensions. Hydrothecae, in mm.: .70x.37; .78x.40; .81x.39;
83x42. Gonangium: .95x39; 1.00x.41.
Fig. 12.—Clytia uni- Fig. 13.—C. univer-
versitatis. Hydrotheea. 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-
ticillata. The pedicils of the hydranths, however, are not arranged
in verticils, and the gonophores are not sporosaes. Young colo-
nies taken in San Pedro Harbor, December, 1901, were previ-
ously identified with ©. denticulata Clark, though certain differ-
ences were noted and the immaturity of the colonies prevented
an accurate determination.
20 University of California Publications. | ZOOLOGY
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, pl. 39, fig. 2.
Calycella syringa, Calkins, 1899, p. 358, pl. 4, fig. 20.
Calycella syringa, Clark, 1876, p. 217, pl. 12, fig. 25.
Calycella syringa, Hartlaub, 1901, p. 358.
Calycella syringa, Nutting, 1899, p. 741; 1901, p. 176.
Calycella syringa, Torrey, 1902, p. 59, pl. 6, fig. 50.
Trophosome. Pedicels borne on stolon, shorter than hydrotheeae.
Margins of hydrothecae frequently reduplicated.
Gonosome. Gonangia on stolon, with acrocysts at maturity; ovate,
smooth.
Distribution. San Diego Bay, 1 to 5 fathoms. Puget Sound;
Berg Inlet and Kadiak, Al. (Nutting). Coal Harbor and Shum-
agin Is., Al. (Clark). White Sea (Mereschkowsky). Northern
Asia (Thompson). Kara Sea (Berg). British Coasts, Iceland
in 100 fathoms (Hincks). Greenland (Levinsen). Helgoland
(Hartlaub).
No gonosome in San Diego colonies, July 15, 1903.
Fam. LAFOEIDAE.
Gen. Lafoea, Lamouroux, 1821.
25. Lafoea dumosa (Flem.).
Serlularia dumosa, Fleming, 1828, p. 83.
Lafoea dumosa, Sars, 1862.
Lafoea dumosa, Hincks, 1868, p. 200, pl. 41, fig. 1.
Lafoea dumosa, Clark, 1876, p. 216, pl. 12, fig. 23.
Lafoea dumosa, Nutting, 1899, p. 747, pl. 64.
Lafoea dumosa, Torrey, 1902, p. 59.
Trophosome. Stem simple and creeping or fascicled and erect.
Hydrotheeae strong, narrowed 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 England Coast (Ver-
rill, Nutting). West Indies, 450 fathoms (Allman). British
Coasts (Hincks). Spitzbergen (Marktanner-T.). North Cape,
Norway (Sars). Helgoland (Hartlaub).
Vor. 2] Torrey—Hydroids of the San Diego Region. pa
Both the erect and creeping forms were found at San Diego.
Overgrown with C. hinckst. No gonosome, June 29, July 15,
1903. :
Fam. SERTULARIIDAE.
Anyone who has had oceasion 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
produce.
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-Turnerestcher (790) dis-
tinguished 18 genera. Nutting has reduced this unusual total
to 12; but that number, I am convineed, is still too large. The
distinetion between Thuwiaria 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, 7hecocladium is distin-
euishable 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 Syntheciwm, which
alone ocurs in American waters, by supporting them on combi-
nations of characters. Syntheciwm is accordingly based upon a
22 University of California Publications. [| ZooLocy
combination of strictly opposite branches, smooth margined hy-
drotheeae, absence of opereula, as well as the intrathecal origin of
the gonangia.
Analysis, however, does not reveal the strength which is
elaimed 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 perisarcal 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
to the character of the margin, or may be wanting altogether.
Among sertularians with smooth round margins, it is often deli-
eate, 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 membrane stretched straight
across the aperture like a drumhead.’’ Sertularella halecina (a
Synthecium aceording to Nutting) possesses a thin drumhead
operculum before the hydranth emerges for the first time, but
is non-operculate in the adult. Such facts only lead inevitably
to the conclusion of Hartlaub (:00, p. 8) that the absence of an
operculum is of no taxonomic consequence. In the third place,
Nutting does not appear to insist that Syntheciwm 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 alternans Allman. It is more probable that Nut-
ting included XS. 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 Syntheciwm. 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
gonangia.
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 S. 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
oceupied (: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
species.
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 S. halecina may prove more convincing. In the majority of
cases, the gonophores of this species arise within hydrothecae.
Oceasionally, 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-
theeal, are held by the same authors to be abnormalities which
may be disregarded in classification. The extrathecal origin of
the gonangia of S. halecina’ 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
extratheeal gonangia must lessen the importance of the usual
condition, and should not be disregarded for this very reason.
‘Tf 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.
24 University of California Publications. | ZOOLOGY
According to Nutting (:04, p. 42), ‘‘it occurs not infrequently
in several widely 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.
Fig. 14.—Sertularella Fig. 15.—S. halecina. Fig. 16.—S. halecina.
halecina. Portion of Portion of stem show- Portion of colony. Show-
stolon with gonan- ing origin of branch ing an extrathecal branch
gium (s); base of one just below hydrotheca (x).
stem, with intrathecal (x).
gonangium (i).
If the branches of S. halecina be now considered, it will be
seen that they, as well as the gonangia, emerge typically from
hydrothecae. In this respect, then, S. halecina is a Thecocla-
dium. Yet this character is not invariable. Occasionally
branches arise independently of hydrothecae. 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 hydrothecae, a familiar
origin of branches in the Sertulariidae. In fig. 16, the branch
is not associated with a hydrotheea in any way. All these cases
(except fig. 17) were found on the same colony; and it is inter-
Vou. 2] Torrey—Hydroids of the San Diego Region. 25
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-
cocladium, the origin of the second and third is typical of Ser-
tularella. Without denying the usefulness of the usual manner
:
“4
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
stem 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. [ ZooLocy
Sertularella halecina is at once a typical Synthecium in the
character of the hydrothecal margin and the intratheeal origin
of the gonangia, a typical Thecocladium in the intrathecal origin
of the branches, a typical Sertularella in the manner of the origin
of both branches and gonangia. Allman refers to the intrathecal
origin of two branches in Synthecitum campylocarpum as an
abnormality, and Nutting speaks similarly of the intrathecal
origin of the only two branches which were present in the mate-
rial from which I deseribed 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
taxonomically insignificant? Which are the abnormal, the insig-
nificant characters in S. halecina? 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.
26. Sertularella halecina.
Sertularella halecina, Torrey, 1902, p. 61, p. 6, fig. 55.
Trophosome. Stems from a creeping stolon rise to height of 30 mm.,
with few branches which originate either within hydrothecae or just below
them. Nodal divisions faint, often wanting. Hydrothecae adnate at base
only, cylindrical, with slight swelling on lower side 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 fathoms; growing on
kelp and among bryozoa. Gonosome present, July, 1901; June,
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 margins of the
hydrothecae are not so distinctly or characteristically everted.
Vou. 2] Torrey.—Hydroids of the San Diego Region. )
+I
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 adecauline side.
Gonosome. Gonangia ovate, covered thickly and completely with
slender spines.
Figs. 19, 20.—Sertularia pedrensis. Hydrotheeae. Fig. 21.— S. pedrensis.
Gonangium.
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, pl. 138, figs. 3-6.
Sertularella tenella, Alder, 1857, p. 113.
Serularella tenella, Hincks, 1868, p. 242, pl. 47, fig. 3.
Sertularella tenella, Hartlaub, 1901, p. 360, pl. 21, figs. 12, 20, 21.
Sertularella tenella, Torrey, 1902, p. 64.
Trophosome. ‘‘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. a 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 Fucus. No gonosome, July 16, 1901,
July 138, 1903. Longest stem, 4 mm.; length of hydrotheca, .4 to
.) mm., breadth, .25 mm.
29. Sertularella tricuspidata (Alder).
Sertularia tricuspidata, Aider, 1856, p. 356, pl. 13, figs, 1, 2.
Sertularella tricuspidata, Hincks, 1868, p. 239, pl. 47, fig. 1.
Sertularella tricuspidata, Clark, 1876, p. 224, pl. 12, figs. 26, 27.
Sertularella tricuspidata, Nutting, 1899, p. 741.
Sertularella tricuspidata, Hartlaub, 1901, p. 359.
Sertularella tricuspidata, Nutting, 1901, p. 183.
Sertularella hesperia, Torrey, 1902, p. 63, pl. 7, figs. 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 internodes, each of which bears a hydrotheca or a branch with
its axillary hydrotheea. Branches irregularly alternate, often branching
profusely either alternately or dichotomously, divided into regular inter-
nodes each of which bears a hydrotheea, 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).
Vou. 2] Torrey——Hydroids of the San Diego Region. 29
30. Sertularella turgida (Trask).
Sertularia turgida, Trask, 1854, p. 113, pl. 4, fig. 1.
Sertularella turgida, Ciark, 1876a, p. 259, pl. 38, figs. 4, 5.
Sertutarella conica, Calkins, 1899, p. 359, pl. 4, fig. 22.
Sertularella nodulosa, Calkins, 1899, p. 360, pl. 5, fig. 29.
Sertularetla turgida, Hartlaub, 1901, p. 360, pl. 21, figs. 5, 6.
Sertularella turgida, Torrey, 1902, p. 64, pl. 7, figs. 59-62; pl. 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. ‘xonangia large, ovate, distally spinose or annulated or
both; aperture small.
J
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 annulated type is reached through a spiny-annulated
condition. The hydrothecae are extremely variable, as regards:
shape, wrinkling and immersion. The internodes vary much in
leneth and thickness.
30 University of California Publications. [ ZooLoGy
Dynamena group.
31. Dynamena cornicina McCrady.
Dynamena cornicina, MeCrady, 1858, p. 204.
Sertularia complexa, Clark, 1879, p. 245, pl. 4, figs. 26-8.
Sertularia complexa, Bale, 1888, p. 769, pl. 18, figs. 1-4.
Sertularia cornicina, Nutting, 1901, p. 359, fig. 56.
Sertularia compleza, Nutting, 1901, p. 360, fig. 57.
Sertularia cornicina, Nutting, 1904, p. 58, pl. 4, figs. 1-5.
Trophosome. Stems short, 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. Hydrothecae tubular, adnate
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. (MeCrady). Woods Hole, Mass. (Nutting).
Pourtales Plateau (Nutting). Yueatan Coast (Clark). Aus-
tralia (Bale).
The colonies 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 synonymous.
D. cornicina is very close to 8. desmoides, from 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 desmoidis, Torrey, 1902, p. 65, pl. 8, figs. 70-72.
Sertularia desmoides, Nutting, 1904, p. 56, pl. 3, 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. ‘wo 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 opereulate 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 I., 42
fathoms: San Pedro, Cal., 13 fathoms. Albatross station 2939,
lat. N. 33° 36’, long. W. 118° 09’ 30’, 27 fathoms (Nutting).
Gonosome present, July, 1901, June 27, 1903. Both robust
and attenuated varieties were obtained.
Vou. 2] Torrey.—Hydroids of the San Diego Region. 31
33. Sertularia furcata Trask.
Sertularia furcata, Trask, 1854, p. 112, pl. 5, fig. 2.
Sertularia furcata, Agassiz, 1865, p. 145.
Sertularia furcata, Clark, 1876a, p. 258, pl. 39, fig. 3.
Sertularia furcata, Torrey, 1902, p. 66, pl. 8, figs. 73-5.
Trophosome. Stems short, unbranched, rising from a creeping
stolon to heght 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
Crum, Cal. (Clark):
Nutting (:04) has identified this species with the Dynamena
pulchella of d’Orbigny from Patagonia, quoting Clark’s deserip-
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 elaim 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,
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.
I 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. [ ZooLocy
Thuiaria group.
34. Sertularia filicula E. & S.
Sertularia filicula, Ellis and Solander, 1786, p. 57, pl. 6.
Sertularia anguina, Trask, 1854, p. 112, pl. 5, fig. 1.
Sertularia labrata, Murray, 1860, p. 250, pl. 11, fig. 2.
Sertularia filicula, Hincks, 1868, p. 264, pl.53, fig. 3.
Sertularia anguina, Clark, 1876a, p. 255, pl. 40, figs. 1, 2.
Sertularia anguina var. robusta, Clark, 1876a, p. 256, pl. 40,
figs. 3, 4, 5.
Sertularia filicula, Torrey, 1902, p. 68, pl. 9, fig. 80.
Sertularia filicula, Nutting, 1904, p. 117, pl. 34, fig. 1.
Trophosome. Stems with alternating branches, pinnately 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.
Gonosome. Gonangia pearshaped, produced somewhat distally, end-
ing with small round aperture.
Distribution. San Diego, 15-25 fathoms; San Pedro, San
Francisco, Cal., shore rocks. Monterey to Point Reyes, Cal.
(Trask). Vaneouver I. (Dawson). Alaska, 10 fathoms; San
Miguel I., Cal. (Clark). White Sea (Mereschkowsky). New
England coast (Verrill). Grand Manan, 20 fathoms (Stimp-
son). Labrador (Packard). Greenland (Levinson). North
Atlantic (Bonnevie).° Norway (Marktanner-Turneretscher).
British shores (Hincks).
Nutting’s treatment of Sertularia anguina Trask is unfor-
tunate. To begin with, the figures of his Abietinaria anguina
(Trask) are so far from typical of S. anguina Trask, judging
either from Trask’s figures, Clark’s figures or all 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. He says the
are from Santa Barbara, Cal., and Ber-
ee
specimens he has seen
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 shown 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 S. anguina Trask of
A A LL TS
Vou. 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 8. filicula E. & S. (:04, p. 68, pl. 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 1s apparently
distinct.’’ I take these statements to mean that if S. anguina
and WN. 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 faet that
Trask’s figure of S. anguina, though erude and containing an
error in showing 4 rather than 3 hydrothecae on the stem
between the bases of successive branches, Murray ’s figure of NS.
labrata, Clark’s figures of 8S. angwina, Hineks’ 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 (Pl. 34, fig. 1) of Abietinaria filicula (Ellis and
Solander) and not, curiously enough, with his figures of Abveti-
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, pl. 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 hydrocladium.
Mesial nematophore reaches level of hydrothecal aperture. Septal ridge
34 University of California Publications. | ZooLoGy
just below supracalycine nematophores and one just above floor of hydro-
theea. .
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 hydrothecae on anterior edge of all but
first and last. One, rarely two hydrothecae between corbula 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.
36 Aglaophenia inconspicua.
Agiaophenia inconspicua, Torrey, 1902, p. 73, pl. 9, figs. 87-89.
Trophosome. Stems stout, in clusters, 35-40 mm. high; divided by
antero-posteriorly oblique nodes into internodes as broad as long. Hydro-
cladia borne on same side of stem, alternate, one from each internode, 3-4
mm. long; divided transversely into equal internodes. A nematophore in
the axil of each hydrocladium and two at its base in a line parallel with
its axis. Hydrothecae deep, slightly compressed, free for not 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 internode. Mesial nemato-
phore reaching nearly or quite to the mouth of the theca. Supracalycine
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 theeate internode between corbula and stem. Sporosacs 6-12.
Distribution. San Diego, 5 fathoms; gonosome present,
July, 1901.
37. Aglaophenia pluma (Linn.).
Sertularia pluma, Linnaeus, 1767, p. 1309.
Aglaophenia pluma, Lamouroux, 1816, p. 170.
Aglaophenia pluma, Hincks, 1868, p. 286, pl. 63, fig. 1.
Aglaophenia pluma, Torrey, 1902, p. 73, pl. 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. Hydrothecae 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).
Vou. 2] Torrey—Hydroids of the San Diego Region. 35
38. Aglaophenia struthionides (Murray).
Plumularia struthionides, Murray, 1860, p. 251, pl. 12, fig. 2.
Aglaophenia franciscana, A. Agassiz, 1865, p. 140.
Aglaophenia struthionides, Clark, 1876a, p. 262, pl. 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. Hydrotheeae 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. Corbulae 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. Diplocheilus, Allman, 1883.
Trophosome. All internodes thecate, each internode with an infra-
calycine mesial nematophore not in contact with the hydrothcea, and a
supracalycine median sarcostyle without definite nematophore; each hydro-
theca wita anterior intrathecal ridge.
Gonosome. Gonangia unprotected.
Allman founded this genus on the foilowing characters: a
duplicature of the walls of the hydrotheeae ‘‘ forming an external
z)
ealyeine envelope,’’ a shield-like 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
hydrotheea, flanked by webs of perisare 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 D. mirabilis. Bale, however, is certainly in error in
finding in Jickeli’s Kirchenpaueria the bond of union. Accord-
ing to Jickeli’s figure (783, pl. 28, fig. 27), the hydroid for which
he erects the genus is an eleutheroplean plumularian—probably
a Plumularia-—with nematophores broken away. The frequent
absence of nematophores ir 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. 75, pl. 10, fig. 95.
Trophosome. Colony with simple stem, divided obliquely into inter-
nodes which vary in length according to age. Hydrocladia alternate, each
from a shoulder process projecting from the middle region of each inter-
node. Each hydrocladium divided more or less obliquely into equal
thecate internodes. Each hydrotheca somewhat 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-
tum about two thirds the length of the hydrotheca from the bottom,
reaching about one third across it at widest point. Cauline nematophores
absent with the exception of single axillary nematophores. Mesial nemato-
phore short, not reaching the base of hydrotheca, expanding into the
form of a sickle shaped segment of a saucer with a diameter two thirds
that of hydrotheca and embracing the internode for half its circumference.
Single median supracalycine sarcostyle, flanked by two webs of perisare
stretched between theca and internode, forming a non-typical median
nematophore.
Gonosome absent.
Distribution. Pt. Loma, Cal., on seaweed and sponges.
The differences which separate D. mirabilis Allman, D. pro-
ducta (Bale) and D. allmani are slight. D. allmani, originally
thought to be identical with D. producta, possesses hydrothecae
with flaring rims and broadly ovate apertures instead of the
compressed form and narrowed apertures of D. producta. It
differs from D. mirabilis in the absence of all cauline nemato-
phores save those in the axils of the hydrocladia, 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.
Vou. 2] Torrey.—Hydroids of the San Diego Region. 37
Gen. Plumularia, Lamarck, 1816.
40. Plumularia alicia.
Plumularia alicia, Torrey, 1902, p. 75, pl. 10, figs. 96, 97.
Trophosome. Stems in clusters, slender, loosely branching, 7 to 13
em. high: divided transversely by faint nodes into short equal internodes.
Hydrocladia atlernate, 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;
theeate internodes with 1 mesial and 2 supracalycine nematophores. Per-
isare 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, pl. 19, figs. 1, 2.
Plumularia megalocephala, Nutting, 1900, p. 57, pl. 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-
cator Reef, 14 fathoms (Allman). Albatross Station 2669, lat.
N. 3° 9% lone. W. 79° 33’, 352 fathoms (Nuttimeg).
‘“The internode interealated between the hydrotheea-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 that the hydrocladia
‘alternate as a rule, but not regularly so in some specimens,
where they are occasionally opposite toward the distal end of the
stem.
The San Diego material consists of two stems, the longest
measuring 100 mm., both unbranched. The stem internodes vary
in leneth, due to the obliteration of one or two nodes, and bear
one, two or three hydrocaldia respectively. The number of their
lateral nematophores varies with their length, from two to four.
There is a pair of nematophores on each basal process, also an
unpaired conical open process from which coenosare projected
in one ease, 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 that the members of every other pair lie in a
plane making an angle somewhat 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 part of the colony it is usually
wanting. Oceasionally a non-thecate internode fuses with a
theeate internode. The theeate internodes frequently bear two
mesial nematophores.
One stem had produced a heteromorphic shoot with several
hydrocladia alternately placed, as is the rule with the basal part
of the stem.
This species appears to be closely allied with P. filicula All-
man, but its hydrothecae are not so deep and its habit is less
regular.
42. Plumularia plumularioides (Clark).
Halecium(?) plumularioides, Clark, 1876, p, 217, pl. 10, figs.
NGA
Plumularia plumularioides, Nutting, 1901, p. 62, pl. 4, fig. 3.
Plumularia plumularoides, Torrey, 1902, p. 78, pl. 11, figs. 103,
104.
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
Vou. 2] Torrey.—Hydroids of the San Diego Region. 39
opposite sides of the stem, one to each internode, having their origin in
a small shoulder-like process just below each joint, divided usually into
regular internodes, though in some cases, short internodes occur hetween
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, pl. 66, fig. 1.
Plumularia setacea, Clark, 1876a, p. 261, pl. 41, figs. 1, 2.
Plumularia setacea, Nutting, 1900, p. 56, pl. 1, figs. 1-4.
Plumularia palmeri, Nutting, 1900, p. 65, pl. 6, figs. 4, 5; 1901,
p- 188.
Plumularia setacea, Torrey, 1902, p. 79, pl. 11, fig. 105.
Trophosome. Stems 5 to 100 mm. long, non-fascicled, divided into
internodes, each bearing a hydrocladium from a distal process. Hydro--
eladia 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 (Hineks). Helgoland (Hartlaub).
40 University of California Publications. [ ZOOLOGY
BIBLIOGRAPHY.
Notr.—Only papers directly concerned with Pacific Coast hydroids or
specifically referred to in the synonymies, are listed below.
Agassiz, A.
1865. North American Acalephae. Ill. Cat. Harv. Mus. Comp. Zool., No. 2.
Agassiz, L.
1860-2. Contributions to the Natural History of the United States.
ii Ve
Alder, J.
1856. A Notice of Some New Genera and Species of British Hydroid
Zoéphytes. Ann. & Mag. N. H., (2), XVIII, p. 353.
1857. A Catalogue of the Zodphytes 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.
1871. A Monograph of the Gymnoblastic or Tubularia Hydroids. Ray
Society.
1877. Report on the Hydroida collected during the Exploration of the
Gulf Stream. Mem. Harv. Mus. Comp. Zool. V, No. 2.
1883. Report on the Hydroida Dredged by H.M.S. Challenger. Pt. I.
Plumularidae.
1885. Description of Australian, Cape and Other Hydroida, mostly
new, from the Collection of Miss Gatty. Jour. Linn. Soe.
Lond., XIX, p. 132.
1888. Report 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. Proce. Linn. Soe. N. S. W., (2), ILI, Pt. I, p. 745.
1893. Further Notes on Australian Hydroids, with Descriptions of
Some New Species. Proc. Roy. Soe. 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 Pacifie Coast of the United States south of
Vancouver Island. Trans. Conn. Ac., III, p. 249.
Vou. 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, ete. 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, ete. London.
Ellis and Solander.
1786. The Natural History of many curious and uncommon Zodphytes,
collected from various parts of the globe. London.
Fewkes, J. W.
1889. New Invertebrata from the Coast of California. Bull. Essex
Inst., XXI; also separate, pp. 3-50, 7 pls.
Fleming, J.
1828. A History of British Animals. Edinburgh.
Hartlaub, C.
1894. Die Coelenteraten Helgolands. Wissensch. Meeresuntersuch., AE
Ite I, jos du@2e
1900. Revision der Sertularella-Arten. Hamburg.
1901. Hydroiden aus dem Stillen Ocean. Zool. Jahrb., Abth. Syst., ete.,
RAV; p. 349:
Hincks, T.
1861-2. A Catalogue of the Zodéphytes of South Devon and South Corn-
wall. Ann. & Mag. Nat. Hist., (3), VIII, p. 293.
1868. A History of the British Hydroid Zodphytes. London.
Johnston, G.
1847. A History of the British Zoéphytes. 2d. Ed., London.
Lamarck, J. B. P. A. de.
1815-22. Histoire Naturelle des Animaux sans vertébres. Paris. (Vol. °
SiG)
Lamouroux, J. F. V.
1812. Bulletin Société Philomatique.
1816. Histoire des Polypiers Coralligénes flexibles vulgairement nommés
Zoophytes. Caen.
1821. Exposition Méthodique des Genres de 1’Ordre des Polypiers.
Paris.
Lesson, R. P.
1836. Tableau de la Famille des Zodphytes Béroides. Ann. Sc. Nat.,
V, p- 204.
Linnaeus, C.
1767. Systema Naturae, 12th ed.
42 University of California Publications. | ZOOLOGY
Marktanner-Turneretscher, G.
1890. Die Hydroiden des k. k. Naturhist. Hofmus. Ann. Hofmus.
Wien, V.
McCrady, J.
1858. Gymnothalmata of Charleston Harbor. Proc. Elliott Soc. 8. C.,
Lp. wos:
Mereschkowsky, C.
1878. Studies on the Hydroida. III. Systematic Facts. Ann. & Mag.
IN; (5), Lops S22.
Murray, A.
1860. Description of New Sertulariadae from the California Coast.
Ann, & Mag. N. H., (3), V, p. 250.
Nutting, C. C.
1899. Hydroida from Alaska and Puget Sound. Proc. U. 8. Nat. Mus.,
XXI, p. 741.
1900. American Hydroids. Pt. I. The Plumularidae. Spee. Bull. No.
4.0. S: Nat Mus:
1901. Papers from the Harriman Alaska Expedition. XXI. The
Hydroids. Proc. Wash. Ac. Se., III, p. 157.
190la. The Hydroids of the Woods Hole Region. Bull. U. S. Fish
Comm., XIX, p. 325.
1902. American Naturalist, XXXV, p. 789.
1904. American Hydroidis. Pt. I]. The Sertularidae. Spec. Bull. No.
4, U. S. Nat. Mus.
Oken, L.
1816. Lehrbuch der Naturgeschichte. Weimar.
Pallas, P.
1766. Elenchus Zodphytorum. Hagae Comitum.
Peron et Lesueur.
1809. Tableau des caractéres génériques et spécifiques de toutes les
especes de Méduses connues jusqu’a ce jour. Ann, Mus.
d’Hist. Nat., XIV, p. 325.
Sars, M.
1835. Beskrivelser og Jagttagelser, etc. Bergen, p. 81.
1862. Bemaerkninger over fire norske Hydroider. Forh. Vidensk. Selsk.
Christiania.
Schneider, K. C.
1897. Hydropolypen non Rovigno, nebst Uebersicht des System der
Hydropolypen im Allgemeinen. Zo6l. Jahrb., Abth. Syst.,
ete., X, p. 472.
Thompson, D’A. W.
1877. The Hydroida of the Vega Expedition. Vega Exp. Vetensk.
Jakttagelser, IV, p. 387.
Vou. 2] Torrey—Hydroids of the San Diego Region. 43
POTey., Hews.
1902. The Hydroida of the Pacific Coast of North America. Un. Cal.
Publ., Zool., I, p. 1. Z
1902a. American Naturalist, XXXVI, p. 987.
IMEI Ue 153
1854. On Some New Microscopic Organisms. Proce. Cal. Ac. Se. 1853,
Ih jos, ILO),
Van Beneden, E.
1844. Recherches sur 1’Embryogénie des Tubulaires, ete. Nouv. Mém.
MNO; Iitib<, ROVINE, fos sk
Verrill, A. E.
1871-2. Report upon the Invertebrate Animals of Vineyard Sound, ete.
Rep. U. 8. Comm. Fish and Fisheries.
Wright, T. S.
1859. Observations on British Hydroid Zodphytes. Edinb. New Phil.
Aone, 5 fy, 2 joy INO),
UNIVERSITY OF CALIFORNIA PUBLICATIONS
ZOOLOGY
WOls Ay IN@s Zp joe Grorteils Veale le December 21, 1904.
CONTRIBUTIONS FROM THE LABORATORY
OF THE
MARINE BIOLOGICAL ASSOCIATION OF SAN DIEGO.
Tn
THE CTENOPHORES OF THE SAN DIEGO
REGION.
BY
HARRY BEAL TORREY.
Ord. CYDIPPIDA.
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.
Fam 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 compressed; rows of
swimming plates reaching from pole to pole; tentacle sheath present.
Three ctenophores taken in Prince Wilham Sound, Alaska, by
Professor Ritter, and now in the collection of the University of
California, belong undoubtedly to Mertens’ Beroe cucumis
(EKuplokanis cucumis, Chun). All are compressed somewhat, so
that the transverse diameters are to each other as 6 to 5. Plewro-
brachia, 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 cireular cross section of Euplo-
kamis, which is rather to be distinguished from Pleurobrachia by
its elongation, from Mertensia by its sight 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
broad.
46 University of California Publications. | [Zoorocy
1. Euplokamis californensis, 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 July. None were more than
25 mm. long. This species is very closely related to E. cucumis,
with which it may prove to be identical. It is near, also, to the
Cydippe elliptica of Eschscholtz, from the tropical Pacifie.
Pleurobrachia, Fleming, 1822.
Body spherical, interradial canals from two stem canals.
2. Pleurobrachia bachei A. Ag.
Pleurobrachia bachei A. Ag., L. Agassiz, 1860, p. 294.
A. Agassiz, 1865, p. 34.
Oesophagus equal to or less than funnel tube in length; tentacle
sheaths distant from funnel, about 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 the exception of the
tentacles, which are yellowish red, and the oesophagus, which is blotched
with deep purple proximally.
Distribution. San Diego to Puget Sound. This species
differs from P. pileus (Fabr.) 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.
Ord. LOBATA.
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.
Vou. 2] Torrey.—Ctenophores of the San Diego Region. 47
Fam. BouINIDAE.
Bolina, Mertens, 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. microptera A. Ag., from the Gulf of Georgia.
Agassiz and Mayer have described another, Hucharis grandi-
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’ cireumboreal 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
accuracy.
Ord. BEROIDA.
OCtenophorae elongated, conical or ovate, compressed, with
large mouth and oesophagus. Tentacles and tentacle canals
wanting. Meridional canals communicate with oesophageal
eanals at the edge of the mouth, and send out numerous branches
which may form a peripheral network.
Fam. BEROIDAER.
With the characters of the order.
Beroe, Browne, 1756.
With the characters of the family.
4. Beroé forskali M. Edw., Chun.
?Beroé rufescens Forskal, 1775, p. 111.
Cydalisia mitraeformis, Lesson, 1843, p. 138, pl. 2, fig. 2.
Idya penicillata, Mertens, 1833, p. 534, pl. 12.
Beroé Forskalii, Milne-Edwards, 1841, p. 207, pl. 5.
Beroé Forskalii, Chun, 1880, p. 309, pl. 14, figs. 3-5; pl. 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. Rows 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. Agassiz, 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.
Vou. 2] Torrey —Ctenophores of the San Diego Region. 49
BIBLIOGRAPHY.
Agassiz, A.
1865. North American Acalephae. Il]. Cat. Harv. Mus. Comp. Zool, No.2.
Agassiz and Mayer.
1899. Acalephs from the Fiji Islands. Bull. Harv. Mus. Comp. Zodl.,
XXXII, No. 9.
Agassiz, L.
1860. Contributions to the Natural History of the United States, ITI
p. 153.
,
Brandt, J.
1838. Ausfuhrliche Beschreibung, etc. Mém. Ac. St. Petersb., (6), IV,
Pp. 237.
Browne, P.
1756. Civil and Natural History of Jamaica.
ChunkiC:
1880. Die Ctenophoren. Fauna tind Flora des Golfes von Neapel.
1897. Die Beziehungen zwischen dem Arktischen und Antarktischen
Plankton.
1898. Die Ctenophoren. Ergebnesse der Plankton Expedition, II:k:a.
Eschscholtz, F.
1829.. System der Akalephen. Berlin.
Fabricius, O.
1780. Fauna Groenlandica. Hafniae et Lipsiae.
Fleming, J.
1822. Phil. Zodlogy.
Forskal, P.
1775. Descriptiones Animalium, ete. Hauniae.
Lesson, R. P.
18438. Histoire Naturelle des Zodphytes. Acalephes. Paris.
Mertens, H.
1833. Beobachtungen und Untersuchungen iiber die Beroéartigen Aka-
lephen. MWiém:. Ac. St. Petersb., (6), IL. p. 479:
Milne-Edwards, H.
1841. Observations sur la structure et les fonctions de quelques Zo6-
phytes, Mollusques et Crustacés des cétes de la France. Ann.
Slob INEhis (2))q 2 WAI, jos 1B)
Morch.
1857. Naturh. Bid. til en Besk. af Groénland.
Vanhioffen, E.
1895. Die Gronlandischen Ctenophoren. Bibl. Zool., VIII (h. 20), p. 15.
1904. Nordisches Plankton. XI. Ctenophoren.
1
EXPLANATION OF PLATE 1.
Fig. 1.—Mertensia ovum.
Fig. 2.—Beroé forskali.
Fig. 3.—Pleurobrachia bachei.
(50)
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UNIVERSITY OF CALIFORNIA PUBLICATIONS
ZOOLOGY
Vol. 2, No. 3, pp. 51-112, Pls. 2-3. January 31, 1905
CONTRIBUTIONS FROM THE LABORATORY
OF THE
MARINE BIOLOGICAL ASSOCIATION OF SAN DIEGO.
wy
THE PELAGIC TUNICATA OF THE SAN
DIEGO REGION, EXCEPTING
THE LARVACEA.
BY
Wo. 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 ease, 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. [ ZooLocy
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. From this duplication of names 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, 1546, the majority, probably, of authors have retained
both the specific names, writing them with a hyphen, thus, Salpa
democrata-mucronata. Others, on the contrary, have retained
but one of the names, but 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 SN.
confoederata-scutigera. I have followed Krohn, with the slight
modification that I have uniformly placed that name first in the
couplet that was proposed at the earlier date. This has reversed
the order in several species, as, for example, S. fusiformis-runci-
nata, this being written by other authors runcinata-fusiformis.
But as fusiformis was proposed by Cuvier in 1804, while runci-
nata 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:
SALPIDAE.
Cyclosalpa bakeri, n.sp.
Cyclosalpa affinis.
Salpa fusiformis-runcinata.
Salpa tilesvi.
Salpa democratica-mucronata.
Salpa zonaria-cordif ormis.
Salpa cylindrica.
Salpa conf oederata-scutigera.
DOLIOLIDAE.
Doliolum tritonis.
Doliolum ehrenbergii.
Doliolum miilleri.
VoL. 2] Ritter—The Pelagic Tunicata. 53
PYROSOMIDAE.
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 eylin-
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-
phie oozooid and blastozooid generations.
Fam. I.—SaAupipak, 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, Cuvier, 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.
Holothuria, Linn, 1758.
Dagysa, Banks and Solander, 1773.
Biphora, Bruguiére, 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
eroups consisting of two parallel series.
Cyclosalpa bakeri n. sp.
(a) Solitary (budding) generation.—Pl. 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,’ Pl. IT, fig. 1; section of body everywhere cir-
cular. Length of largest zooid seen, 41 mm., this with
salpa chain well developed. Test 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 hp of branchial orifice truncate,
lower lip rounded 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 bands except ninth and tenth,
and sometimes eighth, interrupted dorsally, and all except first,
ninth and tenth interrupted ventrally. First trending backward
on dorsal side, the ends becoming nearly parallel and reaching
as far back as the third; also connected on dorsal side with the
posterior lip band 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, Pl. II, fig. 2.
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, Pl. II, which outlines
the form of a specimen alive and swimming. From this observation, and
from the fact that C. affinis 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.
Vou. 2] Ritter—The Pelagic Tunicata.
On
On
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. Guill 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 lke 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
bands.
(b) Aggregate (sexual) generation. Pl. III, 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. [ ZooLocy
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 ean only be said at present
that the close similarity of this species to Cyclosalpa 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 a 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 first and second body muscles.
Branchial orifice terminal, lps (at this 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 definite 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 a little distance from median dorsal line, the branches unit-
ing again about opposite the endostyle to extend into the ventral
peduncle, Pl. III, fig. 7. Third muscle likewise bifureates about
same distance as second from dorsal median line, the anterior
branch extending down to cross the 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,’’ PI. III, fig. 7.
Arrangement on side turned toward posterior end of parent is
—
. enx™~ne ~~
VoL. 2] Ritter——The Pelagic Tunicata.
>)
|
as follows: second muscle not bifurcating ; third bifureating near
dorsal line as on opposite side, the anterior branch again bifur-
eating to send a branch forward which anastomoses with second
muscle, the single band thus produced passing into the ventral
pedunele, Pl. III, fig. 8. In the asymmetry of the muscles this
species resembles SN. rostrata, S. punctata, S. magalhanica of Ap-
stein. The climax of the phenomenon is reached in SN. rostrata.
First and second muscles united on each side by two longi-
tudinal bands, Pl. III, fig. 7. In addition to the dorsal
hp 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 vet
complete, even in the oldest zooids found. With little doubt the
final form is a cirele 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. [| ZooLocy
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., figs. 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
band.
C. bakeri appears 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. bakeri 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. baker.
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
hp 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, pl. figs. 2A-C, solitary genera-
tion; 2D-H, aggregate generation.
Cyclosalpa affinis Blainville, 1827.
Salpa affinis Meyen, 1832, p. 407.
Salpa affinis Traustedt, 1885, p. 357, Pl. I, 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.
60 University of California Publications. | ZooLocy
(a) Solitary (budding) generation—F ig. 9. Body distinetly
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
Fig. 9.—S. affinis, solitary generation.
asperities; a pair of lateral and somewhat dorsal appendages at
‘
the posterior end. No lineaform ‘‘glandular’’ organs. Leneth
of full grown zooids, 80 to 100 mm. Branchial orifice directed
somewhat ventrad, lips prominent, upper overarching. Atrial
orifice without lips, directed somewhat dorsad, armed, particu-
larly in older specimens, with a pair of latero-dorsal appendages.
Body muscles eight, all excepting last 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-
tinetly 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 (fig. 11). Gill long and narrow, extending from slightly
in front of the anus to the esophagus, terminating, consequently,
—-.
VoL. 2] Ritter —The Pelagic Tunicata. 61
ti
Fig. 11.—Hypophysis of C. affinis.
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 body muscle band.
Fig. 10.—C. affinis, aggregate generation.
(b) Aggregate (sexual) generation.—Fig. 10. Agegregations
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 consisteney, 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, all continuous across
the dorsal side, and all 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 first atrial.
A well developed band in each lip of the branchial orifice meet-
ing in a wide angle on each side. First atrial band nearly as
broad as body bands; the other atrials, 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 em. 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 yellow, nearly uniform throughout. Heart conspicuous,
immediately in front of intestinal ring on ventral side. Placenta
A A a eS
Vou. 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. affinis 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, ean be fully appreciated only by examining the living
swimming animal. Preserved specimens have more the form of
the figures of the authors above mentioned. Again, the two
64 University of California Publications. | ZooLocy
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 em. 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, Pl. 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, 1904, in
the last named locality, and was reproducing actively, both sex-
ually and asexually.
Salpa fusiformis-runcinata Cuvier.-Cham.
Salpa fusiformis Cuvier, 1804, p. 23, fig. 10.
Salpa runcinata Chamisso, 1819, p. 16, Pl. figs. 5A-51.
Salpa runcinata-fusiformis Krohn, 1846, p. 112.
Salpa runecinata-fusiformis, Leuckart, 1854, p. 3 ad seq. Pl. I, figs.
6, 8, 16, 17, 18; and Pl. IT, figs. 1, 3, 4; 5, 13, 15, and 18.
Salpa runcinata-fusiformis Traustedt, 1885, p. 370, Pl. 2, figs.
29, 30, 31.
Salpa runcinata-fusiformis Herdman, 1888, p. 76, Pl. 6, figs. 5-12.
Salpa fusiformis Apstein, 1901, p. 1117, figs. 6a and 6b.
Fig. 12.—S. fusiformis-runcinata, solitary generation.
(a) Solitary (budding) generation —Fig. 12. Nearly eylin-
drical, somewhat larger at the atrial end, both orifices terminal,
truneate in general effect at both ends. Length of largest speci-
os Si _
ee
ee
Vou. 2] Ritter—The Pelagic Twnicata. 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 distinetly 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
66 University of California Publications. [ ZooLoGY
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.—S. 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 at 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
Vou. 2] Ritter—The Pelagic Twnicata. 67
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.
Fig. 14.—S. fusifornis-runcinata, echinate form. Postero-
dorsal view, showing serrations of test, and muscle bands.
pop.
1
'
1
|
i
!
i}
; t
, ' 1
pap. : i S772.
s.m0.T lr
Mig. 15.—S. fusiformis-runcinata, echinate form, ventral view
of surface.
68 University of Califorma Publications. | ZOOLOGY
Having worked over a large quantity of material of S. 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 ealled 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 echination of the test, the other the
arrangement of the body muscles. Figs. 14 and 15 are dia-
erams, though made with special care, from the examination of
three specimens which agreed almost perfectly in these partic-
ulars. Fig. 14 is a postero-dorso-dextral ‘view. It shows two
double rows of echinations on the dorsal surface that begin some
distance behind the anterior end and extend to the posterior end,
but do not terminate in spines. lLaterally from these is a row
on each side on the edge of a prominent ridge—almost a fin—
of test, this ridge extending the entire length of the body and
terminating posteriorly in two prominent processes, 7. and 7. p.
d. p. On the ventral side are two sets of submedian rows (Fig.
15, s. m. r. and s. m. 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. v. 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 fusi-
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
ar he een tases waaay
a ig el Oa
Vou. 2] Ritter—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 ease 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 seareely 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
fusiformis-runcinata.
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-
70 University of California Publications. | ZOOLOGY
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-runci-
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, Pl. 86,
figs. 1-5.
Salpa costata-tilesii Krohn, 1846, p. 114.
Salpa costata-tilesii Traustedt, 1885, p. 379, Pl. 1, figs. 10 and 11;
and Pl. Il. figs. 38-41, and 47.
Salpa costata-tilesii Herdman, 1888, p. 60, Pl. 4, figs. 1. 4, 8.
Salpa costata Brooks. 1893, p. 10 (particularly), Pl. IV, fig. 4;
Pl. VIII, fig. 4,
Salpa Tilesii Apstein, 1894, p. 16; 1901, p. 111 10, figs. lla, 11b.
*
\ ; Fi f | | eee hie AVP D> 2
= held Les ee ars
: | Sper \ 9 ¢ “
f
ooernad |
Fig. 16.—S. tilesii-costata, solitary generation.
(a) Solitary (budding) generation.—Fig. 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 em., exclusive of appendages, which were
47 mm. Test thick and firm, particularly on the ventral side,
— ———'— —~a= a eee ep
———_————- = @&
VoL. 2] Ritter.—The Pelagic Tuncata. 71
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. Guill 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
tuberele, on which the rather large, irregularly triangular mouth
mouth is situated.
Fig. 17.—Costata-tilesii, aggregate generaion.
(b) Aggregate (sexual) generation.—F ig. 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 em. 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 em. 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, and
not extending more than half way down to the mid-ventral line.
The first three drawing together, but not 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, but 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 bands in the ventral lip, the dorsal and ventral lp bands
crossing and intermingling in a complex but somewhat variable
way at the angles of the orifice. Two short longitudinal bands
in dorsal lip. Numerous delicate bands in atrial siphon, all
confluent with a longitudinal bantl on each side. Endostyle
slender, nearly straight, reaching back 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 body muscles.
a
Se ee
Vou. 2] Ritter—-The Pelagic Tunicata. 73
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 democratica Forskahl, 1775, p. 113, Pl. 36, fig. G (solitary
gener. )
Salpa mucronata Forskahl, 1775, p. 114, Pl. 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 Leuckart, 1854, p. 3, et seq., Pl. I,
figs. 1, 3, and numerous others.
Salpa democratica-mucronata Traustedt, 1885, p. 365, Pl. 2, figs.
25 and 28.
Salpa democratica-mucronata Herdman, 1888, Pl. VILI, figs. 1-10.
Salpa democratica Brooks, 1893, pp. 6-16, particularly for anat-
omy. Pl. 2, especially.
Thalia democratica-mucronata Herdman, 1899, p. 748.
Salpa mucronata Apstein, 1901, p. IIT 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
74 Universily 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 distinetly higher
and broadly notched in middle. Atrial orifice nearly terminal,
but inclined a little upward, without lps. <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-
eate, the sixth particularly so. A single rather broad muscle
band 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. Guill 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 from surface view wholly disconnected
from it; rather small, somewhat triangular in dorsal view.
Nucleus very small, ovate, situated far back, ight 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.
Vou. 2] Ritter.-—The Pelagic Tunicata. (5)
(b) Aggregate (sexual) generation—Fig. 19. ‘Agreeing
with solitary generation in general form, size, orifices, thickness
and transparency of test, endostyle, gill, ganglon 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
nucleus.
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. [| ZooLocx
Salpa zonaria-cordiformis, Pall.-Quoy et Gaim.
Holothurium zonarium Pallas, 1774, Fasl. X, p. 26, Pl. I, figs. 17,
A, B, and C.
Salpa polycratica Forsk., 1775, p. 116, Pl. 36, fig. F.
Salpa zonaria Cham., 1819, p. 12, Pl. figs. 1A to G.
Salpa cordiformis Quoy et Gaim., 1327, p. 225, Pl. 8A, figs. 3-6.
Salpa cordiformis-zonaria Krobn, 1846, p. 112.
Salpa cordiformis-polycratica Vogt, 1854, p. 7.
Salpa cordiformis-zonaria Traustedt, 1884, p. 382, Pl. II, figs.
18, 19, 20, and 21.
Salpa cordiformis-zonaria Herdman, 1888, p. 70, Pl. VII, figs. 1-9.
Salpa cordiformis Brooks, 1893, p. 10 (particularly); Pl. IU,
figs. 2 and 3; Pl. IV, figs. 3, 5, 6; Pl. VIII, fig. 5.
Salpa zonaria Apstein, 1894, p. 19; 1901, p. ITT, 10, figs. 10a, 10b.
Fig. 20.—S. cordiformis-zonaria, solitary generation.
(a) Solitary (budding) generation.—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; somewhat quadrilateral, though
this perhaps due to shrinkage, and may be characteristic of pre-
served animals only. Long axis almost perfectly straight. Wholly
colorless except for light lemon yellow of nucleus. Maximum
length of zooids so far obtained in our area, 35 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 the third, considerably longer, situated in the median ven-
tral plane and directed somewhat ventrad as well as backward.
Body muscle bands six in number, exceedingly broad, all
quite parallel with one another in the course around the body;
all interrupted both dorsally and ventrally, and by about the
~]
ba |
VoL. 2] Ritter—The Pelagic Twnicata.
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 shghtly into the pharyngeal cavity, situated exactly in the
sagittal plane. Ganglion rather small and eye spot seareely 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.—8S. zonaria-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 Californa 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
bifureates on each side. Sixth band bifureates on right 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 right side. (The
asymmetry of this muscle in this species may be compared with
that of the muscles of Cyclosalpa bakeri.)
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. Gill 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.
i ee a ee
Vou. 2] Ritter-—The Pelagic Tunicata. 79
Salpa cylindrica Cuvier.
Salpa cylindrica Cuvier, 1817, p. 22, figs. 8 and 9. (Solitary
generation. )
Salpa cylindrica Sav. 1816, p. 124, Pl. XXIV, figs. 2,, 2., 23.
(Tasis cylindrica in description of figures.)
Salpa cylindrica Traustedt, 1884, p. 377, Pl. II, figs. 35, 36, 37.
Salpa cylindrica Herdman, 1888, p. 72, Pl. VII, fig. 10.
Salpa cylindrica Brooks, 1893, Pl. ITI, figs. 5, 6, 7; P1.VIII, fig. 2.
Salpa cylindrica Apstein, 1894, p. 16.
Fig. 22.—S. cylindrica, solitary generation.
(a) Solitary (budding) generation. Wig. 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 bands 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
hip and two in ventral; a segment of a cireular 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 leneth 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) Aggregate (sexual) generation.—Fig. 22’, copied from
Brooks. Having seen no specimen of this generation, I give a
translation of Traustedt’s Latin diagnosis: ‘‘Body ovate; ex-
tremities terminating in conical, searcely 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
eoast. I 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 Calli-
fornia. Curiously enough, all the specimens so far seen, with
one doubtful exception, were of the solitary generation.
Salpa confoederata-scutigera Forsk.-Cuvier.
Salpa confoederata Forsk., 1775, p. 115, Pl. 36, fig. A.
Salpa scutigera Cuvier, 1817, p. 18, figs. 4 and 5.
Salpa octofora (?)* Cuvier, 1817, p. 20, fig. 7.
‘While I 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] Ritter-——The Pelagic Tunicata. 81
Salpa (Pegea) octofora, Sav., 1816, p. 124, Pl. 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, Pl. II,
figs. 23, 24.
Salpa scutigera-confoederata Herdman, 1888, p. 84, Pl. IX, fig. 9.
Pegea confoederata Lahille, 1890, p. 12, text, figs. 1, 2, 3A, 3B.
Salpa scutigera Brooks, 1893, pp. 6-16, anatomy; Pl. IV, figs. 1-7
Salpa confoederata Apstein, 1894, p. 12, Pl. Il, 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.
ANY a
anh Mins —
Fig. 23.—S. confoederata-scutigera, aggregate generation.
(b) Aggregate (sexual) generation— Fig. 23. Agerega-
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. [ ZooLocy
red or iron rusty pigment widely present in the mantle, but most
pronounced at two ends, and more abundant posteriorly than
anteriorly.
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 bands on the dorsal side behind the branchial
orifice, the apices directed backward, and the dorsal limb of each
reaching well into the dorsal lp to serve as a retractor of
it. (Dorsal lmb 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,
straight except for a distinct ventral curvature just short of the
posterior end, the posterior termination separated from the
nucleus by the length of the pericardium. Guill relatively rather
short, beginning but little in front of the first 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, shghtly
compressed dorso-ventrally, projecting but shghtly, 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 area, 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, June, July, and August.
Fam. IJ].—Do.uiouipAE Bronn, 1862.
Thaliacea in which the body form is typically and perfectly
that of a barrel; the body muscle bands are complete rings; and
Vou. 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 nucleaform; 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
Dolchinia not occurring, so far as known, in our area, are not
taken into account in the diagnoses. )
Three species, namely, D. tritoms, D. ehrenbergu and D. mil-
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,’’ @.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 has
begun.
84 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 :
Gonozooid
|
Embryo = Beginning of cycle
|
Oozooid
}
Bud
\
\
\
\
Trophozooid |
\
\
Phorozooid \
Gonozooid
Embryo = Beginning of cycle
A great variety of nomenclature has been employed by dif-
ferent writers to designate the several generations of Doliolum.
That here used, though not without objections, appears to me
to be the best of any yet proposed. It is in the main from Herd-
man, 1888 and 1904.
The chief synonyms found in the literature of the subject
are as follows:
Gonozooid (Herdman); sexual animal (Krohn, Grobben,
Uljanin) ; sexual generation A (Keferstein und Ehlers).
Ooozooid. (Herdman) ; asexual animal (Krohn, Gegenbaur) ;
generation B (Keferstein und Ehlers); first nurse generation
(Grobben) ; nurse (Uljanin) ; blastozooid (Herdman, ’88).
Trophozooid (Herdman) ; lateral buds, generation C' (Kefer-
stein und Ehlers); lateral buds (Grobben) ; nutritive animals
(Fol. Uljanin).
Phorozooids (Herdman); median buds (Krohn, Gegenbaur,
Grobben) ; generation C™ (Keferstein und Ehlers) ; foster-ani-
mals (Uljanin).
Vou. 2] Ritter—The Pelagic Tunicata. 85
Doliolum tritonis Herdman.
Doliolum denticulatum Herdman, 1883, p. 101, Pls. 18, 19, and 20.
Doliolum tritonis Herdman, 1888, p. 47, Pl. 3, fig. 3.
Doliolum tritonis Traustedt, 1893, p. 4, Pl. I, fig. 10.
Doliolum tritonis Borgert, 1894, p. 19, Pl. 5, figs. 17 and 18.
Doliolum tritonis Borgert, 1901, p. III 3, fig. 3.
Fig. 24.—Doliolum tritonis, gonozooid.
(a) Gonozooid.—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 lttle 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
edge on a 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 pylorie 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) Oozooid.—(See Figs. 27, 28, and 29 of D. ehrenbergii and
miller, this generation, which have the essential features of the
present species.) Body of the usual doliolum form; muscle bands
nine, first and ninth relatively very narrow, seventh band 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? (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
VoL. 2] Ritter—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 internal 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. miilleri and D. ehren-
bergu, with which it occurs in this region. Old individuals (of
this species, probably) reaching a leneth 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 deseribed 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 gonozooid of D. ehren-
bergu I assume to belong to this latter species. Similarly, those
in which the organ is the same as that in D. miilleri, 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
milleri, 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 searcely recognizable. This is likewise true of the
00zo01d 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
88 University of California Publications. | ZooLoGY
an oozooid constantly occurring 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 7mm. 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 to 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 be attained by the oozooids of any of the species.
Uljanin, *84, has recorded the occurrence in the Mediterranean
of a nurse measuring 30 mm. in length, and this he assumes
belongs to D. ehrenbergii. The identification is, however, by no
means certain. Asa matter of fact, in the present state of knowl-
edge, I do not believe it possible to assign with certainty any
oozooid, after the degeneration 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
these old nurses is obvious.
Fig. 25.—D. tritonis, Phorozooid.
(ec) Phorezooid.—F ig 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 lttle 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-
————
Vou. 2] Ritter—The Pelagic Twnicata. 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.
Phorzooid. Gonozooid. Phorzooid. Gonozooid.
Dge lav eth eh, s opens (o) ohs <irs auaps 35 (Sd PCR CECI ROEEEESS CORE 40
Mpreh cao erste, ae ate! stoys 35 AR frat a ayer Oansrsye docgahkona) ane 31
Se ea iar gAaSH Sy ache tol sl 1 is 42 Si vavekstetanncmarohensy avisteis ois 37
CHa Popecase’s te cea S siehie 33 MOM weiaie/auo a tais st Ste Slee a 90
DET atone erdete 3 Oo Md Masher cl taaustiers sie hep asebaneay 35
Gi A acbadecks tls hs Ste uiebavs 30 1 WY re areets cicr ee ee 83
Oe aispsontseauec ro tenciis snakes 38
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. Pedunecle very broad at distal end and narrowing
toward the body of the zooid; length of largest specimens seen,
90 University of California Publications. [ ZooLocy
3 mm.; 2 mm. broad in broadest part. Endostyle forming
considerably less than a right angle with the main axis of the
‘body. 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 sheht 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, d.m., 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, ly., need no special comment for the present pur-
pose. The peripharyngeal band, p.ph.b., 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 excretory. 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 strueture, and particularly the
physiology, of this generation of Doliolum 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 em.
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] Ritter.—The Pelagic Tunicata. 91
to ehrenbergu or miilleri, 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-
clusion.
D. tritons 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 Uljanin (not Krohn).
Doliolum Gegenbauer, 1856, Pl. XVI, figs. 12 (?) and 13 (2),
and fig. 15.
Doliolum Gen. 2B and 4B, Keferstein und Ehlers, 1861, p. 68, Pl.
IX, figs. 5 and 7; and Pl. X, fig. 4.
Doliolum denticulatum Grobben, 1882, pp. 23-41, Pl.1, figs. 3, 4,
and 5; Pl. 2, figs. 7 and 8.
Doliolum ehrenbergii Uljanin, 1884, pp. 132-133, Pl. 5, figs. 1 and
3; Pl. 12, fig. 8. (Numerous other figures are assigned to
. ehrenbergu, but I here refer to only those that undoubt-
edly represent the species as here understood.)
Doliolum ehrenbergii Herdman, 1886, p. 46, Pl. III, figs. 5 and 7.
Doliolum ehrenbergii Lahille, 1890, p. 65, figs. 47, 48, 49, and 50.
(a) Gonozooid. Unknown.
he g 0
Fig. 27.—D. ehrenbergii, 00z001d, before loss of internal organs.
(b) Oozooid—Fig. 27. Leneth from .925 mm. to 2.77 mm.,
between disappearance of larval characters and beginning of
_
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. Branechial 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 sixth muscle
band. Intestine rather long, extending back 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 between third and fourth muscles.
Otolith sae 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
4
pals fe
Fig. 28.—D. ehrenbergii, oozooid, after complete degeneration
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 milleri, and this accounts for the fact that while t77-
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 gegenbaurti zooids in the same
92 University of California Publications. | ZOOLOGY |
VoL. 2] Ritter—The Pelagic Twnicata. ' 98
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,
1904:
No. Lengh of Zooid. Width of Muscle. Condition of Intestirfe.
UE etoeege eis Cb 2.22 mm. .20 mm. organs intact
isi Aer roca ore 2.59 mm. .18 mm. organs far degenerated
Big Sino came Gere 2.60 mim. .129 mm. organs intact
Gh, Such Sa eRe eRe 2.77 mm. .148 mm. organs intact
Descenscpecanec she pir2 3.33 mm. 27 mm. organs nearly gone
\Digraoneiore A cman 4,25 mm. .o7 mm. organs wholly gone
Ucediare Rieter atte 5.00 mm. oo 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 ehrenbergui 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 gegenbaurvi either
to D. denticulatum or to any form supposed to belong to this
species, the real question is, Do all the forms assumed by Uljanin
to belong to D. denticulatum, and hence ealled by him gegen-
bauru, 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. | ZOOLOGY
named species? I agree with Borgert that the latter is the case.
Since, consequently, D. gegenbaurti 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 Pl. XVI, fig. 15, I have
little doubt. I am strongly of the opinion, too, that ‘‘Gen. 2B and
4B”’ of Keferstein und Ehlers, and shown in their Pl. IX, figs.
5 and 7, and Pl. X, fig. 4, likewise belong to the same species, as
do also Grobben’s D. denticulatum, shown in his PI. I, figs. 3
and 4, particularly, and pretty certainly also in fig. 5. As to his
fig. 7, Pl. 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 has D. tritonis. So far it has not been taken in the fall and
winter months, unless some of the few old oozooids at present in
doubt as to species belong here.
(e) 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 Pl. IT, fig. 8. 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 deseribed and figured by
this author apparently differs from the trophozooid assigned by
me to D. tritonis, fig 26, in the following 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.
VoL, 2] Ritter —The Pelagic Tunicata. 95
Doliolum miilleri Krohn.
D. milleri Krohn, 1852, p. 58, Pl. II, fig. 4.
D. nordmanni Krohn, 1852, p. 59, Pl. II, fig. 6.
Doliolum sp. Gegenbauer, 1856, Pl. XV, fig. 8.
Doliolum Gen. 3B, Keferstein und Ehlers, 1861, p. 68, Pl. X, fig. 3.
Doliolum miilleri Grobben, 1882, pp. 55-65, Pl. II, figs. 9 and 10;
Pl. II, 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: Pl. 4, figs.
1, 3) 4,nand_ 9, larvees Ply, fig. 11; PL 8, fig. 10; Pl. 9,
fos Pl ti; ho 9: Pil 12) fess 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, I give the diagnosis of all the generations, relying on
Uljanin, the most recent writer, chiefly, for all the generations
except the oozooid, this bemg made mainly from my own obser-
vations.
(a) Gonozooid.—Lengeth 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 intermusecular 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.
96 University of California Publications. [| ZooLoGy
SS SL
Fig. 29.—D. miilleri, oozooid, before degeneration of internal
organs.
(b) Oozooid. Fig. 29. Maximum length before beginning of
degeneration of intestine, something less than 2 mm.; test rela-
tively thick and 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 otoecyst differing in no easily recognizable way
from the corresponding parts in D. ehrenbergu. Intestinal tract
U-shaped, esophageal mouth very near the center of the branchial
membrane, whole tract in fifth intermuscular space.
Dorsal process short and thick, with but few buds, sometimes
having a pronounced kink near the base, but never, so far as
observed, with the top-shaped terminal piece characteristic of
D. ehrenbergu.
In this, as in the other species, little of positiveness is known
about the old ‘‘nurses.’’ Several times catches containing
oozooids 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. 1B,’’ Pl. X, fig. 1. These I suspect belong to the present
species. I do not imagine this pecular form to be wholly dis-
tinective. It is probably due to contraction; but it certainly does
not occur frequently in old nurses of what I suppose to be f77-
tonis and ehrenbergu.
Vou. 2} Ritter—The Pelagic Tunicata. 97
(ec) Phorozooid.—Entirely similar to gonozooid except for
absence of gonads and presence of ventral process which is rela-
tively long and eylindrieal.
(d) Trophozooid—Small, but long in proportion to breadth,
and with a relatively long peduncle.
As already said, D. miilleri is the least abundant of our spe-’
cies. Thus far it has been taken in mid-summer only, off San
Diego.
Order I1.—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.
Fam. PyrosoMIDAE T. Rupert Jones, 1848.
Pelagic colonial Ascidiacea, with the colony in the form of a
hollow eylinder 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 eylinder of the colony, as
above indicated. Embryo arising from the egg, known as the
‘‘eyathozooid,’’ 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.
98 University of California Publications. [ ZooLoGy
Genus Pyrosoma Peron.
But one genus in the family, hence generic characters same
as those of the family.
Pyrosoma Peron, 1804, p. 437, Pl. 72, and all succeeding writers.
Pryosoma giganteum Lesueur. Fig. 30.
Je giganteum Lesueur, 1815, p. 70, Pl. I, figs. 1-15.
P. giganteum Savigny, 1816, p. 207, Pl. IV, fig. 7, and Pls. XXII
and XXIII, many figures.
P. giganteum Keferstein und Ehlers, 1861, pp. 72-77, Pl. XII, figs.
4,5, 7, and 8.
P. giganteum Paneeri, 1872, pp. 1-25, Pls. I and II.
P. giganteum Herdman, 1888, pp. 26-29, Pl. I, figs. 4-21.
P. giganteum Seeliger, 1895, pp. 61-62, Pls. I and II; Pl. IV,
figs. 3 and 4. Fig. 30.
Fig. 30.—Pyrosoma giganteum colony, 4.5 em. 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 atrial orifices of the zooids corresponding to
the processes being on the more convex side of the processes.
The processes, as a rule, with an oblique, more or less distinct
plane, with finely serrate edges at their summits. A velum or
diaphragm of test shutting off to a variable extent the opening
of the colony. Measurements of three largest preserved colonies:
first, length 25 em., greatest thickness near open end 2.5 em.;
second, length 25 em., greatest thickness 3.5 em.; third, length
19 em., greatest thickness about middle of length, 3 em.; largest
colonies observed, measured in life, 60 em. long, 40 em. long,
and 35 em. long. Thickness of wall of largest colonies about
6mm. Color varying from an entire absence of pigment and ex-
treme transparency to livid pink, due to pigment in the oozooids.
a a me ee ea ei
VoL. 2] Ritter—The Pelagic Tunicata. 99
Hl W
HUT
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iil
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Fig. 31.—Single zooid 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 sae in full
erown 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
longex.
Largest number of branchial stigmata seen, thirty-two, on
each side of sae; 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. [ ZooLocy
shaped; intestinal loop narrow, the anus 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, after much perplexity, to call our
one species of Pyrosoma, P. giganteum, I must confess that the
decision as between giganteum and atlanticum has little more
value to my mind than it would have had 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 specifie 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 case of the zooids, often within the same colony. In
fact, I am compelled to question the actual existence of both
atlanticum and giganteum as distinet species. For example,
Savigny, 1816, 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 gigantewm ecylin-
drical. This difference appears to have been chiefly relied upon
by Herdman, ’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] Ritter —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
ereater 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 gigantewm. 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
youn colonies agree entirely with Seeliger’s account of the
young giganteum.
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 ean
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 em. long. It is, however, true
that one sees colonies of say 20 mm. length some of which are
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 Mareh on through the summer and autumn months. 1!
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 Californa Publications. [ ZooLocy
BIBLIOGRAPHY.
The more important literature of the Pelagic Tunicata.
Agassiz, A.
1866. Description of Salpa Cabotti Desor. Proce. Boston Soc. Nat.
Hist., Vol. XI, pp. 17-23. Good figures of Salpa democratica-
mucronata.
Apstein, Carl.
1894. Die Thaliacea der Plankton-Expedition. B. Vertheilung der
Salpen.
Ergebnisse der Plankton-Expedition der Humboldt-Stiftung.
Bast Baas:
New: S. floridiana, S. verrucosa, S. magalhanica, S. fusiformis
var. echinata proles gregata. S. virgula redescribed. Special
attention to distribution, geographical, vertical, seasonal.
1901. Salpen, in Nordisches Plankton, p. III 5-IIT 10.
Banks and Solander.
1773. Hawkesworth’s Voyages. Vol. II, pp. 2-3. The deseription of
Dagysa by Hawkesworth, but attributing the observations
and name to Banks and Solander, admits of no question
that the animals described were salps.
Blainville, de H.
1827. Dictionnaire des sciences naturelles, Tome 47, pp. 94-123. Ge-
eric name Cyclosalpa proposed.
Borgert, A.
1894. Die Thaliacea der Plankton-Expedition C. Vertheilung der
Doliolen.
Ergebnisse der Plankten-Expedition. Bd. I, E. a.
1901. Dolioliden, in Nordsches Plankton, p. III 1-IIT 4.
Brooks, W. K.
1875. Embryology of Salpa. Proc. Boston Soc. Nat. Hist., Vol. 18,
p. 198. Author’s view that solitary salpa is the female, and
chain salpa the male, hence not a true alternation of gen-
erations, first set forth.
1893. The Genus Salpa. Memoirs from the Biological Laboratory of
the Johns Hopkins University, Il, Johns Hopkins University
Press, Baltimore, pp. 1-303. Most comprehensive study of
the genus yet published. Part. I.—‘‘A general account of
the life history of Salpa. Part I1.—The systematic affinity
of Salpa in its relation to the conditions of primitive pelagic
life; the phylogeny of the tunicata; and the ancestry of
the Chordata. Part III.—A eritical 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] Ritter —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 deseription 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: Tunieata Lmk. Die Klassen u. Ordnungen des
Thierreichs. Bd. III, 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 eir-
cumnavigatione terrae auspicante Comite U. Romanzoff duce
Ottone de Kotzebue, Annis 1815, 1816, 1817, 1818 peracta.
Fasciculus primus. De Salpa. Berolini. pp. 1-24.
New species Salpa affinis (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-
spiciunter 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. Mémoire sur les Thalides et sur les Biphores. Annales du Mu-
séum 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. Mémoires pour servir a l’histoire et 1’anatomie des Mollusques.
Mémoire sur les Thalides (Thalia. Brown), et sur les Biphores.
(Salpa. Forskaohl), 22 pp., 1 pl. A reprint of the 1804
memoir.
Delage, Yves et Edgard Hérouard.
1898. Trataité de Zodlogie Coneréte. T. VIII, les Procordés. 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. Deseriptiones Animalium Avium, Amphibiorum, Piscium, Insee-
torum, Vermium; quae in itinere orientali observavit.
Salpa, p. 112. Genus instituted and defined as belonging to
the mollusca: S. maxima, S. pinnata, S. democratica, S. mu-
cronata, S. confoederata, S. africana, S. punctata, S. fasciata
S. sipho, S. solitaria, S. polyeratica.
Gegenbaur, C.
1854. Ueber die Entwickelung 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 nebst Bemerkun-
gen tiber die Larven dieser Thiere. Zeitschr. f. wiss. Zodl.,
Bd. VII, p. 283-314.
Recognized the difference between the median and _ lateral
buds on the dorsal process, without, however, finding how
these are related to one another and to the ‘‘nurse’’; and
other important new facts. .
Grobben, Carl.
1882. Doliolhim und sein Generationswechsel, etc. Wien, 1882. One
of the most important contributions to the knowledge of
the genus.
Herdman, W. A.
1882. Report on the Tunicata collected during the Cruise of H.M.S.
Triton in the Summer of 1882. Trans. Roy. Soc. Edinburgh
Vol. XXXII, pp. 93-117.
1888. Report on the Tunicata collected during the Voyage of H.M.S.
Challenger, Part III, Pyrosomidae, Thaliacea, Larvacea.
Challenger Reports, Zodlogy, Vol. XXVII, pp. 1-163, Pls.
1-11. New salpidae: S. musculosa, S. mollis, S. nitida, S.
quadrata, S. echinata. New Doliolidae: D. affine, D. challen-
geri, D. krohni, D. tritonis. New Pyrosomidae: P. spinoswm.
All new and numerous old species fully described and fig-
ured, and relationships discussed.
bf
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 Jasis, Savigny, 1816.
1904. Ascidians and Amphioxus. The Cambridge Natural History,
Vol. VII. Macmillan and Co.
Huxley, T. H.
1851. Observations upon the Anatomy and Physiology of Salpa and
Pyrosoma. Philos. Trans. Roy. Soc. London, part II, pp.
567-593. In addition to the important discussion of the
Vou. 2] Ritter—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. Remarks upon Appendicularia and Doliolum, two genera of
the Tunicata. Ibid., pp. 595-605.
Jones, T. Rupert.
1848. 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. III Ueber die Anatomie und Entwicklung
von Doliolum, pp. 56-71, Pls. IX, X, XI. Important. IV
Bemerkungen tiber die Anatomie von Pyrosoma, pp. 72-77
el pexXaliie
Krohn, A.
1846. Observations sur le génération et le développement 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, Pl. 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. miilleri, D. nordmanni
and D. troschelwi described as new.
Lahille, F.
1890. Contributions a 1’étude anatomique et taxonomique des Tuni-
ciers. Thésés, Tolouse.
Lesson, R. P.
1830. Voyage autour du monde, exécuté per ordre du roi, sur la cor-
vette de S.M. la Coquille, 1822-25. Zodlogie, t. II, Paris,
1830, pp. 256-279.
Lesueur.
1815. Mémoire sur quelques nouvelles espéces d’animaux mollusques
et radiaires receuillis dans la Mediterranée prés de Nice.
Bull. d. Science de la Société Philom., Paris. T. 3, p. 281,
BIR Vee tioee ce:
1815. Mémoire sur l’organisation des Pyrosomes, et sur la place qu’ils
semblent devoir occuper dans une classification naturelle.
Bull. d. Science. de la Société Philom., Paris, 1815, p. 70, Pl.
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 Octacnemus patagoniensis.
Panceri, Paolo.
1872. Gli organi luminosi e la luce dei Pirosomi e delle Foladi. Atti
della R. Acad. della Scienze fisiche e mathematiche de
Napoli, Vol. V, No. 13, p.51, Pls. 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. giganteum.
Peron, M.
1804. Mémoire sur le nouveau genre Pyrosoma. Annals du Muséum
d’hist. natur, T. IV, p. 437, Pl. 72. Pyrosoma atlanticum
described.
Perrier, Edmond.
1899. Traité de Zodlogie. Tasc. V, Amphioxus-Tuniciers.
Quoy et Gaimard.
1834. Voyage de découvertes de 1’Astrolabe. Zodlogie, t. III, pp. 559-
602, Pls. 86-89. Many species of Salpa and Doliolum elab-
orately, but in many instances not accurately, illustrated;
many of the figures not identifiable.
Salensky, W.
Beitrige zur Embryonalentwicklung der Pyrosomen. Zodl.
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-César.
1816. Mémoires sur les Animaux sans Vertébres. 2d partie, Ist fasc.,
p- 124, Pl. 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.
Vou. 2] Ritter —The Pelagic Tunicata. 107
1895. Die Pyrosomen der Plankton-Expedition. Ergebnisse der
Plank.-Exped. der Humboldt-Stiftung, Bd. IT, E. b. Fullest
most resent treatise on the structure and classification and
distribution of the group. New species: P. aherniosum, P.
minimum, and varieties levatum and tuberculosum of P. at-
lanticum.
Todaro, F.
1902. Sopra gli organi escretori delle Salpidi. Rend. Acead. Lincei
(5), Vol. 11, Sem. 1, p. 405-417. In addition to deseription
of renal organ in Salpa, proposes genus Helicosalpa.
Traustedt, M. P. A.
1885. Spolia Atlantica. Bidrag til Kundskab om Salperne. Vidensk.
Selsk. Skr., 6 Raekke, naturvidenskabelig og mathematisk.
Afd. II, 8 p. 339-400. Most useful paper on the classifica-
tion of Salpa yet published.
1893. Die Thaliacea der Plankton-Expedition A. Systematische Bear-
beitung. Ergebuisse der Plankton Expedition, Bd. IT, Weide
Uljanin, Basilius.
1884. Die Arten der Gattung Doliolum im Golfe von Neapel, ete.
Fauna und Flora des Golfes von Neapel, Monographie X.
The most extensive treatment of the genus yet published.
Vogt, C.
1854. Recherches sur les Animaux inférieurs de la Méditerranée.
Second mémoire, Sur les Tuniciers nageans de la Mer de
Nice. Mém. de 1’Institut 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 in the text figures as well as in those
of the plates.
an.—anus.
at. o.—atrial orifice
br.—branchae.
br. o.—branchial orifice.
coe.—Cecum of intestine of sexual
zooid.
el.—eleoblast.
end.—endostyle.
emb.—embryos.
es.— esophagus.
d. p.—dorsal process of nurse.
gn.— ganglion.
gl.—** gill.’?’
ht.—heart.
hy.—hypophysis.
i. c.—Intestinal appendages.
int.—intestine.
nu.—* ‘nucleus. ’’
oc.— otoeyst.
ov.—ovary.
p. a.—post abdomen.
p. br, b.—peripharyngeal band.
pd.—peduncle for attachment of
sexual zooid to stolon.
ped.—peduncle.
pla.—placenta.
s.—stomach.
sto.—stolon.
tes.—testis.
v. p.—ventral process of phorozooid.
[108]
PLATE Il.
All figures of Plates II and IIT are of Cyclosalpa bakeri.
Figs 1, 2, 3.—Lateral, dorsal, and ventral views, respectively, of the
solitary generation, all drawn mainly from preserved specimens.
Fig. 4.—Outline of same, lateral view, made from a living, actively
swimming specimen.
Fig. 5.—Intestinal tract, ‘‘gill,’’ endostyle, peripharyngeal band, hypo-
physis, and ganglion with its sense organ, seen from left side.
Fig. 6.—Anterior end of ‘‘gill,’’ with hypophysis.
{110]
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PLATE III.
Figs. 7 and 8.—Left and right side views, respectively, of zooids of
sexual generation, taken from the stolon.
A :
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[112]
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No. 4.. Basket Designs of the radians of Noppenters California, ~
by A. L. Kroeber. Pages 60, Plates 7, January, 1905. ‘Price, -75 .
Vol. 3. The Morphology of.the ape Language, OY EunY Earle Goddard.
Pages 344, June, 1905. : Price, 3.50 |
ANTHROPOLOGICAL MEMOIRS.
Vol. I, Explorations in Peru, by Max Uhle (in preparation).
No. 1. The Ruins of Moche.
No. 2, Huamachuco, Chincha, Ica. Spe ;
No. 3. The Inca Buildings of the Valley of Pisco.
UNIVERSITY OF CALIFORNIA PUBLICATIONS
ZOOLOGY
Vol. 2, No. 4, pp. 113-233, Figs. 1-62. October 14, 1905
CONTRIBUTIONS FROM THE LABORATORY
OF THE
MARINE BIOLOGICAL ASSOCIATION OF SAN DIEGO.
IV.
THE PELAGIC COPEPODA OF THE
SAN DIEGO REGION.
BY
C. O. ESTERLY.
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
deseribed 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.
Ritter 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. [ ZooLocy
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 mawillac; (5) The ante-
rior maxillipeds (second maxillae); (6) The posterior maxilli-
peds (maxillipeds). The thoracic segments bear the swimming
feet (four pairs), and also a fifth pair of feet which usually are
modified in both sexes, and at times are absent in the female.
The cephalic appendages from 3 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 bears the inner and outer rami (respectively
endopodite and exopodite). Neither ramus is more than three-
jointed, and in many forms the number of joints is reduced by
fusions.
The anterior antennae in the Gymnoplea consist of 25 joints,
but this number may be reduced by fusion. The joints
earry bristles and sense organs, or aesthetasks. The joints in
an appendage, or a part of one, are numbered from the base
to the tip. Among the Gymnoplea 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.
ih
VoL. 2 Esterly— Copepoda of the San Diego Region. 1s
or not. In the Podoplea some of the males have anterior erasp-
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 Gymnoplea 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 Abbé 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 Gymnoplea 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
eases, 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. [ ZooLocy
It may be remarked that little defense is needed for translat-
ing bodily from Giesbrecht, in view of the fact that whatever
could be written concerning any form known to him has been
so well stated that one could not improve upon it.
Following is a list of the species treated in this paper:
GYMNOPLEA. PAGE
Ie Acantig tonsa MDaAm ae. a sis <ieun wuss ted te oles eter ratetevets cireeneaet fete ee 204
Do VALI GCUST UN MUALIES A ESLAGLY, wears erate siete tone: Sete ttetanat tate aiciete tener nea 145
8. Arietellus setosus Giesbrecht 2.2 = earn ces ces Ses male «ems = eye 189
4. Augaptilus longicaudatus Giesbrecht .............---+--++:> 188
5. Calanus finmarchicus Gunner ..... 20-0. cccee eee sree een eeece 125
65 CALAIS OMGCTIS WANA ene ~io's et oeatel es ets ote ate take) eel acest ental) ciate 128
= GOlANUS MNANOT. GICSDEGGING) .'- cia) c ioe etacieuete leis Gia veetaiweatale tee wi tlerorete cs 126
8: OCalanus robustior Giesbrecht <<< <<... <t- sn sierwis eianle © ale wiv vies 129
9: \Calanus: Genwicorms DADA Kiwin vc sieiw tes ola cle © oie ola nie nis see Sreioks 127
10. Candacia bipinnata Giesbrecht ........2..c cece sree eee eeers 195
PU CONALOU CUT UAB 5. o's ce isare ore mish s, cet Nel abel ae 6! oer etn scone ce al a ee 196
Tos Candacia aetmopica Dama cscs < e cieve wie cin © y= mints witere) sata siehels 196
13. Candacia pectimata Brady .......-cccceccecerr sees seeeeres 193
14. Centropages bradyi Wheeler ..........ceec ee ce ce ercecreaces 172
15. Clausocalanus arcuicornis Giesbrecht .............-.-see-0- 142
16. Hucalanus attenitatius Dama sec secs wes el nysce ela o o'n mis eaten wieieie tare 135
17. Eucdlanus crassus Giesbrecht:.... 6.6 ccc cee renee ween 134
18. Bucatanus. elongatus Dama 5. a sscis soe sine Mis eels siecle 131
19. Eucalanus subtenuis Giesbrecht ...........ce cere es eeceecces 135
20: Buchaets acuta Gaesbrocht 5 oca0scscc.ocns us deu sates oo 157
1. Puchacia media Gresbrecht cc ...e <cim vleim © ei ie ein erwin so sia) aniiaite 160
22. Huchaeta spinosa Giesbrecht ....... 220s nee ne eee ces ee ee nee 159
D3 UCU CLIC. GQINOCIIE | GIROSDIGCID i evere vious pore, #1 shia else ee wer el cael re 155
24. Huchirella galeata Giesbrecht ..........-2- seer eeeeceneeenas 155
95. Euchirella messinensis Giesbrecht ....'..........ceceeceteees 151
26. Huchirella pulchra Lubbock ...... 2... ec ce cee ee ee wee eee 153
OT; EUCRANELUL TOS ENOL CoLANIS cacti abe tates inlets te to role Serie olyalls slant ol cctiae eel 152
28. Gaidius pungens Giesbrecht .........+-.2s2-ee esse ee eres eee 146
99. Heterorhabdus clausi Giesbrecht .....- 2... cee e nce ence ean 185
30. Heterorhabdus longicornis Giesbrecht .................+..--- 186
31: Hererorhabdus. panilligen (Glaus) 2 2. <a correc oe ste eens 184
32. Heterorhabdus spinifrons Claus) ~..- 2.2.02 as nee eg eee ere = 182
33. Labidocera trispinosa, Wew species ...............-+-++----= 200
34. Lucwoutia flawicormais Clans) vo) tec keke) ola nte sl cane alee) kel steels fo 180
35. Mecynocera. claust Thompson ......-1--...-...+..-.----..-- 137
36. Mietridia Uueens: BOCCK. cra aces ates icteric etcetera ae aii
37. Metnidia Doecks Giesbrechit cso cic fet hereto ent tenis ts ry eee 178
38. Paracalanus marvus Claus, S2/.teree = ete =e ete eee al 140
39: Phyllopus bidentatus sb rade tere coca selene astern hn nes 191
40. Pleuromamma abdominalis Lubbock 174
Vou.2] Esterly.—Copepoda of the San Diego Region. aly
PAGE
1 LCUrOMaMmMNa <OTGCilise ClAUIBs 55s Pests « sie gore ds- lerele.s.siw/ ele cee 6 aloe 175
42. Pleuromamma giphias Giesbrecht ......a.0..005000.00 00200 176
43. Khincalanus nasutus Giesbrecht .......4..Zi00.ceceeceecees 136
Aae scoleciinma bradyy Giesbrecht) 2... s0cee ses aeeee.. sence se 165
io, Scolectinmmsaande Mubbock wyrcie. oe 40 at dase 6 0h ose id gee 164
AG SColeciihnin Pacifica, Mew SpeCies . ..c<e-d-aaclees acest eases 168
aif scoleciinima persecans Guesbrechty.... << ses... ssoecaccs + ans 166
ARES COLE CULTS (SUIUILUS SEOUL) ase a vz said a aaa e ate scacal acntete ere hd 170
49. Scolecithrix subdentata, new species ...........0.000eceeeees 167
0) Cineeneananeh Tonto Gieslortxlls 5s anccassnecanohonnodudanbece 148
ol Undeuchacta minor Giesbrecht: 2 2.0 sess sens dan Gao be ses 149
PODOPLEA.
De CLYLEMNEStNG, TOStmata Giesbrechueemeretetie ee ess se ee saee 214
Bain COMI ConnGnniis Cueslomeyelar 5 5o00n0cccnascboonunnoanoane 226
Si, (CORMGECNS: CCRUSIUS: IDEMB, . bo nocosuodsnabdodabosouenkeooude 225
Dos MLeTpeE Ault mOns GLESHTeCHE «acy. sels c <njscie 6 oo mtes «eae Sine 212
Bes JMO TORAH (Enel ococdosaoocnesdaooubenscasoae lal
Dio Qa ion Wein Cueslownxet .oooco0ccdagnboeoonenoaas5n bo nena 209
Dore OM NON PU UMnAy CRA MBAIEE, sia. 301s. disc ovat anh 8 ek SS Seal ois So vec 207
Beh, Onencn Gog Gnas CHSIORAO NK Socoovdoondouasouadseanoasooce 216
GU Oncacae minim Giles breech cere reicieecentciinioeists eat set male
Ol, Sapoploirune Cinema) IDMWES 555 c000ccnnceouuanoocrannvendooeus 221
C2 SG NOG, WHOS DATIGY 5 clo omnod ane bdo no dacdooueod sae oGunoe 219
CoE SCD NNUnUNanLOMOe MN ewa SDCCLES Ur fee se cle cla elaeiaicin Acie cacie > 223
His Savanna, WeMinuan Csr 565 o000cano sed condo o0b0G0de 222
GYMNOPLEA.
Gymnoplea (sub-order) Giesbrecht, 1892, p. 41.
Gymnoplea (tribe) Giesbrecht, 1898, p. 7.
I Sub-order.
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 cf
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
KEY TO THE GENERA OF THE GYMNOPLEA.
(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. 44d.......... 19
2 inner ramus; OL Lrstsroot),.o-jOLULOd ot meme hte een iee eee eee 3
2.) Loner ramus ob first’ £006 -2-jOIntediaw. «yejac ners oe eae eee 32
2. inner ramus of first footie -joimbed) fia lod: eee lel ee ee 38
3. A black or brown knob on the first segment of the cephalothorax
oO
in the antero-lateral angle, on the right or left side; figs. 33a,
ERLE eray dio niche a Sa ee yate Rea an ewe) er oe ee eA ts eee *Pleuromamma
This! knob VAbSeMba ary. 5 ke cetietsesite eto eens ers erel o Siete cteke alertness t
. First joint of inner ramus of second foot with proximally curved
hooks ‘on thesinner margins fos SOD. oie eteres vesine «ieee *Metridia
2 Dhis' joint, Jike the rest, bearing a bristles: o/s... -\-11- se © brea 5
. Terminal joint of outer ramus of third and fourth feet with two
spines or thorns on outer margin, and one terminal bristle; fig. ld 6
. Terminal joint as above, but with three spines on outer border;
HPS). LTO LOD 2 coker ec tave sin morte Mayen Acie eee re averted eee 6a
6. Terminal bristle of outer ramus of third and fourth feet with
nroad smooth borders ne: 1d oes cre aise ree eae eter *Calanus
6a. Terminal bristle bearing teeth or spines on outer border; figs.
Oe DIP oxare ot a aha. 4, Bele Sick letgphe tc 1a ale seuel ore, s Lae atel s eitirstia tee ancl seee fea eae f
7. One bristle of left ramus of furca much longer and thicker than
Ghee OLher <LOrcal (DRISleS) 1... 04> wsitorale + erctero cls ane ereryeloe eects 8
feebuncals bristles: SymiMetrCala. ciccs sen 2 crete: wie eee Crete ste mele 9
8. Mandibular blade with three or four teeth, the ventral one
hooked and separated from the others by a wide space; fig.
ERS siesta seotels age cert oe Rieck eve euebcuhe. Sahu SECA ete, Sehiace *Heterorhabdus
8. Mandibular blade with at least 8 teeth..................... Disseta
9! AmtervoraAncenn ge SymMMenLICAl . hs spe sis o/es mm lois,ciets’s = fester stetaaiets 10
9.,Anterior antennae (as ymimenm cally i. chy- see cision wines an sae es 14
LO! Rami of ftth feet 3-jommteds mips S263 seo e ss wisis ees eiclege coeteereis 11
1O:) Rami of fitthy feet. a —jommbe Merwe ss ate eel vol ofc paras eo oot *Augaptilus?
10. Outer ramus 3-jointed, inner ramus 2-jointed............ Isochaeta?
10. Outer ramus 3-jointed, inner ramus 1l-jointed............... Isias?
10. Outer ramus 3-jointed, inner ramus lacking............. *Phyllopus?
10. Outer ramus 1-jointed, inner ramus rudimentary; fig. 42b *Arietellus?
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 vhorder: figs sOU,eoRey ar ee cet peel scce ter acter thee leet tte 12
12. Terminal joint of inner ramus of fifth foot with five bristles;
ik) Sasol Morea Sho arcane xi Gee oc Soo ae 0 *Lucicutia?
12. This joint, withvat, least) sixes bristles: orto ore artnet eterno eer 13
13: Abdomen with 4 seomentsi 7. )icte <tc sete tee tar ener ete Haloptilus?
13. Abdomen’ with! 3) SeSmMents tse cele elelsiaietetet tated a *Augaptilus?
Vou.2] Ksterly—Copepoda of the San Diego Region. 119
(4 Gxraspinrsantennea ony te hte Sid es o> trey ieqie +o lislereiss)aiste)<' <i s.6 15
(4 Graspinc antenna somwletbesider 7s cyeeeseriies saucer sbole suche cle eres eeenets' «)< 7
15. Inner rami of both feet of fifth pair 3-jointed, with plumose
LORSIISIPIGSISe Tica 2 NOOR eleor olen. cin ec HE Oia c ODES Oo DRIOiC Ee Dio. coe bias 16
15. Inner rami rudimentary, without plumose bristles ............ Isiasé
16. Inner rami of feet of fifth pair unlike, the right foot with a
ORCOTSM, sclera tes tole ons arte ee tak wale Giscss atsmerneersteretat terene rane *Centropagess
HG Heimie re reaman yallalkseiseiti og, Gry alld eet yoneteyovetemen steer eet elicate) okaltel *Augaptiluss
Pe Bommramivoteach toog on fib he palces-fOUNTed erp utemiletnake lel: 18
17. Both outer rami 3-jointed, the inner rudimentary........ * Arietellust
17. Rami of left fifth foot 3-jointed, of the right 2-jointed.. *Lucicutiat
18. Inner ramus of maxilla present, distal bristles of anterior max-
iiped makedwor: Set. wathespIMesise-ss ct ct sale ele) ole soicin =i Haloptilusg
18. Inner ramus of maxilla lacking; distal bristles of anterior max-
illiped with mushfoom-shaped appendages (cf. fig. 41D)
Js aD STOIC DOS ORDO OIDIOIRC Oe Oran UID ena CrO CPAs *Augaptiluss
HOMME TAT AIMS Olea ties tetO Opts] OLMLC Cayepeya ass) ete storey eilelio) cierto) sieliela ciara’ 20
WG), Toaiaeie WAMU) OEE Taste sto Bommel Seco oéoeoscna600cn0 cco cboncc 25
1G), IGaiMCie wee aaNOKS) COLE ToISIE stOKONA dleafoMGYCl CounsoacoedoocouabEeoe Eurytemora
20. Head without dorsal cuticular lenses or lateral hooks............. 2]
20. Head with one or two pairs of cuticular lenses and hooks on each
STGIGY, dba oO OIG IOC GEO ENE: Oise es Ot oo notte PRE CHO ROCCE SOO ON eNO 23
21. Rami of posterior antennae about equal in length....... Parapontella
PN. WaaaAeIe TATE TaN, IME WOE Geo soo danccoboasoopDcGGooeUaHaeG 22
22. Abdomen with asymmetrical outgrowths ............... Pontellopsis
22, Aoclomean charmmeriaGel 5 oo5pcesbaoooudgsboodsaooomuCds 4c Pontellina
23. Head with two pairs of eye lenses..........:.......-.- Anomalocera
OS), Wiesel yyairle lohe Chaves jopwle ChE NEMS! G5 ac5ccoccvcouc0 bbe mcononceoor 24
24. Last two thoracic segments distinct; anterior antennae of female
24-jointed; terminal portion of grasping antennae of male
Ee OT ACS cate lane ca son saes one ce eheres emia) «een ial se =) Geka ss aera vaxct™ a rar ate Pontella
24. Last two thoracic segments fused; anterior antennae of female
22-jointed; terminal portion of grasping antennae of male
cleo Ret |g Ine Ein do 4 5 na oie eo On OCI ICI Ivellopsis
25. Head with one pair of cuticular lenses; fig. 48a.......... *Labidocera
25. Head without dorsal cuticular lenses .........----.-++++--+-s0: 26
96. Inner rami of fitth feet jommted....-...-....--4------- *Centropages
26. Inner rami of fifth feet 1-jointed or absent...............-++--- 27
27. Second maxilliped longer than the first ...........-++.++--+4-- 28
27. Second maxilliped shorter than the first ........--++-.+++e+e0-- 29
28. Furea long and narrow, at least six times as long as broad... . Temora
29. First joint of maxillipeds with few short bristles; fig. 47h. .*Candacia
29. This joint bearing long bristles set with spines............+++-- 30
30. Outer ramus of posterior antennae less than half as long as the
TMP. TATE; sovodescocodeponasoopeeneugcacsnasuooducT * Acartia
30. Outer ramus more than half as long as the inner...........+--- 31
31. Posterior maxilliped 6- 7- jointed...........+.++2++--+e-- Calanopia
31. Posterior maxilliped 3-jointed ..........-2.+eeee eee recess Tortanus
39. Both feet of fifth pair with inner.rami, which are 2- or 3- jointed,
bristles. plumose ....... 6. ee eee eet e eect eee nett eens *Lucicutia
120 University of California Publications. | ZooLoGY
32. Inner rami without plumose bristles, or lacking; the entire foot
may beabsent on one or sbobhysides. cer scale eee ioe sien ee 33
33. Furea long and narrow, at least six times as long as broad....Temora
33. Furca at most three times ag lone as broad. 5... <csc~ ec se eae 34
34. Middle joint of outer ramus of third and fourth feet with two
bristles; terminal joint with seven; fig. 12a................. 35
34. Middle joint with one, terminal with five...................:.- 37
35. Outer border of outer ramus of swimming feet not denticulate. .
wo avelie aveievevapsiace'a oy dus sobeletnsal atetapoteka uae uOtsye iets cei nia tey sesso. sieht Calocalanus
35. Outer border on rear pairs denticulate; fig. 12a.......:......... 36
36. Fifth foot lacking in female, or knob-like; in the male only the
left, fifth. oot: present is. /cvecte we ecsielcreisterccs een eee ets a ay Acrocalanus
36. Fifth foot in female 2-jointed (fig. 12c); in the male the right is
2-jointed, the left 5-jointed (fig. 12e)................ *Paracalanus
37. Ourer ramus of first foot 3-jointed; fifth foot absent in female...
rises chs by pane ante Ohara) NEEM tpib tata loll diets (een er eo paRePEE Ts ket ore eNO *Eucalanus
37. Outer ramus of first foot 2-jointed; fifth foot present in female
GH LOD oie teres acho arate tae oss, yagec esa ter se itches ke anteater *Rhincalanus
38. Inner ramus of second foot 3-jomted.....-...-...+6.-- *“Mecynocera
38. Inner ramus of second foot 2-jointed (fie. 280)............<6.-..- 39
38. Inner ramus of second foot l1-jointed (fig. 25€) ................ 51
39. Terminal joint of outer rami of second to fourth feet with five
Pmisules = oninmer an am eu tes apiet oa laxeteelerovel sie tans erere reise net Spinocalanus
39. Terminal joint with four bristles on inner margin (fig. 14))..... 40
40. Surfaces of rami of second to fourth feet without very large
spines; appendages of anterior maxillipeds have the form of
DTISLIGS OLA OOKS i aia retsis ics, ousie Hosieves okt srekonehe ay Otis aeetniet oRcvoha: a aiets hate 41
40. 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. 28b); part of the appendages of the anterior maxilliped
vermitorm (ie. o0d) ‘or -pencillate scr. <itelests ore byes ste reniene 49
41. Basals and outer rami in second and third feet broader than in
the fourth pair, the second basal irregularly toothed on the
FATES (HEE MI) oY oy oto Ue cen Cy Hs aaa FS) Pee a OI ee 8 eI ot Oyo *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
LTS pM NT Aros ck caalst oye seapoucyolanetes simte ewer ein el lela keeueta ayetaris Ctenocalanus
423 ThesevotLethesnsval ishapereatectsiicercae oc real erates Shee a eee eee 43
43. Fifth pair of feet symmetrical 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....
PP Pas PCa EE ey IA ta OTOL pop nolo an og cmowtoct Gaetanus?
Aa. Head =wathout: Ghis spine ge cite seacets)cierar-tol rete ene Ren Meieyetetet esta netaters 45
Ade vith LOOT MACK IAD sper cre: ous ie orate eect! helene ote 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
een A eae Ole GIS OO Oo CO CIOE Drepanopus?
46. Terminal bristles not as long as, or but little longer, than basal
fori Ure IES Roe el atsrars ASiaraccNiS to DoS S.crolo ore Stephus?
Vou.2] Esterly—Copepoda of the San Diego Region. 121
47. Fifth feet, especially the left, with several apical appendages. ..
g Oth Canoe AROS AE OLE Chad Oop OueRD PRCANG ICID OOS ner oe RCo eNO Stephusg
47, Fifth feet slender, stylet-like, with a few short needles or with
but one needle or hook-like appendage. .¢.%..........o20.0-- 48
48. Fifth feet slender, stylet-shaped, about as long as the abdomen...
5, 5 S-otto craic yO RL DIG Die eraraer Nui mis ean o Craukonn iene Pseudocalanus4
48. These shorter than the abdomen, the right foot with a terminal
INGYOLE 5 refage Rec REG eR aeons Mee Oy Geel “Wins ae Seat rp a ee Drepanopust
fo cephalothorax, broad to globular 222.100 aasoosse ase ens esos Phaenna
49. Cephalothorax elongate, ellipsoidal (figs. 26a, 29).............. 50
DO a Ehead distin chy trom, borax, Was coe Lie ete ave kee ele Xanthocalanus
50 tead fuged with thorax’ (figs. 26a,°27)). .< iene 2 ne conan 56 *Scolecithrix
SPP t esr OOts LACKING. Vier. as, se eoaetne A size en aenv ee Mme See eee io apsticc de, as 52
ep L Hae OOt MORESOMEN: S7..ctr.< hiatal etorsiers a Sait st tale carte miele teres senate oes 59
52. Last thoracic segment prolonged laterally into a long, pointed
PROCESS ACH Shea, SU tarsi sleet cecle ete teriatcl kisah aL ev 53
52, Lateral angles of last thoracic segment rounded or slightly pointed
(CaicseeallG cea) eon aetna eracsce ee aed eters Suet mt LS aio 56
53. Bristles of the sixth joint from the last in the anterior antennae
Ehnickwan(distrans merselyzerinoe dimen eies esd crernie seers Bradyidius?
Se}, MUNESS lous, Gh Ta@ Wel WOM. Soo ole oaaoacadenugopcudGusan6 54
o4. Rostrum with two heavy teeth (fig. l4a)............... * Aetideus?
54. Rostrum with one point (fig. 150) or lacking.................. 55
Sas Owiise weweMMIS| Che Tudshy TOON Bsomamiecl FogoonancoucaboouccgDc Chiridius?
DI Oucer TaMUs oO fins oot 2-joimbeds iow W5q)e ea. seh ee *Gaidius?
56. Rami of posterior antennae about equal in length........ *EuchaetaQ
56. Outer ramus at least 1144 times as long as the inner ramus
(GIN OCI aesdevencusyer fay ope ener eters ee te ene Sterere tenes Stayt heiar apes, auelarancatleir 57
57. Inner border of first basal of fourth foot naked or feathered..... 58
57. Inner border with teeth or spines (figs. 19b, 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
bluntyprocessess (mot spimes))i tans... sas ss esse Chirundina?
59. Last thoracic segment on each side with a strong point (fig.
NAC asap tN Pape Maton ee rica ceusctiel sy asc ek dite) ah dito er lepps telcos saver ont clove * Aetidiuss
59 hast ibonacresseament noun ded silos 20 Gees ree ciel is 60
60. Right fifth foot with forceps (figs. 20b, 21b)............ Euchirellag
60. Right fifth foot ends in a stylet without forceps (fig. 23a).... 61
61. Inner rami of posterior antennae at most equal in length to the
OMFS Leeds SenchO Gi GLES.O1 SO OICATE PSEOW Onl REET IRON ONO ed Euchaetag
. Inner rami of posterior antennae over half the length of
Ce MOUECI, acy seve. sSareh syokor Siemens oe aoe tateda ce w ahsnele is aie tousle *Undeuchaetat
122 University of California Publications. | ZooLoGy
Fam. CALANIDAE.
Dana (subfamily), 1852.
Claus, 1868, 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;
that of the male with 5, often with very short anal segment.
Anterior antennae of female, 16- to 25-jointed. Outer ramus of
second antennae one-half to four times 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 5-, 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. 300).
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. 1d),
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
absent.
Sub-fam. CALANINAE.
Calanina Giesbrecht, 1892, 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-
Vou.2] Hsterly— Copepoda of the San Diego Region. 128
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 Giesbrecht, 1892, pp. 45, 88, 725.
Calanus Dahl, 18946, p. 61.
Calanus Dahl, 1898, p. 13.
Calanus Wheeler, 1899, p. 164.
2 Head free, or fused with thorax; fourth and fifth thoracic
segments not fused. Abdomen with 4 segments, genital segment
symmetrical, furea occasionally asymmetrical. Anterior antennae
25-jointed, terminal joints with long, plumose bristles; the anten-
nae vary in length, in some eases not reaching the posterior end
of the body, in others extending beyond the furea (fig. 5a).
Rami of posterior antennae of about equal lengths, outer ramus
7-jointed. Inner ramus of maxilla 3-jointed. Anterior maxil-
hped 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 sealpelliform, its
margin smooth (fig. 1d). 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 lke the others.
4 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
parts 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. 1b, ¢), 2 basals on each side,
right foot (fig. 16) with 3-jointed outer and inner rami, the left
(fig. le) with 3-jointed outer ramus without plumose bristles,
inner ramus reduced or lacking.
KEY TO THE SPECIES.
91. Anterior antennae extend beyond end of cephalothorax for at
least halt ts Hon ebhs (Hep) ele cnis pret sinix eieteriede w sfelole myemye |e ere 2
1. Anterior antennae do not extend beyond end of cephalothorax,
OT OM yet OL a Rew | OLNCA ail MOSUsts x ot orice stats seaside eno te 4
2° BYIstles: OL LULA SyIMMeGriGal \. -yalcigis\ tant asvayetd enews ieee C. tenuicornis
2. Furea with an elongated bristle on left side.................... 3
3. Ventral surface of genital segment very strongly convex (fig. 5c)
at ee EM erentecn, Sphere ae pn sietn oiiiene diet aE ee eee aed enamoche: oteete iets C. robustior
3. Ventral surface of genital segment rather slightly convex (fig.
GT ates cB ety cents cies a oayakal sets sats as hg cna mie oped Rela aoe LenS atatate C. gracilis
4. Head not fused with thorax: cephalothorax with six segments
(Gites) ee Bec ney ROR as CO Ee ea ckaIC cue C. finmarchicus
4, Head fused with thorax: cephalothorax with five segments (fig.
23) Reasae Puc ame Fee Tie URIs CIO teen tac teks otc eke co eee eee C. minor
41. Anterior antennae longer than body by at least six joints..........
PE ey A POS TCI RYO ET rrr ok eel eS aca coe Cc. tenuicornis
1. Anterior antennae not longer than body, or but slightly so....... 2
2. Outer margin of terminal joint of outer ramus of second to
fourth Leet) demi Gulate sot ones eters syeye fener stereos delet ele etslalote 3
2. Outer margin of same smooth or feathered...............-.+---- 4
3. Inner rami of fifth feet similar in structure.............. C. gracilis
3. Inner ramus of left foot of fifth pair shortened and without bristles
ea OC re TC qc. o Spe acne ett ae rae C. robustior
4. Outer ramus of right fifth foot with plumose bristles on inner
isck 1g? 0 i APNE pany terete etek fre or tieh er pau nr C. minor
4. Outer ramus of right fifth foot without bristles on inner margin
(fig. 1b); outer ramus of left foot (fig. le) less than twice as
long as that of the right .............-2--+-20.- C. finmarchicus
Vout.2] Hsterly—Copepoda of the San Diego Region. eS
1. Calanus finmarchicus Gunner.
Monoculus finmarchicus Gunner, 1765, p. 175, figs. 20-30.
Calanus perspicaz Dana, 1852, p. 1071; 1855, pl. 74, figs. la-c.
Cetochilus helgolandicus Claus, 1863, p. 171, pl. 26, figs. 2-9.
Calanus finmarchicus Brady, 1883, p. 32, pl. 1, figs. 1-10.
Calanus finmarchicus Giesbrecht, 1892, pp. 89, 218, pl. 6, fig. 19;
pl. 7, figs. 32, 33; pl. 8 figs: 3, 15, 21, 31, 335; 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 140. (d) Outer ramus of third foot of male
«140. St., terminal bristle. Se., outer marginal bristle.
(e) Basals of fifth foot of female 195.
2 Head not fused with thorax, front and lateral portions of
fifth thoracic segment rounded, fureal 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 witha 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. [| ZooLocy
First basal of fifth pair with coneave dentate inner border
(fie: e).
é Head not fused with thorax, anterior antennae straight ;
outer ramus of right fifth foot without bristles on inner margin
(fig. 1b), terminal bristles sometimes dentate, thorn-like. Basals
and proximal joint of outer ramus of left foot (fig. 1c) elongate,
terminal joint of outer ramus shortened. Outer ramus of right
foot (excluding terminal bristle) reaches 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, while
the entire body of some animals is brightly colored.
Leneth: Both sexes, 2.6-3.1 mm.
Oceurrence: Probably the commonest species in the San
Diego region, occurring abundantly in nearly all collections with
the larger nets; in some eases C. finmarchicus is almost the only
species, and is very often predominant.
2. Calanus minor Claus.
Cetochilus minor Claus, 1863, p. 172, pl. 26, figs. 1-8.
Calanus valgus Brady, 1883, p. 33, pl. 3, figs. 1-7.
Calanus minor Giesbrecht, 1892, p. 90, pl. 6, figs. 3, 16, 22; pl. 7,
figs. 6-22; pl. 8, figs. 1, 9, 19, 30; 1898, p. 15.
Calanus minor Wheeler, 1899, p. 165, fig. 2.
Fig. 2.—Calanus minor. Female, lateral, X31.
° 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.
Vou.2] Ksterly—Copepoda of the San Diego Region. 127
4, 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.
Leneth: Female, 1.8-2 mm.; male shghtly smaller.
Occurrence: Not at all abundant, but coming in most catches
with C. finmarchicus.
38. Calanus tenuicornis Dana.
Calanus tenuicornis Dana, 1849, p. 278; 1852, p. 1069; 1855, pl.
73, figs. 10a, 10D.
Calanus tenuicornis Giesbrecht, 1892, pp. 90, 129, pl. 6, figs. 12,
HS plat, fes..0, LO 23s ple 8) esas s27 3 SO8tsn. 18:
a
b
Fig. 8.—Calanus tenuicornis. (a) Male, lateral, X31. (b) 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 114 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.
4 Anterior antennae as in Q, 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. [ ZooLoey
Length: Female, 1.8-2 mm.; male, 1.5-1.8 mm.
Occurrence: Fairly abundant, both in summer and winter
collections.
4. Calanus gracilis Dana.
Calanus gracilis Dana, 1849, p. 278; 1852, p. 1078; 1855, pl.
74, fig. 10.
Cetochilus longiremis Claus, 1863, p. 171, pl. 26, fig. 1.
Calanus gracilis Brady, 1883, p. 35, pl. 5, figs. 1-6; pl. 6, fig. 10.
Calanus gracilis Giesbrecht, 1892, pp. 90, 128; pl. 6, fig. 1; pl. 7,
fig. 26; pl. 8, figs. 2, 4, 6-8, 12, 16, 26; 1898, p. 17.
ene tore
Fig. 4.—Calanus gracilis. (a) Genital segment of female, lateral, x83.
(6) Outer margin of first basal of anterior maxilliped of
female 140.
2 Head fused with thorax; forehead and sides of last thoracic
segment rounded. Left side of furca with one elongated bristle.
Anterior antennae at least 114 times as long as the body. There
is a process at the base of the inner marginal bristle of the second
basal of the second foot (ef. fig. 5d). The proximal outer marginal
spine of terminal joint of outer ramus in second to fourth feet,
respectively, divides the margin into portions as 1:1, 4:3, 4:3 in
length. First basal of fifth pair with feathered inner margin.
4&4 Head separate from thorax, anterior antennae straight,
joints 1 and 2, 24 and 25 fused. Mouth parts greatly reduced.
Outer border of third joint of outer ramus denticulate in second
to 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.
Vou.2] Esterly—Copepoda of the San Diego Region. 129
5. Calanus robustior Giesbrecht.
Calanus robustior Giesbrecht, 1888, p. 332; 1892, pp. 91, 129;
pl. 7, figs. 15, 19, 25, 30; pl. 8, fig. 34; 1898, p. 18.
Calanus comptus Scott, T., 1893, p. 26, pl. 5, figs. 46-50; pl. 6,
figs. 1-5.
b
b2. Sj rid.
Fig. 5.—Calanus robustior. (a) Female, lateral, X36. (b) Outer mar-
gin of first basal of anterior maxilliped of female >< 140.
(c) Genital segment of female, lateral, X83. (d) Inner
ramus of second foot of female 185. B.2, second basal
of foot. Ri.1, 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 (ef. figs. 4a and 5c),
and the first basal of the anterior maxilliped has a bulging
protrusion on the outer border (ef. figs. 4b and 5d).
4 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. EUCALANINAE.
Eucalanina Giesbrecht, 1892, p. 45.
2 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; furea 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 outer rami with smooth edge, that of the first pair as in the
succeeding three pairs; fifth pair absent or uniramous; if present,
with from three to five joints.
4 Body, especially the head, shortened (fig. 6c); anterior
antennae without reduction in number of joints; furea as in the
female. The mouth parts may be stunted. Fifth pair of feet
not well developed, left foot uni- or biramous, right uniramous
or lacking.
1. Genus Eucalanus Dana.
Calanus Dana, 1848, p. 11; 1849, p. 278.
Eucalanus Dana, 1852, p. 1047.
Eucalanus (in part) Lubbock, 1856, p. 13; 1860, p. 160.
Calanella Claus, 1863, p. 174; not Eucalanus Claus, 1881, p. 325.
Eucalanus Brady, 1883, p. 37.
Eucalanus Giesbrecht, 1888, p. 333; 1892, pp. 46, 131, 739; 1895,
p. 246; 1898, p. 19.
Eucalanus Wheeler, 1899, p. 166.
Anal segment and furea fused, latter 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 8- jointed and shorter than inner. Mandible of female longer
than maxilla; second basal of mandible makes with the outer
ramus a eylindrieal body on which the inner ramus articulates
proximally to the outer ramus (fig. 7¢). 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)
Vou.2] Ksterly—Copepoda of the San Diego Region. Bul
both are uniramous. The left 4-jointed, the right 1- to 4- jomted
or lacking. Head appendages of male retrograded and modified,
body shortened.
KEY TO SPECIES.
AN GON Will 8) OR Zh TSGUNTMIS. coco cobs asadococens0e000g500e g
AN KOON Yanda (3) CeAMIEMISs bo a ooe conc cor oud ecooUnBU OD REDO ODOC 4
91. Two segments between anal and genital segments (fig. 6a)......
Jo sb ow eco dub noamomtinn Cmome bod Gd 6 ning. Go-cto molec E. elongatus
1. One segment between genital and anal segments (fig. 7b)....... 2
2. Inner border of second basal of mandible divided into two
approximately equal portions by the insertion of the inner
rep (Cie WO) soasasaccdescosouogeocecHodqon yon E. attenuatus
Ze eroximalsporiion: much) lono er thangs taller pottetspeneltalcieleNelensl-i-r1 3
3. Two terminal bristles of left side of furca longer but hardly
thicker than on the right side; genital segment (fig. 8d) much
broader hans lone oniton-shia Peds eis itt ello sell lelelct en 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,
TROMUACNG! I IMO, Gosoopoaueade mss ugecgooSonoDoUDbE E. subtenuis
$1. Both feet of fifth pair present ..........-.- 2. sees eee e ee eee eee 2
Ie Rireht foot of dikth pair aWSeMba ca. <a oc we lem seems lene mea 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
IMAG 5 ocogdousscooesas boobn Ub Gobo Um UOGooD ODOR Gar EB. 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 clongatus Dana, 1848, p. 18; 1849, p. 278; 1852, p. 1079;
1855, pl. 75, figs. la, b.
Eucalanus elongatus Giesbrecht, 1892, pp. 131, 149, pl. 11, figs.
2, 7, 12, 20, 25, 32, 36; 1895, p. 246; 1898, p. 20.
2 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. See-
132 University of California Publications. — [ZooLocy
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.—Eucalanus elongatus. (a) Female, dorsal, X15. (b) Female,
lateral, X15. (¢) Head of male, dorsal, X37. (d) Fifth
foot of male, X40; left foot at left of figure.
4 Pronounced secondary sexual characters; right fifth foot
present, left as long as the fourth foot exclusive of terminal
bristle.
Coloration: Very transparent, with a small though varying
amount of red in the body, usually limited to a single oil-drop
near the posterior end 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 7144 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.
Vou.2] Hsterly—Copepoda of the San Diego Region. 133
2. Eucalanus attenuatus Dana.
Eucalanus attenuatus Dana, 1848, p. 18; 1849, p. 278; 1852, p.
1080; 1855, pl. 75, figs. 2a-e.
Calanus mirabilis Lubbock, 1856, p. 16, pl. 5, figs. 1-6.
Calanella mediterranea Claus, 1863, p. 176, pl. 28, figs. 6-11.
Eucalanus attenuatus Giesbrecht, 1892, pp. 131, 150, pl. 3, fig. 1;
pl. 11, figs. 1, 11, 13, 16, 18, 24, 40; pl. 35, figs. 3, 6, 17,
25, 34, 37; 1898, p. 20.
fta.seg. ri.
Fig. 7.—Eucalanus attenuatus. (a) Head of female, dorsal, X20. (0)
Abdomen of female, X31. Gen. seg., genital segment. F.-+-a.
seg., furca and anal segment. (¢) Mandibular rami, female,
x31. Ri., inner ramus.
Q 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 114
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 EZ. elongatus; first joint of inner ramus of posterior
maxilliped with 3 bristles, second with four.
4 Pronounced secondary sexual characters; right fifth foot
present, left considerably shorter than the fourth foot.
Coloration : Similar to that of HL. elongatus; 1 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
mm.
Occurrence: A few come in the hauls with elongatus, but are
not nearly so common. They were especially abundant in June
and July, 1903.
134 University of California Publications. [ ZOOLOGY
3. Eucalanus crassus Giesbrecht.
Eucalanus crassus Giesbrecht, 1888, p. 333; 1892, pp. 132, 151;
pl. 11, figs. 8, 10, 17, 21, 22, 38; pl. 35, figs. 4, 20, 26-28;
1898, p. 22.
Fig. 8.—Eucalanus crassus. (a) Fourth foot of female X83. (b) Head
of female, lateral, X18. (c) Head of female, dorsal, X18.
(d) Abdomen of female, ventral, 83.
2 Genital segment (fig. 8d) much broader than long, onion-
shaped ; between it and anal segment but one free segment. Fore-
head (fig. 8c) flatly rounded, furea 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 and 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-
men.
Length: Female, 3 mm.
Occurrence: San Diego, June 16, 1904, one female.
Vou.2] Hsterly—Copepoda of the San Diego Region. 135
4. Eucalanus subtenuis Giesbrecht.
Eucalanus subtenuis Giesbrecht, 1888, p. 3383; 1892, pp. 132, 150,
pl. 11, figs. 4, 23, 42; pl. 35, figs. 9-11, 18, 29, 30; 1898,
15 ale
Fig. 9.—Eucalanus subtenuis. (a) Head of female, dorsal, X20. (b)
Head of female, lateral, x 20.
2 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.
4 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.
Rhincalanus Dana, 1852, p. 1082; 1855, pl. 76, figs. 2a-d.
Rhincalanus Brady, 1883, p. 40.
Rhincalanus Giesbrecht, 1888, p. 334; 1892, pp. 47, 152, 761;
1898, p. 22.
Rhincalanus Scott, T., 1893, p. 30.
2 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-
136 University of California Publications. [ ZooLocy
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 at end of third joint, plumose on inner
border.
4 Anterior antennae shortened; fifth foot on the left side
2-jointed.
Left. fifth foot Diramous oc. a: ass ovav uaee se ee 4
Both feet ot itthpair gmiramous, 2.12 cs. eee 2
1. Rhinealanus nasutus Giesbrecht.
Rhincalanus nasutus Giesbrecht, 1888, p. 334; 1892, pp. 152, 160;
pl. 3, fig. 6; pl. 9, figs..6, 14; pl. 12, figs. 9-12, 14, 16, 17;
pl. 35, figs. 46, 47, 49; 1898, p. 22.
|
Fig. 10.—Rhincalanus nasutus. (a) Female, dorsal, X18. (b) Fifth foot
of female X 260.
2 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. 100.)
4 Right fifth foot with strongly curved bristle at end; left
with long outer ramus reaching almost to end of inner ramus.
Vou.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.
Leneth: Female, 3 mm.; male, 2.7 mm.
Oceurrence: A female was first seen, San Diego, June 10,
1904, but two or three occur in most catches when Hucalanus is
abundant.
>
3. Genus Mecynocera I. C. Thompson.
Leptocalanus Giesbrecht, 1888, p. 334.
Mecynocera Thompson, I. C., 1888a, p. 150.
Mecynocera, Giesbrecht, 1892, p. 160; 1898, p. 23.
Mecynocera Wheeler, 1899, p. 167.
Furea 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. First pair of feet with outer ramus of three
joints, inner of one joint; fifth pair present, with five joints on
either side. & Unknown.
2 Head distinct from thorax; rostral threads delicate ; abdo-
men short, with three segments; genital segment and furea 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
‘amus. 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. C., 1888a, p. 150, pl. 11, figs. 1-4.
Leptocalanus filiformis Giesbrecht, 1888, p. 334
Mecynocera clausii Giesbrecht, 1892, p. 160, pl. 5, fig. 1; pl. 11,
figs. 43, 45; pl. 35, figs. 21, 22; M. clausi, 1898, p. 23.
Mecynocera clausii Wheeler, 1899, p. 167, fig. 5.
The only species of the genus.
Coloration: Exceedingly transparent, without pigment in my
specimens.
138 University of California Publications. [ ZooLocy
Length: Female, 0.9-1 mm.
Occurrence: The only specimens I have were collected Deeem-
ber 30, 1903, on the ‘‘Banks’’ off Point Loma.
NS
<—
a=
f
i
jaa
[)
io \
]
/
1
a |
||
y,
J
\
A
(ee
\
(ease
Pp | Srema
Fig. 11.—Mecynocera clausi. Female, dorsal, x 45.
Sub-fam. PARACALANINAE.
Paracalanina Giesbrecht, 1892, p. 48.
2 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 mner
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.
Vou.2] Esterly—Copepoda of the San Diego Region. 139
4 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 Claus, 1868, p. 172.
Paracalanus Boeck, 1864, p. 8.
Paracalanus Claus, 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; sealpelliform 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. 120) with 4 seg-
ments; the last joint of anterior antennae less than 114 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).
2 Head fused with first thoracic segment, and fourth thoracic
segment with fifth. Rostrum produced into two thin filaments.
Genital segment and furea 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 joimt 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.
4 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.
140 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 Claus.
Calanus parvus Claus, 1863, p. 173, pl. 26, figs. 10-14; pl. 27, figs.
1-4.
Paracalanus parvus Claus, 1881, p. 327, pl. 3, figs. 1-16.
Paracalanus parvus Bourne, 1889, p. 145, pl. 11, figs. 1-3.
Paracalanus parvus Giesbrecht, 1892, pp. 164, 170; pl. 1, fig. 5;
pl. 6, figs. 28-30; pl. 9, figs. 5-11, 25, 27, 31, 32; 1898, p. 2a.
ps.
Fig. 12.—Paracalanus parvus. (a) Fourth foot of female 195. fKe.3,
third joint of outer ramus. (b) Female, dorsal, X83. (c)
Fifth foot of female 410. (d) Second joint of inner
ramus of second foot of female X195. (e) Fifth foot of
male. Ps., left foot. Pd., right foot.
2 Inner bristle of furea barely longer than the furea. Ante-
rior antennae reach, when brought to the sides of the body, per-
haps to the posterior border of the third abdominal segment.
,
4
,
:
:
‘
Vou.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.
4 Fifth foot asymmetrical (fig. 12e); compare also generie¢
description.
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.
@ 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, furea
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.
& Unknown in Spinocalanus and Ctenocalanus. Abdomen
with shortened anal segment; anterior antennae and head
appendages in some eases like those of the female, in others as in
the Paracalaninac. Fifth pair of feet: the right, 1- to 5-, the left
5- jointed.
142 University of California Publications. [ ZOOLOGY
1. Genus Clausocalanus Giesbrecht.
Calanus Dana, 1849, p. 278; 1852, p. 1047.
Calanus Claus, 18638, p. 172.
Eucalanus Claus, 1881, p. 325.
Drepanopus (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
reduced.
2 Head fused with thorax and the fourth with the fifth
thoracic segment. Abdomen with four segments, genital segment
and furca symmetrical. Anterior antennae extend beyond the
thorax, 23-jointed. Outer ramus of posterior antennae 114 times
as long as the inner, the former 6-jointed, with short bristles on
the proximal joints. First joint of inner ramus of mandible
with a very small, sack-like appendage. Maxilla and maxillipeds
as in Calanus, outer marginal bristle lacking on anterior maxilli-
peds. Outer rami of swimming feet with 3 joints; inner ramus
of first 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 uniramous, 3-jointed.
4 Head fused with first thoracic segment, and elongated at
expense of free thorax rings. Rostrum suppressed ; abdomen with
five segments, anal very short. Anterior antennae with joints
8-10, 13-16, 20-21, 24-25 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, uniramous and with 5 joints, right short, 1- to 3-jointed.
1. Clausocalanus arcuicornis Dana.
Calanus arcuicornis Dana, 1849, p. 278; 1852, p. 1056; 1855,
pl. 72, fig. 9a-b.
Calanus mastigophorus Claus, 1863, p. 173, pl. 27, figs. 5-8.
Vou.2] Esterly.—Copepoda of the San Diego Region. 143
Clausocalanus arcuicornis Giesbrecht, 1888, p. 334; 1892, pp. 186,
1985 pl ios 1a pig; fig 7; pl.) 10; figs: 3-8, 14, 16, 17,
19; pl. 36, figs. 29-31, 34; 1898, p. 27.
Clausocalanus arcuicornis Wheeler, 1899, p. 171, fig. 9.
b
Ee
: 2)
Fig. 13.—Clausocalanus areuicornis. (a) Male, lateral, X45. (b) Second
basal of second foot to show toothed distal margin, 410.
(c) Fifth foot of male X83.
2 Genital segment longer than the two following. Furea
about as long as broad. No aesthetask on fourth, sixth, eighth,
eighteenth or twenty-second joints of the anterior antennae.
& 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. 138c).
Coloration: Not very transparent, with red pigment in vari-
ous places on the posterior part of the body and on genital seg-
ment.
Occurrence: San Diego, June 25, 1904, one male.
Sub-fam. AETIDIINAE.
Aetidiina Giesbrecht, 1892, p. 52.
Aetidiinae Wolfenden, 1903, p. 263.
2 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 chitinized, usually with
one point, seldom with two or lacking. Genital segment and
furea 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 at most 24 as long as the second
basal. Inner ramus of first swimming feet always 1-jointed; that
of the second almost always 1-jointed, while in the second and
third the inner ramus is 3-jointed; the form of the swimming
feet as in the Clausocalaninae; inner marginal bristle of first
basal long and plumose.
4 Known in Aetideus, Euchirella and Undeuchaeta. Charac-
ters like those of Clausocalanus; oceasionally the twentieth
and twenty-first joints of one of the anterior antennae are fused.
Left foot of fifth pair 5-jointed (if the right is lacking, or stylet-
like, in which ease the right is claw-like).
1. Genus Aetideus Brady.
Aetidius Brady, 1883, p. 75.
Aetidius Thompson, 1888b, p. 142.
Aetidius Giesbrecht, 1892, pp. 53, 213.
Aetideus Wolfenden, 1903, p. 266; 1904, p. 116.
Aetideus Giesbrecht, 1898, p. 31.
2 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.
———E————— Oe
Vou.2] Esterly—Copepoda of the San Diego Region. 145
4 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, pl. 10, figs. 5-16.
Aetidius armatus Giesbrecht, 1892, p. 213, pl. 2, fig. 6; pl. 14,
figs. 1-13; pl. 36, figs. 6-9.
Aetideus armatus Giesbrecht, 1898, p. 31.
Aetideus armatus Wolfenden, 1903, p. 266.
Fig. 14.—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.
Leneth: 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, pl.
iO), eS 1G tsk
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
3-jointed. |
146 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. 15d).
1. Gaidius pungens Giesbrecht.
Gaidius pungens Giesbrecht, 1895, p. 249, pl. 1, figs. 1-4; 1898,
p- 32.
Gaidius pungens Wolfenden, 1903, p. 266.
‘
Fig. 15.—Gaidius pungens. Female. (a) Dorsal, X18. (b) Head, lateral,
x45. (ce) Tube-like processes on inner distal portion of
second basal of fourth foot 195. (d) First foot «195.
Ri., inner ramus.
2 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-hke (fig. 15c).
& Unknown.
Coloration: Transparent, with little or no pigment.
Length: Female, 3 to 3.5 mm.
Vou.2] Esterly—Copepoda 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, pls. 15, 16.
Undeuchaeta Wolfenden, 1903, p. 267.
2 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-joimted, inner
ramus l-jointed. Outer ramus of posterior antennae at least 114
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.
4 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 34
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. 16f) 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. 16f. Re. 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 Huchaeta, 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 Fuchirella, as does the division
(though indistinct) 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 deseribed 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 Giesbrecht’s the number of
joints is 23. In his description of the male, Sars gives the num-
ber of joints of the anterior antennae as 22, but in his drawing
(pl. 16, fig. 2) there are but 21. The fifth pair of feet in the
male of U. spectabilis is very different from that in the San
Diego specimens, a striking point being that both the right and
left feet 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, pl. 21, figs. 5-11.
Undeuchaeta major Giesbrecht, 1888, p. 336; 1892, pp. 227, 232,
pl. 37, figs. 56, 57, 59; 1898, p. 34.
2 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).
& (new) Compare generic description.
Coloration: Female not especially transparent, with red pig-
ment on basals of posterior maxillipeds, and in mouth region.
Male: plumose bristles of furea steel-blue; those of posterior
antennae and mandible red; on the feet the bristles are faintly
red on the outer ends.
‘— }. 2 ee a ee ees
Vot.2] EKsterly—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
Loma.
reg on
Re eres
d e c.
Fig. 16.—Undeuchaeta major. (a) Female, lateral, X20. Ab.1, 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 195. (f) Distal portion
of left fifth foot of male. Re.2, Re.3, respective joints of
outer ramus. 5
2. Undeuchaeta minor Giesbrecht.
Undeuchaeta minor Giesbrecht, 1888, p. 335; 1892, pp. 228, 232,
pl. 14, figs. 31-34; pl. 37, figs. 55, 58; 1898, p. 34.
OTe.
S
\\\ 2 . . <
a
Fig. 17.—Undeuchaeta minor. Female. (a) Lateral, x20. (b) Second
foot X83.
150 University of California Publications. | ZooLogy
2 Head without crest (fig. 17a), genital segment with a spine
on the dorsal surface.
Coloration: Similar to that of U. 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 part) Lubbock, 1856, p. 21.
Calanus (in part) Lubbock, 1856, p. 15.
Undina Claus 1863, p. 186.
Euchaeta (in part) Brady, 1883, p. 59.
Euchirella Giesbrecht, 1888, p. 336; 1892, pp. 54, 233, 743; 1898,
p- 34.
Euchirella Cleve, 1900, p. 4.
Euchirella Wolfenden, 1903, p. 267.
Rostrum present in most species, simple; lateral angles of
last thoracic segment not pointed. Inner ramus of posterior
antennae 14 to 14 as long as the outer ramus, the two proximal
‘ joints of which are fused. Inner and outer rami of the maxilla
short, the former provided with heavy hooked bristles. Outer
ramus of the first pair of feet 2-jointed, that of the second to
fourth pairs 3-jointed. Inner ramus of 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 (ef. figs. 18d, 19d).
2 Head not always distinct from thorax, last two thoracic
segments fused., Abdomen with four segments, genital segment
and fureal bristles symmetrical or 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
jong 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.
& Head oceasionally 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;
Vou.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.
KEY TO SPECIES.
Brisa] ihe SE CUO) Pget2 C1S( 8) eet tase RE Red, Cates ae ee ye ee ea io)
BE UEt Mab OGRE EESCIIE. 55 'crctae A rales cha dirceaeta cme rermeiies am Na cast wares aus 4
Q1. Head without crest, and rostrum one-pointed (fig. 19a) ........ 2
[Head wath erest, rostrum present (figs 200)06 $4.06 vs. 0secks ooo a
2. Outer ramus of posterior antenna about twice as long as inner..
POCAIDToOOoGoOCGoDonObDoOOOooCoO DTU o OOO OOo DUM COO OC Haga nS 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.
HSE) Spire ny so RSRee PS cc sroy he cts enc ec OAR GNS SoS] gears Mtaton fl ohare es crer ance E. messinensis
a Heaiewat he low erest (fe 20M) sa. ca catia ers sree) saa eee a, E. pulchra
a Eleagewith iioh Cresta (Ho. 226)... vtec eo oe8.s Cccass ss he ene E. galeata
@ JAG aA NONy Cressy (Ge PASO) Geis Gwomue bon GomoG oes onoood Gone 2
Meade wabhOut seresben(He: ZUG) jf. 2 sr. sis: oj eieiord-csceve: srnlelenet sue tere jacsichals 8 ats 3
2. Forceps-like terminal portion of right fifth foot longer than the
JORISEUL joronron@in, (Galery ISA) s Gagaccncoucace Ruste cet ekareeMenene E. messinensis
2. Terminal portion (forceps) of right fifth foot shorter than the
ORAL TNOMaON, Ciey HUM), aeiocs Secoonadaudoo bongo otas E. pulchra
3. Fifth foot short, the right about four times as long as the second
basalts: broad Chios 20) oo ysncla erstcetsl arses cistla dita ehs vec. ee & E. amoena
3. Right fifth foot six times as long as the second basal joint. .E. rostrata
1. Euchirella messinensis Claus.
Undina messinensis Claus, 1863, p. 187, pl. 31, figs. 8-18.
Euchirella messinensis Giesbrecht, 1892, pp. 232, 244; pl. 15, figs.
12, 16, 21, 24; pl. 36, figs. 14, 15, 18, 24, 25; 1898, p. 35.
2 Forehead with rostrum, without crest (fig. 18a); genital
segment asymmetrical, with sac-like appendage on left side of
dorsal surface (fig. 18) ; third terminal bristle on right side of
furea 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.
4 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.
Leneth: Female, 4.5 mm.; male, 4 mm.
Le University of California Publications. [ ZOOLOGY
Occurrence: July 9, 1903, one male; July 22, 1903, one
female.
Fig. 18.—Euchirella messinensis. (a) Head of female, lateral, X15. (b)
Abdomen of female X15. (c) Head of male X30. (d)
Fifth foot of male X20.
2, Euchirella rostrata Claus.
Undina rostrata Claus, 1866, p. 11, pl. 1, fig. 2.
Euchaeta hessei Brady, 1883, p. 63, pl. 20, figs. 1-13; pl. 23, figs.
11-14.
Euchirella rostrata Giesbrecht, 1892, pp. 233, 245, pl. 15, figs. 3,
13, 25; pl. 36, figs. 19, 20; 1898, p. 36.
Euchirella rostrata Cleve, 1900, p. 4, pl. 2, figs. 1-12.
2 Front without crest, with rostrum, abdomen symmetrical.
Inner ramus of posterior antennae 14 as long as outer ramus;
second joint of inner ramus with 8-6 bristles. First basal of
fourth pair of feet (fig. 195; 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 at least to the point of
the first bristle on the outer border of the third joint of the
ramus.
4 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 14 as long as outer. First basal of fourth foot
without triangular lamellae. (Cleve 1900).
Vou.2] Hsterly—Copepoda of the San Diego Region. F533
The male was described by Cleve, 1900, and is identical with
Euchaeta hesser Brady.
Coloration: Red pigment as in EF. messinensis, but more abun-
dant, especially on swimming feet.
= Bil
Fig. 19.—Euchirella rostrata. (a) Female, lateral, X18. (b) Fourth
foot, female, X83. B.1, first basal, showing lamellar pro-
cesses.
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, pl. 4, figs. 5-8; pl. 7, fig. 6.
Calanus latus 2 Lubbock, 1856, p. 15, pl. 2, fig. 12; pl. 11, figs.
8-11.
154 University of California Publications. [ ZOOLOGY
Euchaeta pulchra Brady, 1883, p. 638, pl. 14, fig. 7; pl. 20, figs.
ilbyealye ale
Euchirella pulchra Giesbrecht, 1892, pp. 233, 244, pl. 15, figs. 22,
23, pl. 36; figs. 13, 27; 1898, p. 36.
d
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, 83. i., inner ramus.
(d) First basal of fourth foot of female 140.
2 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 24 as long as outer ramus; second joint of
inner ramus with 6-5 bristles. First basal of fourth pair of feet
(fig. 20d) 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.
Vot.2] Hsterly.—Copepoda of the San Diego Region. fre
On
OV
& Considerably like EF. 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 H. 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;
pl. 15, fig. 20; 1898, p. 36.
Fig. 21.—Euchirella amoena. Male. (a) Lateral X20. (b) Fifth foot
x 45.
Q Unknown.
& 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; pl.
15, fig. 18; pl. 36, figs. 22, 26; 1898, p. 36.
2 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 25 as
156 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.
& 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.
a
Fig. 22.—Euchirella galeata. Female. (a) Lateral X9. (6b) First basal
of fourth foot X83.
Sub-fam. EUCHAETINAE.
~
Euchaetina Giesbrecht, 1892, p. 55.
2 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 feet 2-jointed, of the
second to fourth 3-jointed; inner ramus of first and second pairs
1-jointed, of the third and fourth pairs 2-jointed.
3 Abdomen as in the Clausocalaninae. Outer ramus of first
pair of feet 3-jointed; fifth foot on each side with 2-jointed basal,
and biramous; inner ramus of left stylet-like, of right truncate;
left outer ramus 3-jointed, right 2-jointed.
1. Genus Euchaeta Philippi.
Euchaeta Philippi, 1843, p. 54, pl. 4, fig. 5.
Euchirus Dana, 1846, p. 183.
Euchaeta Dana, 1848, p. 20; 1849, p. 279; 1852, p. 1084.
Euchaeta Claus, 1863, p. 163.
Euchaeta Giesbrecht, 1892, pp. 55, 245, 740; 1895, p. 251; 1898,
Dass
2 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
4
(od)
Vou.2] Hsterly—Copepoda of the San Diego Region. 1
of posterior antennae about 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.
4 Head fused with thorax; abdomen with five segments, ana!
segment short; innermost bristle of furea 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.
23a).
KEY TO SPECIES.
J Earlre(ol ne SOO RMEET OFS (CNN acto oer Cre MeC LG Ole o Abin ae Sie Gi OCIS AiO ean fe)
JRA EGINE MHOYONE. OREN Ne ers-oetaln eae So ooo oid a ebitoe od colds ie OImeini eco 4
41. Terminal joint at each foot of fifth pair, with long straight or
slightly curved stylet; elevation for frontal organ not pro-
TIRUNGHB IED” 5% Gi oa ed cele haw BU CIO oo GOO OOmorn SLAG pion oro bane E. acuta
91. Hairs of frontal organ on a low elevation (fig. 25b)............. 2
1. Hairs of frontal organ on an elevation which extends toward the
fer OMityp (UO LAD) cre tataye srevekete cick sie leks iene NsFo: eres oie 6] 4) let Cxaiyn siaeve ts o/s 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
Giles; BH) scpesgcanoodotonssaadgpqoornonebootegueseucnaune 3
3. Genital segment with a knob-like protuberance in front on the
IGE BIC Gee cdcos at docls pbodsoD oud Gonos Con ocomopnednaT BE. 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
GReey asin (CUE MEU KO coe ec clan CIpIAeicO On Olceced Ok minInImicacieRcrO e Cria 5
SAnterionantennaesloneer tham boOdiye sc aces see E. spinosa
Or
1. Euchaeta acuta Giesbrecht.
Euchaeta acuta Giesbrecht, 1892, pp. 246, 262, pl. 16, figs. 6, 10,
14, 18, 21, 27, 39 ;pl. 37, figs. 47, 48, 52; 1898, p. 38.
2 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. Furea
158 University of California Publications. | ZooLoGy
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.—Euchaeta acuta. (a) Fifth foot of male X37. Ri. dz., inner
ramus of right foot. Re. 3, dx., third joint of outer ramus.
Re. 1, 2, sn., first and second joints of outer ramus of left
foot. Proc., process. Sph., spermatophore. (b) Second-and
third joints of outer ramus of left fifth foot of male «140.
Parts as in a. (c) Second foot of male X60. (d) Maxilla
of male X60. B.1, first basal. B.2, second basal. Le.1, first
lobe of outer margin. fi., inner ramus. FRe., outer ramus.
First lobe of outer border of maxilla (cf. fig. 25f) with six
bristles( one very small), second basal with three, fused second
and third joints of inner ramus with four. Outer border of
Vou.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, pl. 16, figs. 12,
26, 34,47; pl. 37, figs. 31, 34, 35, 50; 1898, p. 39.
2 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. 245). Second terminal bristle
of the furea longer than the other terminal bristles, dorsal bristle
much thicker than the terminal ones. Anterior antennae reach
beyond the end of the furea 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. 24d).
& Unknown.
Coloration: Red in cephalothorax, sometimes on furea.
plumose bristles of maxillipeds same color. Eggs blue.
Leneth: Female, 6 mm. or over.
160 University of California Publications. [ ZooLocy
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.
i Senses *
Fig. 24.—Huchaeta spinosa. Female. (a) Head, lateral, X15. (b) Abdo-
men, lateral, X48. Gen. seg., genital segment. (c) Outer
ramus of second foot X45. (d) Basals, inner ramus, proximal
joints of outer ramus of fourth foot X45.
3. Euchaeta media Giesbrecht.
Euchaeta media Giesbrecht, 1888, p. 337; 1892, pp. 246, 263, pl.
16, figs. 13, 36; pl. 37, figs. 39, 40; 1898, p. 39.
2 Elevation on front of head low (fig. 25d); genital
segment asymmetrical with processes in the region of the orifice
and a flap on the right side of the segment behind the orifice
(figs. 25c, d) ; furea (fig. 25g) as in E. acuta. Anterior antennae
extend a little beyond the posterior border of the genital segment.
Vot.2] Esterly—Copepoda of the San Diego Region. 161
First outer marginal lobe of the maxilla (fig. 25f) 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
pair.
nor: S!
let lth rsd
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.1, first lobe of outer margin. Li.1, first lobe of
inner margin. Ri.2, 3, fused second and third joints of inner
ramus, bristles not shown. (g) Furea, dorsal, X45. Si., inner
marginal bristle.
4 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 EL.
162 University of California Publications. | ZooLoGy
acuta. Three or four females were taken during June and July,
1904.
The specimens which I have placed in this species correspond
to Giesbrecht’s descriptions; but the females have hairs on the
ventral sides of the abdominal segments, and Giesbrecht does
not mention these nor figure them (1892, pl. 37, figs. 39, 40). The
outer marginal lobe of the maxilla is always provided with eight
bristles, but one of these is very inconspicuous and much shorter
than the others. As these animals correspond very closely in
other respects to Giesbrecht’s specimens, especially in the form of
the genital segment, I have thought best to include them under
his species, even though there are slight differences.
Sub-fam. SCOLECITHRICIN AE.
Scolecithricina Giesbrecht, 1892, p. 55.
2 Head commonly fused with first, and fourth with fifth
thoracie segment; rostrum with two usually soft filaments; abdo-
men with four segments, symmetrical. Eighth and ninth joints
of anterior antennae always fused, and occasionally other joints.
Outer ramus of posterior antennae 6-jointed. Blade of 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. 306), which occasionally are pencillate
at the end; lobes of appendages closely crowded together. Inner
ramus of posterior maxillipeds at most only as long as the second
basal. Inner rami of swimming-feet jointed as in the Clauso-
calaninae and set with spines; fifth foot rudimentary or absent.
& Abdomen with shortened anal segment, number of joints
of anterior antennae reduced, the twentieth and twenty-first
often fused only in one. Other head appendages like those of
the female, or specifically modified. Left fifth foot 5-jointed,
oceasionally with inner ramus, the right 4-jointed (rudiment of
inner ramus sometimes present) or lacking.
a
Vou.2] Hsterly—Copepoda of the San Diego Region. 163
1. Genus Scolecithrix Brady.
Undina (in part) Lubbock, 1856, p. 21.
Scolecithrix Brady, 1883, p. 56.
Scolecithriz Giesbrecht, p. 337; 1892, pp. 56, 265, 264; 1898, p. 42.
Lophothriz Giesbrecht, 1895, p. 254.
Amallophora (in part) Seott, 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-
tinet, aesthetasks well developed, more numerous in male. Bitiny
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-jomted, seldom
absent; fifth foot of male uniramous on each side, or the left
biramous and the right uniramous, or both biramous.
KEY TO SPECIES.
I JBIGEKCl TANOM CHES poeodcdoudeemoaenoadomacce 5 Ute OmbanO Ow oc 2
Il, JELeRGh WANN meh (Gi, Zs) Saakaoaccagcocbucb: usoboonoonuUdDe 4
2. Anterior antennae of female 19-jointed; right of the male 17-,
acy Wein; TUSHSHOuUMHEh oc benossadcadodobs doa avo nGo cK UMaDUCma od 3
2. Anterior antennae of female 23-jointed............... S. subdentata
2. Number of joints unknown; for characters compare description
anil 306%, BO) cos ona dab ogowoooonpde cas Coo ebe se Gboon nT cor S. pacifica
3. First joint of outer ramus of first foot with a thorn-like bristle
Gm Omer mame (NE 26) os eccyo cle crere a ere wie oe ole «wis «els ate ee S. danae
3. This joint without the bristle ..........2.---.-..-2..-.0> S. bradyi
4. Anterior antennae of female 23-jointed; fifth foot (fig. 26e) ;
right antenna of male 17-jointed, left 18-jointed; fifth foot
(Cie PCED ae poise oo dot SenOie os Ole in Oreo a S. persecans
164 University of California Publications. [ ZooLoGy
1. Scolecithrix danae Lubbock.
Undina danae Lubbock, 1856, p. 21, pl. 4, figs. 6-9.
Scolecithriz danae Brady, 1883, p. 57, pl. 17, figs. 1-12.
Scolecithrix danae Giesbrecht, 1888, p. 333; 1892, pp. 265, 283, pl.
pl. 13, figs. 4, 9, 14, 17; pl. 37, fig. 6; 1898, p. 42.
hin
Fig. 26.—Scolecithriz danae. (a) Female, lateral, X20. (b) Genital seg-
ment, female, lateral, X83. (c) Outer ramus of first foot
of female X140. (d) Fifth foot of male X83. fRe.1dz.,
first joint of outer ramus of right foot. Re.sn., outer ramus of
left foot. Ri.sn., inner ramus of left foot.
2 Fourth thoracic segment separate from fifth, latter with
rather flat, rounded lateral angles. Third and fourth segments
of the abdomen broader than long, genital segment with ventral,
shovel-shaped process (fig. 265), 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 (ef. fig. 29c). First
-—- —_e SS eae
Vot.2] Ksterly.—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.
4 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,
plete. ple doy Hes. oe 7 bi 2d 28s pk Sitios. dl,
2, 9; 1898, p. 42.
/
Fig. 27.—Scolecithria bradyi. Female X31.
9° 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, furea 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 jomt
of outer ramus of first pair without outer marginal bristle; fifth
foot very small.
4 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.
166 University of California Publications. [ ZooLoGy
Length: Female, 1.4 mm.
Occurrence: June 14, 1904, one female.
3. Scolecithrix persecans Giesbrecht.
Scolecithrix persecans Giesbrecht, 1895, p. 253, pl. 3, figs. 6-12;
1898, p. 48, fig. 9.
Fig. 28.—Scolecithrix persecans. (a) Male, lateral, X9. (b) Second foot,
male, X45. (c) Left fifth foot of male X83. (d) Right
fifth foot, male, X83. fe.3, third joint of outer ramus.
(e) Fifth foot of female.
4 Tead with rather high crest (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 maxilliped 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 (fig. 285) ; spines on outer margin of the two prox-
Vou.2] Esterly—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.
@ Anterior antennae 23-jointed, reaching to end of furea;
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
leneth 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 deseribed, 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. 28d).
4. Scolecithrix subdentata n. sp.
© 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 34 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 prom1-
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
168 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. 290).
& Unknown.
ri ——\W
Fig. 29.—Scolecithriz subdentata, n. sp. (a) Female, lateral, X31. (6)
Fifth foot, female, X195. (c) Maxilla 140. B.2, second
basal. i., inner ramus. fe., outer ramus.
Approaches S. dentata Giesbrecht in form of last thoracie
segment, but the indentation is not as deep as in that species.
The fifth foot is much as in dentata, but more oval and rounded.
Distinct from dentata in possessing an outer marginal bristle
on the first joint of outer ramus of first foot, and in the number
of joints of the anterior antennae. The bristles of the maxilla
distinguish S. subdentata most sharply. S. subdentata has the
same number of joints in the antenna as S. longicornis Scott and
S. auropecten Giesbrecht.
Length: Female, 1.48 mm.
Occurrence: San Diego, May 31, June 14, June 23, 1904.
5. Seolecithrix pacifica n. sp.
2 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,
a
Vou.2] Hsterly—Copepoda of the San Diego Region. 169
outer with five (fig. 30d). 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. 830c) 2-jointed, with a short distal bristle and a
very long proximal one.
Fig. 50.—Scolecithrix pacifica, n. sp. (a) Female, lateral, X31. (bd)
Anterior maxilliped 140. (c¢) Fifth foot X195. (d) Max-
illa X83, parts as in fig. 29c.
& 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.
170 University of California Publications. [ ZooLoGy
6. Scolecithrix similis T. Scott.
Amallophora dubia var. similis Scott, T., 1893, p. 56, pl. 4, figs.
19-23.
Scolecithrix similis Giesbrecht, 1898, p. 46.
S. similis (?) Wolfenden, 1904, p. 119, pl. 9, figs. 5, 6.
Fig. 31.—Scolecithriz similis. (a) Male, lateral, X31. (b) Fifth foot,
male, X45.
4 First abdominal segment short, second long, twice the
length of the third, which is shorter than the fourth. Right
anterior antenna 18-jointed, left 23-jointed (Scott). Last two
thoracic segments fused. First basal of fourth foot with a
plumose inner marginal bristle; both feet of fifth pair biramous
(fig. 310).
Length: Male, 2.6 mm.
Occurrence: San Diego, June 23, 1904.
The antennae of the single specimen were broken, but the
|
rd
form of the abdomen and fifth feet warrant one in identifying
it with Scott’s species, at least provisionally.
Fam. CENTROPAGIDAE.
Centropagidae Giesbrecht, 1892, p. 58; 1898, p. 52.
2 Head always distinct from thorax; rostrum 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 24 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-
* a yy wohtee ee wl awe ene Ee
a a o@
——————————OoUEE— OOO ee
Vot.2] Hsterly.—Copepoda of the San Diego Region. ek
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- lke) or rudimentary, inner ramus 1-jointed or
lacking, outer ramus 1- to 3-jointed.
4 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.
Sheght sexual differences occasionally in form of last thoracic
segment and swimming feet.
Sub-fam. CENTROPAGINAE.
Centropagina Giesbrecht, 1892, p. 59.
2 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.
& 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 Dana, 1852, p. 1103.
Ichthyophorba Lilljeborg, 1853, p. 184.
Tiaptomus Lubbock, 1857, p. 403.
Ichthyophorba Claus, 1863, p. 198.
Centropages Brady, 1883, p. 81.
Centropages Giesbrecht, 1892, pp. 59, 303, 731; 1898, p. 53.
Centropages Wheeler, 1899, p. 172.
Centropages Thompson and Scott, 1903, p. 247, pl. 1, figs. 19-25.
2 Head separate from thorax, fourth thoracic segment from
fifth. Abdomen with three segments, genital segment asymmet-
172 University of California Publications. | ZooLocy
rical. Anterior antennae 24-jointed; joints 24 and 25 fused.
Outer ramus of posterior antennae 7-jointed and almost 1144
times as long as the inner ramus. The distal bristles of the ante-
rior 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 a spinous feathering;
inner ramus well developed, 5-jointed. Rami of swimming feet
usually 3-jointed, but 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 bristle of
second joint of outer ramus thorn-like and fused with the joint
(fig. 32c.)
4 Sexual peculiarities in the form of the abdomen, right
anterior antenna, and fifth pair of feet. The abdomen is com-
posed of five segments; anal segment in most species very short ;
genital opening on the left. Right anterior antenna a grasping
organ. Joints 19-21 and 22-23 fused, geniculation between the
18th and 19th. Inner marginal bristles lacking on outer ramus
of left fifth foot, joints 2 and 3 fused. The outer ramus of the
right foot 3-jointed, both distal joints metamorphosed into a 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.
Centropages violaceus Brady, 1883, p. 83, pl. 37, figs. 1-14.
Centropages bradyi Wheeler, 1899, p. 174, fig. 12.
2 Second joint of outer ramus of fifth foot with a stout
smooth spine (fig. 32c). Sides of inflated genital segment with-
out spines or knob-shaped projections. Furea symmetrical, with
a peculiar short, truneated, peg-shaped projection (fig. 320)
between insertions of the two outer bristles. (Wheeler, 1899).
4 Joint 17 of right anterior antenna with smooth anterior
border, not serrate; joints 19 and 20 fused, separated from joint
21; joint 18 with accessory series of teeth on lower surface
(Wheeler, 1899).
Vou.2] Esterly.—Copepoda of the San Diego Region. a3
Coloration: Opaque, with a large purplish spot in middle of
body.
Length: Female, thorax, 1.6 mm.: abdomen ?
Occurrence: June 10, 1904, one female.
:
Fig. 32.—Centropages bradyi. (a) Female, thorax, lateral, X45. (b)
Abdomen, ventral, after Wheeler 1899. (c) Fifth foot
x83.
SW
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 furea) 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. dorsispinatus (Thompson,
1903, p. 247, pl. 1, figs. 19-25), but differs widely in other
respects.
Sub-fam. TEMORINAE,
Temorina Giesbrecht, 1892, p. 60.
2 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.
4 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. Genus Pleuromamma Giesbrecht.
Diaptomus Lubbock, 1856, p. 27.
Pleuromma Claus, 1863, p. 195.
Pleuromma Brady, 1883, p. 45.
Pleuromma Giesbrecht, 1892, pp. 61, 347, 757.
Pleuromma Dahl, 1893, p. 105.
Pleuromma Wheeler, 1899, p. 176.
Pleuromamma Giesbrecht, 1898, p. 108.
Easily recognizable by a dark-pigmented knob on the right
or left side of the first thoracic segment (figs. 33a, 34a). Furea
at most twice as long as broad. Rami of the first to fourth pairs
of feet 3-jointed, first joint of outer ramus of third pair with a
deep notch in the outer border; terminal bristle of outer ramus
of third pair short and bent outward; first joint of inner ramus
of second pair with hooks on inner border, on right and left foot
in the female, usually on one side in male. Fifth pair in female
rudimentary, 2- to 4-jointed, in male 5-jointed on each side,
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, pl. 10, figs. 1-8.
Pleuromma abdominale Claus, 1863, p. 197, pl. 5, figs. 1-6, 13,
14; pl. 6, fig. 1-10.
Pleuromma abdominale Brady, 1883, p. 46, pl. 11, figs. 1-13.
Pleuromma abdominale Giesbrecht, 1892, pp. 347, 357, pl. 5, fig.
8; pl. 32, figs. 3, 5, 13, 22, 25-30; pl. 33, figs. 43, 44, 48,
49, 52.
Pleuromamma abdominalis Giesbrecht, 1898, p. 109.
Vou.2] Ksterly—Copepoda of the San Diego Region. AS:
® 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. 330).
eee
Fig. 33.—Pleuromamma abdominalis and P. gracilis. (a) P.a., female,
lateral, X31. (0b) P.a., fifth foot X45. (c) P.g., fifth foot
195.
¢ 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,
1904.
2. Pleuromamma gracilis Claus.
Pleuwromma gracile Claus, 1863, p. 197, pl. 5, figs. 7-11.
Pleuromma abdominale (in part) Brady, 1883, p. 47, pl. 2, figs.
UENMGe joll, Bil, aero, Way Wee
Pleuromma gracile Giesbrecht, 1892, pp. 347, 357, pl. 5, fig. 7;
pl. 32, figs. 6, 18-20; pl. 33, fig. 41-47.
Pleuromamma gracilis Giesbrecht, 1898, p. 110.
2 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).
176 University of California Publications. [ ZooLoGy
& 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. abdominalis.
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 P. gracilis are without doubt mature, since
females are often found with attached spermatophores.
3. Pleuromamma xiphias Giesbrecht.
Pleuromma sxiphias Giesbrecht, 1889, p. 6; 1892, pp. 347, 367,
pl. 32, fig. 14; pl. 33, figs. 42, 45, 50.
Pleuromamma wxiphias Giesbrecht, 1898, p. 110.
Fig. 34.—Pleuromamma sxiphias. (a) Female, lateral, X18. (b) Head
of female, lateral, X48. (c) Second basal, and proximal
joints of rami of second foot, x48.
Vou.2] Esterly—Copepoda of the San Diego Region. A by Gri
Allied to P. abdominalis, but the front of the head anterior
to the rostrum is prolonged into a prominent process (fig. 340).
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 with 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 Robertson, 1878, p. 126.
Metridia Giesbrecht, 1892, pp. 61, 339, 749; 1897, p. 254; 1898,
p. 105.
Metridia Dahl, 1894a, p. 10.
Metridia Wheeler, 1899, p. 175.
Metridia Wolfenden, 1904, p. 125.
(See also T. Scott, 1893, p. 42, pl. 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. Furea 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. pl. 8,
figs. 1-3.
Metridia armata Brady, 1878, p. 42.
Metridia hibernica Giesbrecht, 1892, pp. 340, 357, pl. 33, figs. 2,
12; 16, 22, 28,36, 39.
Metridia lucens Dahl, 1894, p. 11.
Metridia lucens Giesbrecht, 1898, p. 106.
2 Cephalothorax 124 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 34 as long as the preceding.
Furea 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 lttle over 14 as long
178 University of California Publications. | ZooLoGy
as the joint. Fifth foot 3-jointed and with three rather long
bristles on end joint (fig. 35c).
© a
Fig. 35.—Metridia lucens and M. boeckii. (a) M.l., female, lateral, 20.
(b) M.1., second basal and first joint of inner ramus of second
foot of female to show hooks, K195. (c) M.L, fifth foot
of female X195. (d) M.b., fifth foot of female 195.
4 Grasping antenna on right side. Fifth foot: second joint
of outer ramus of left foot without, first joint of outer ramus
of right foot with a long, thorn-lke bristle.
Length: Female, 3.2 mm.
Occurrence: Very common, summer and winter.
2. Metridia boeckii Giesbrecht.
Metridia boeckii Giesbrecht, 1889, p. 5; 1892, pp. 340, 346, pl. 33,
figs. 8, 19, 31, 37; 1898, p. 107. ;
2 Like M. lucens, but furca is as 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 (fig. 35d).
& Unknown.
Length: Female, 2.5 mm.
Occurrence: One or two in catches with MW. lucens.
i re a a
~
Te)
vou.2] Esterly—Copepoda of the San Diego Region. 1
It should be noted that not a male specimen of Metridia has
been taken in any eatch, so far as I have examined them, and
rather particular attention has been paid to this point.
Sub-fam. LucIcurTmuNas.
Leuckartiina Giesbrecht, 1892, p. 62.
2 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
ramus.
& (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.
Leuckartia Claus, 1863, p. 182.
Leuckartia (in part) Brady, 18838, p. 50.
Leuckartia Giesbrecht, 1892, pp. 62, 358; 1895, p. 25
Lucicutia Giesbrecht, 1898, p. 110.
Lucicutia Steuer, 1904, p. 596.
Lucicutia Wolfenden, 1904, p. 121.
oO
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
180 University of California Publications. ‘| ZooLowy
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 tube-like process. The bristle on
the inner margin of the second joint of the outer ramus of the
fifth pair has the form of a curved awl (fig. 360).
4 Sexual peculiarities in the form of the abdomen, posterior
antennae and fifth pairs of feet. Abdomen with five segments,
genital opening on right side. The left antenna is a grasping
organ, geniculating between joints 18 and 19; joints 19 to 21,
22 and 23 fused. Fifth pair of feet with 2-jointed basals; rami
of left foot 2-jointed, of right 3-jointed. The distal joint of the
outer ramus of the right foot is hooked and may be moved toward
the proximal joints.
1. Lucicutia flavicornis Claus.
Leuckartia flavicornis Claus, 1863, p. 183, pl. 32, fig. 17.
Leuckartia flavicornis Giesbrecht, 1892, p. 358, pl. 5, fig. 4; pl.
19, figs. 2, 3, 15, 17, 21, 23, 29, 38; pl. 38, fig. 38, 40.
Lucicutia flavicornis Giesbrecht, 1898, p. 111.
ri.
VS —=—
Fig. 36.—Lucicutia flavicornis. (a) Female, lateral, X18. (b) Fifth
foot, female, X83. Ri., inner ramus. (c¢) Outer margin of
outer ramus of third foot 140.
Q Anal segment shorter than the preceding; second terminal
bristle of furea thick, twice as long as abdomen. The anterior an-
Vou.2] Hsterly—Copepoda of the San Diego Region. 181
tennae reach beyond middle of the furea, 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.
& Terminal portion of grasping antenna (joints 19-25) some-
what !onger 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.
Heterochaetina 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 distinet. 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.
4 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. [ ZooLoGy
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, 1863, p. 180.
Heterochaeta Brady, 1883, p. 48.
Heterochaeta Giesbrecht, 1892, pp. 64, 372, 745; 1895, p. 259.
Heterochaeta, Aurivillius, 1899, p. 38, figs. 4, 5.
Heterorhabdus Giesbrecht, 1898, p. 113.
Heterorhabdus Wolfenden, 1904, p. 124.
2 Cephalothorax with five segments, rostral filaments soft,
situated on a papilla. Abdomen with four segments, left half of
furea 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 eight 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 maxillipeds 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 first basal
joint. All the feet have 3-jointed rami; inner marginal bristle
of the first basal in pairs one to four, and of the distal basal joint
in the first 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 (fig. 38e).
4 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.
KEY TO SPECIES.
Abdomen 4-segmented, fifth foot symmetrical.................- 2
Abdomen 5-segmented, fifth foot asymmetrical................ 5
-1. Third joint of outer ramus of third foot of same form as in
second and fourth pairs) (fies 400). ar etait H. longicornis
ee Pett
ee ee ee
VoL.2] Hsterly—Copepoda of the San Diego Region. 183
1. This joint in third pair broad and oval, terminal bristle short-
Med (OA SEO) imeers aa ctersie ssh lcr sce si ctw lens ouckelere velo enersts eo ters lo are eels fel 2
2. First basal of posterior maxilliped with a very long, heavy
bristle in the middle of the inner border; rostral papilla with
2 [oonini; (i, BiWMeso60cc0s0ccns00end suo5ad0asGon abe H. spinifrons
2. As above, but rostral papilla without point (figs. 38), ¢)........ 3
3.2 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
Jorenisiiel, 25 Iivorisay stovare (ites GE). 66 oacnaocehecccsneaease H. clausi
3.9 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 Claus.
Heterochaeta spinifrons Claus, 1863, p. 182, pl. 32, figs. 8-9,
14, 16.
Heterochaeta spinifrons Giesbrecht, 1892, pp. 372, 382, pl. 10,
figs. 1, 3, 11, 16, 19, 31; pl. 39; figs. 42, 43, 51, 52, 54.
Heterorhabdus spinifrons Giesbrecht, 1898, p. 114.
Fig. 37.—Heterorhabdus spinifrons. Female. (a) Head, lateral, X83. (6)
Posterior maxilliped 167. 8.1, first basal.
The papilla on front of head ends in a sharp point; anterior
antennae reach beyond the end of the furea 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 14 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. 370). Hooks at the end of both outer rami of the
184 University of Californa Publications. | ZooLocy
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
ramus.
Coloration; Transparent and colorless.
Length: Female, 3.4 mm.
Occurrence: June 23, 1904, one female.
2. Heterorhabdus papilliger Claus.
Heterochaeta papilligera Claus, 1863, p. 182, pl. 3, figs. 10-13, 15.
Heterochaeta papilligera Giesbrecht, 1892, pp. 372, 382, pl. 20,
figs. 4, 7, 10, 15, 17, 23, 35, 36; pl. 39, figs. 40, 53.
Heterorhabdus papilliger Giesbrecht, 1898, p. 114.
7 ---Ve2,
pS
Fig. 58.—Heterorhabdus papilliger. (a) Female, lateral, X31. (6b) Head,
female, lateral, X83. (c) Head, female, dorsal, X83. (d)
Outer ramus of third foot of male X83. (e) Fifth foot of
female 140. fRe.2, second joint of outer ramus. (f)
Right mandibular blade of male X83. (g) Anterior maxil-
liped, female, X83.
ne
VoL.2] Esterly.—Copepoda of the San Diego Region. 185
Papilla on front of head elongated but not ending in a point
(figs. 380, c). Anterior antennae when at the sides of the body
reach about to the end of the furea. Fourth lobe of the anterior
maxillipeds (fig. 389) 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 (ef. fig. 37b). 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.
Heterochacta claus Giesbrecht, 1889, p. 2; 1892, pp. 372, 382, pl.
20, fig. 2, 28, 37, 38.
Heterorhabdus clausi Giesbrecht, 1898, p. 115.
Fig. 39.—Heterorhabdus clausi. Fifth foot, male, 82.
186 University of California Publications. [ ZooLoGy
Like papilliger, but the anterior antennae reach somewhat
beyond the end of the furea. 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 .
hooked; second basal of the right fifth foot 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 left side.
Length: Male, 2-2.5 mm.
Occurrence: San Diego, July 22, 1903, one male; June 23,
1904, one male.
4. Heterorhabdus longicornis Giesbrecht.
Heterochaeta longicornis Giesbrecht, 1889, p. 2; 1892, pp. 373, 383,
pl. 20, figs. 14, 21, 25, 26; pl. 39, fig. 44.
Heterorhabdus longicornis Giesbrecht, 1898, p. 116.
Heterorhabdus zetesios Wolfenden, 1902, p. 367.
Heterorhabdus longicornis (male) Wolfenden, 1904, p. 124, pl.
9, fig. 34.
Fig. 40.—Heterorhabdus longicornis. Male. (a) Third foot 140. (6b)
Right fifth foot X140. (c) Left fifth foot «140.
2 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
a en
Vou.2] Hsterly—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.
4 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) Claus, 1863, p. 176.
Augaptilus Giesbrecht, 1889, p. 3; 1892, pp. 65, 400, 724; 1898,
p- 120.
(See also T. Scott, 1893, p. 36, pl. 2, figs. 25-37; Steuer, 1904,
p- 597.)
2 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 (ef. fig. 410). 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.
& 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).
188 University of California Publications. [ ZooLocy
1. Augaptilus longicaudatus Claus.
Hemicalanus longicaudatus Claus, 1863, p. 179, pl. 29, fig. 3.
Augaptilus longicaudatus Giesbrecht, 1892, p. 414, pl. 27, fig. 31;
pl. 28, figs. 11, 19, 23, 31, 35, 38; pl. 2, fig. 22; pl. 39, figs.
37, 48; 1898, p. 123.
Augaptilus longicaudatus Scott, 1894, p. 34, pl. 1, figs. 24-26; pl.
2, fig. 5.
Fig. 41.—Augaptilus longicaudatus. Male. (a) Dorsal X18. (b) An-
terior maxilliped X83. (c) Right fifth foot X83. (d)
Left fifth foot x83.
2 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 14 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 end inner ramus as 7:6:5. Outer ramus of
fifth foot 2-jointed.
he a i ees
a
Vot.2] Hsterly.—Copepoda of the San Diego Region. 189
4&4 Grasping antenna on left. Fifth foot, fig. 41¢, d.
Coloration: Transparent, without pigment.
Length: Male, 3.39 mm.
Occurrence: June 10, 1904, 1 male.
3. Genus Arietellus Giesbrecht.
Arietellus Giesbrecht, 1892, pp. 66, 415.
Rhinealanus (part) T. Scott, 1893, p. 31.
Arietellus Giesbrecht, 1898, 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; furea, and appendages with long, richly plumose
bristles. Anterior antennae of female and the right one of male
at most 20-jomted, joints 1 and 2, 21-25 fused; grasping
antenna on the left, 19-jointed, terminal portion 2-jointed.
Inner ramus of postericr 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, pl. 29, figs. 1, 3-7,
9-13; pl. 39, figs. 34-36; 1897, p. 254; 1898, p. 124.
With the characters of the genus.
Coloration: Terminal expansions of plumose fureal bristles
red, the remaining portion black. Body orange red, bristles on
posterior antennae and mouth parts, deep red.
Leneth: 5.5 mm.
Occurrence: One female was taken at San Diego, Dee.
22, 1903.
190 University of California Publications. [ ZooLocy
Fig. 42.—Arietellus setosus. Female. (a) Dorsal X9. (b) Fifth foot
X83. (c) Distal portion of one of the bristles of the an-
terior maxilliped «140.
4. Genus Phyllopus Brady.
Phyllopus Brady, 1885, p. 78.
Phyllopus Giesbrecht, 1892, pp. 66, 419; 1898, p. 124.
Phyllopus Wolfenden, 1904, p. 124.
2 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-
7 a... oe oO tst—‘_aOS
Vou.2] Esterly—Copepoda of the San Diego Region. Nit
einal 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. 430).
é 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, pl. 5, figs. 7-16.
Phyllopus bidentatus Giesbrecht, 1892, p. 419, pl. 18, figs. 25-33;
pl. 38, fig. 35; 1898, p. 124.
Phyllopus bidentatus & Wolfenden, 1904, p. 124, pl. 9, fig. 16.
Fig. 43.—Phyllopus bidentatus. Female. (a) Lateral X18. (b) Fifth foot
X195.
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 Califorma Publications. — [ZooLoey
Length: Female, 2.2 mm.
Occurrence: San Diego, May 31, 1904, one female.
Fam. CANDACIIDAE.
Candacidae Giesbrecht, 1892, p. 67.
Candaciidae Giesbrecht, 1898, p. 126.
2 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 and hooked. Posterior maxilliped as in Calanus
but small and weak. Inner ramus of anterior pairs of feet 2-
jointed; fifth pair rudimentary.
& Genital orifice on left; 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 Dana, 1846, p. 184.
Ifionyx Kroyer, 1848-49, p. 601.
Candace Dana, 1849,\p. 279; 1852, p. 1109.
Candace Lubbock, 1856, p. 29.
Candace Claus, 1863, p. 189.
Candace Streets, 1877, p. 139.
Candace Brady, 1883, p. 66
Candace Thompson, 1888), p. 148.
Candace Giesbrecht, 1892, pp. 67, 423, 729.
Candace Wheeler, 1899, p. 177.
Candacia Giesbrecht, 1898, p. 126.
2 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 24-jointed, proximal segments thick-
ened, anterior border toothed. Rami of posterior antennae
short, outer ramus slender, second joint elongated, terminal
enes very short. Basal of mandible large, rami short, blade
with 2 teeth. Anterior maxilliped elongate, without lobes, dis-
Vou.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.
4 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.
KEY TO SPECIES.
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 ther joubt (to. 44.0). --le ae-- 2
1. This bristle much shorter than the designated portion of the outer
loomsler’ (Gite, AIC c do eeaogo ouae one co cop bos au0ddd so euacaddS
2. Genital segment of female longer than broad........... C. pectinata
2. Genital segment of female broader than long, male not known..
Sain A ET IE ee aoc CAPE finer en Sec. ein lap ease ese eet ot aie C. bipinnata
3. The thick proximal portion of the anterior antennae is 7-jointed
(Cie anata cieaste ets cle Sr oeaieaber Yexsheiei cin dnejele.e (= mallet # "equals 4
Ze PRES) MOKWMOM, G-jOTNGER sects cer ciel ire 2-6 2 wits wee ene Seg a= oe 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.
AGC tay is orators ioe as er meeieg as cata atels wloks olen he leenshet “Asie = 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. 47b)..
An aL TER OP ee eG. Groth CAC RT RS CRC CE ONC On CR ROTOR LO C. aethiopica
1. Candacia pectinata Brady.
Candace pectinata Brady, 1878, p. 49; 1883, p. 67, pl. 30,
figs. 1-15.
Candace pectinata Giesbrecht, 1892, pp. 424, 439, pl. 4, fig. 3; pl.
91, figs. 2, 12; pl. 22, figs. 9, 17, 18, 31, 43-46; pl. 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
194 University of California Publications. [ ZooLocy
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. 440), without inner marginal
bristle; right fifth foot of male with forceps.
Fig. 44.—Candacia pectinata. (a) Last thoracic segment and first and
second segments of abdomen, male, X45. (b) 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 19 of grasping antenna, usually a
blackish brown.
Length: Females average 2 mm., males 1.9 mm.
Oceurrence: Rather common; both sexes are taken in sum-
mer and winter.
Se
Vou.2] Esterly—Copepoda of the San Diego Region. 195
2. Candacia bipinnata Giesbrecht.
Candace bipirnata Giesbrecht, 1889, p. 5; 1892, pp. 424, 439;
pl. 22, fig. 20; pl. 39, figs. 27, 29.
Candacia bipinnata Giesbrecht, 1898, p. 129.
Fig. 45.—Candacia bipinnata. (a) Cephalothorax and genital segment
of female, dorsal, X31. (b) First eight joints of anterior
antennae of female X83. (c) Last thoracic segment and
abdomen of female X20.
Q Like C. pectinata, but genital segment (fig. 45a) is broad
and has a wing-like expansion on each side. § Unknown.
Coloration: Much as in C. pectinata.
Leneth: Female, 2.6 mm.
Oceurrence: Taken usually with C. pectinata, but in fewer
numbers.
196 University of California Publications. [ ZooLocy
83. Candacia curta Dana.
Candace curta Dana, 1849, p. 279; 1852, p. 1116; 1855, pl. 78,
figs. 6 a-d.
Candace curta Giesbrecht, 1892, pp. 424, 439, pl. 21, fig. 15; pl.
22, figs. 12, 24; pl. 39, figs. 8-10, 12.
Candacia curta Giesbrecht, 1898, p. 129.
c
Fig. 46.—Candacia curta. Male. (a) Last thoracic segment, and genital
segment, lateral, X83. (b) Same, dorsal, X31. (c) Fifth
foot X83. Right foot at right of figure.
Allied to C. pectinata, but right side of genital segment in
female has a ventral projection; fifth foot of female with two
heavy teeth on the end, and one on the inner border. Proximal
joint of inner ramus of first foot with but two inner marginal
bristles.
Coloration: As in preceding species, with very slight varia-
tions.
Length: Male, 1.5 mm. Giesbrecht gives 2.4-2.65 mm.
Occurrence: San Diego, Jan. 4, 1904, one male.
4. Candacia aethiopica Dana.
Candace ethiopica Dana, 1848, p. 23.
Candace melanopus Claus, 1863, p. 191, pl. 33.
Candace ethiopica Giesbrecht, 1892, pp. 424, 439, pl. 4, fig. 13,
pl. 21, figs. 1, 9; pl. 22, figs. 1, 6, 13, 14, 32, 40-42; pl. 39,
HOS ual als:
Candacia aethiopica Giesbrecht, 1898, p. 128.
Vou.2] Esterly—Copepoda of the San Diego Region. 197
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. 472). 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. 47g).
Fig. 47.—Candacia aethiopica. (a) Female, dorsal, 165. (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, 60. (e)
Fifth foot of female X130. (f) Left fifth foot of male
x130. (g) Forceps of right fifth foot of male X130. (h)
Anterior maxilliped of female 130.
Coloration: Dorsal surface of cephalothorax, excepting an-
terior portion of head and the last thoracic segment, black brown,
198 University of California Publications. [ ZooLocy
distinguishing the species at 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.
Fam. PONTELLIDAE.
Pontellidae Giesbrecht, 1892, p. 68; 1898, p. 131.
2 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 pairs of cuticular lenses and one unpaired lens. Anterior
antennae 16- to 24-jointed, the two terminal joints always fused.
Second basal and first joint of inner ramus fused, terminal joints
of outer ramus shortened. Mandible on 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
on the proximal. First basal of 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.
3 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 orifice on left side,
erasping antenna on right, middle joints much or slightly
broadened; joints 19 and 21 and 22 to 25 fused. Fifth pair of
feet rarely with rudiment of inner ramus; forceps of right foot
incomplete or very powerful.
Sub-fam. PONTELLINAE.
Pontellina Giesbrecht, 1892, p. 68.
2 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
Vou.2] EKsterly—Copepoda 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
2-jointed.
& 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
middie 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 (sub-genus) Lubbock, 1853a, p. 25; 18530, p. 202.
Pontella Claus, 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. [ ZooLocy
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 5, sometimes asymmetrical in the female. Anterior
antennae of female 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 inner ramus. Inner ramus of swimming feet
2-jointed, outer ramus 3-jointed; fifth foot of female on each
side with 2-jointed basal portion, rami 1-jointed, though the
inner ramus may be rudimentary. Fifth foot of male 4-jointed
on each side, right without inner ramus, left at times with rudi
mentary inner ramus. The two terminal joints of the right foot
form a powerful forceps.
1. Labidocera trispinosa n.sp.
2 Cephalothorax symmetrical, evenly rounded in front;
erest, 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 3 segments,
genital longer than the last two, asymmetrical, with a prominent
blunt wing-like process on the right side (fig. 48d); middle
abdominal segment with a knob-like projection on the left in
front. Furea 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.
489), 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.
& Eye lenses larger, and more nearly contiguous (fig. 48a).
Last thoracic segment on right side with a long slender spine,
a
eee
Vou.2] Hsterly—Copepoda of the San Diego Region. 201
curving dorsally, (fig. 48), 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. 481). Fifth foot, fig. 48e, f.
A 18, 19.21.
Fig. 48.—Labidocera trispinosa, n. sp. (a) Male, dorsal, X18. (b) 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. (h) Joints 17, 18,
19 to 21, 22, of grasping antenna of male, 160.
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. brunescens 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, ete.
Coloration: Rather transparent, intestinal contents light
green, thorax and abdomen yellowish with green 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, 1892, p. 73.
Cephalothorax usually with five, seldom with six segments,
rostral filaments slender or lacking, last thoracic segment with
rounded or pointed sides. Eyes without dorsal chitin lenses.
2 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 eylindrical, 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 hooked
bristles of anterior maxillipeds long and strong, seldom so on
proximal portion of the appendage. Posterior maxilliped as in
the Pontellinae but 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.
& 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
eS ae eg A i NE A ma ee
Vou.2] Esterly—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.
Dias Brady, 1883, p. 72.
Dias Lilljeborg, 1853.
Dias Claus, 1863, p. 191.
Dias Brady, 1883, p. 72.
Acartia Thompson, 1888a, p. 149; 1888b, p. 141.
Acartia Giesbrecht, 1892, pp. 75, 506, 721; 1898, p. 150.
Acartia Dahl, 1894ce, 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.
° 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
bristies, and with three inner marginals on third joimt. 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. 49d)
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.
204 University of California Publications. [ ZooLoGy
& 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 furea 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 part and a right foot of four joints, a left of
three, each uniramous. The right especially has the form of a
claw, which, because of a process on the joint preceding the
terminal, becomes an incomplete forceps.
1. Acartia tonsa Dana.
Acartia tonsa Dana, 1848, p. 26.
Acartia tonsa Giesbrecht, 1892, pp. 508, 522; pl. 30, figs. 7, 24,
34; pl. 43, figs. 6, 10; 1898, p. 154.
a)
;
Fig. 49.—Acartia tonsu. (a) Female, dorsal, X45. (b) Abdomen of male,
dorsal, X55. (c) Fifth foot of male X138. Dz., right foot.
(d) Fifth foot of female 138.
ex.
b
Rostral filaments present, last thoracic segment rounded
laterally ; abdomen of male (fig. 495) with spines on the second
segment; anal segment with lateral hairs. Anterior antennae of
female without thorns, not reaching to the posterior border of
6 ee, > ~<a te
Vou.2] Hsterly.—Copepoda of the San Diego Region. 205
the genital segment. Middle joint of fifth foot of female about
as broad as long (fig. 49d); terminal portion of foot as long
as the rest of the appendage, straight, toothed posterior to
middle; plumose bristle as long as terminal 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. 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.
KEY TO THE GENERA OF THE PODOPLEA.
(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
LOU, JEN ee Hee Geo mob Aon Diao A Oooo to mob e-cicho y clown Por anO teccke 2
1. Inner rami of third and fourth feet l-jointed............ Mormonilla
DS binder TeANeNvis) Git THAR WOOL, JENN Sooonoedasouecloacs bvoobaagcodc 3
Ze tinereramusvOrminst: Loot) o-joImted (fo wooC)/ aes so ee eee = 14
3. Head with two large chitinous lenses (figs. 62, 57b)............ 4
35 lel@nicl qyailioui: Claniiomey IOGEEE poop aasseccnacccacounDuDEmos DDoS 7
4. Inner ramus of fourth foot 2- or 3-jointed (fig. 59b)............. 5
4. Inner ramus of fourth foot 1-jointed or replaced by bristles
(Gils, VOAO)) sao bbs food bade bias cocidaduiodsclcle co pe Sioa toes 6
5. Abdomen with four or five segments which are broadened later-
Billi ((Caiaes!, HG, BI) > coe clkckoooooeoosc vetoes aqecog6 *Sapphirina
206 University of California Publications. | ZooLocy
5. Abdomen with two segments, not broadened................. Corina
6. Eye lenses separated by at least their diameter; the last two
oat |
13.
13.
14.
14.
15.
thoracic segments without lateral sharp prolongations..... Copilia?
. Eye lenses placed close together, last two segments of the
anterior portion of the body prolonged into lateral pointed
Processes’ (fis: sO, GAD) re arccoayshn wicks mumieltic oreteleters ora re reons *Corycaeus
. Entire body much flattened, furca very long and stylet-like. .Copiliag
. Body of various shapes but more rounded; if at times depressed,
Mever leatelvce) 5.1 "ire ls ratio ete rae le a ee a ee 8
. Outer ramus of first foot 1-jointed, postero-lateral angles of
fourth segment of body prolonged into processes (fig. 54a)....
a eR a AP PUN A rie nur hrhnd RU RN ate Sets She *Clytemnestra
» Outer ramusof first. foot 2 to'3-joImbed « % joes foe wim aya whet atau os 4 0M 9
. Outer ramus of posterior antenna 1-jointed; furca very short,
each ramus with a very long bristle twice as long as the body at
least; rami of furea and the two bristles fused in the median
line; remaining furcal bristles stunted................. Aegisthus
.
. Outer ramus of posterior antenna 3-jointed; furea short, rami
separate (fig. 52c), each with a long bristle, at least as long
as the body, and at least twice as long as the other bristles
GLUES eB) eh a as at ayege sachets erste tere aftetet teat atarones ratelaaee *Microsetella
. Outer ramus of posterior antenna lacking; furea longer than
LONG, AMT HOV ALAO 212 sects ojsurhs tale laipo re Sis ete oo eeim eae steerete eters 10
. Anterior and posterior maxillipeds alike in structure, both with
LORE SPINOUS iPLISGLABn cre ot aler<apeotin = sotoisy aieie lec ioy Sina teen ora *Oithona
. Posterior maxilliped with few or no short bristles and a terminal
hgolk: ;(On¢aeidse) (hes: S505 56D) so. oe clm sw ee eee eins 11
. Fifth pair of feet 1-jointed, with two lancet-shaped appendages
at the end which have dentate borders; body elongate. .Lubbockia
. Fifth pair 1- or 2-jointed or knob-like, with naked or plumose
DEIShLOS ss body, MOLE: TOMUBU A ra 5 atayet aie niece ain mieialenalinetete olay ate wtetans 12
. Anterior antennae with very long and thick aesthetasks on the
terminal joimts; fitth feet) 2-jomied 2... oc ss) 2w 6 aieieoferee = Ratania
2. Anterior antennae with numerous pencillate aesthetasks on the
proximal joints; fifth foot a protuberance............. Pachysoma
2. Anterior antennae with few and very delicate aesthetasks; fifth
foot reduced to a small rod or kneb, or at times to one
DEISGIO? 2.5 15 waata ya ele mista iaiayatein siete Seda Note Miah auaois pee nye iol oft oNel Unio 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 1144 times as long as the
first and second Joints Losers 5... er nie ere eter *Oncaea
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
OL Lhe —proxwMalGiOLOtS (apts tate ions olepenel = eter Peano see Conaea
Front of head with two great chitinous lenses.............- Miracia
Head: -withowt slenses; jeune oo) arene cele t-perelat=/ poke siete otek net tater 15
Forehead conical, rounded in front; body very narrow; outer
ramus of posterior antennae lacking ...............+.+-- Setella
%
ee ee
— a ee
vou.2] Esterly.—Copepoda of the San Diego Region. 207
15. Forehead pointed (fig. 53a); body broad; outer ramus of pos-
terior antenna 1-jointed... ... 2.5 coc cee see cen ce eee eee esas 17
16. Furea with separate rami (about twice as long as broad) and
bristles much shorter than body (fig. 53@)............-- *Buterpe
16. Rami of the fuca very short and with the two unusually long
Vayerss Ales) Feoee eel shay qelave) semVerolreenat IbhiNe Goo Gap oc odDDUDoOUDOoC Aegisthus
Fam. CYCLOPIDAE.
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.
® 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.
4 Front of head blunt (fig. 51a) ; first and second abdominal
segments not fused (fig. 500, 51a), bristles of furea_ 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, 1848.
Oithona plumifera Dana, 1852, p. 1099, pl. 76, figs. 4a-e.
Scribella scriba Dana, 1849, p. 279.
Githona spinirostris Claus, 1863, p. 105.
Oithona plumifera Giesbrecht, 1891, p. 475; 1892, pp. 537, 548;
pl. 4, fig. 10; pl. 34, figs. 12, 13, 22, 25, 27-29, 32, 33,
44-47; pl. 44, figs. 1, 7, 12-15.
Githona plumifera Wheeler, 1899, p. 186, fig. 22.
2 Front ending in a somewhat ventrally directed, pointed
beak, but visible in dorsal view. Furea shorter than anal seg-
208 University of California Publications. [ ZooLocy
ment, 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. 50d) 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.
se ae ae ow ©
Vou.2] Esterly—Copepoda of the San Diego Region. 209
on the first and second joints, two on the third; proximal bristle
of outer margin of third joint of third and fourth pairs reduced.
4 Genital segment broad (figy505). 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: Giesbrecht shows red pigment in body, and
especially in long bristles of anterior antennae, furea, feet and
mouth parts, while other animals may be colorless. All speci-
mens I have seen are colorless.
Leneth: Female, 1-1.4 mm.; male, 0.75-1 mm.
Occurrence: Not as abundant as O. 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, pl. 4, fig. 8; pl. 34, figs.
10, 11, 20, 24, 26, 34, 35, 42; pl. 44, figs. 2, 4.
Fig. 51.—Oithona nana. (a) Male, dorsal, X83. (b) Third joint of outer
ramus of first foot of female X83. (c) Third joint of outer
ramus of fourth foot of male 83.
210 University of California Publications. | ZooLoGy
2 Front blunt; furca as long as the anal segment, hardly
twice as long as broad, outer marginal bristle about as long as
the furea. Anterior antennae’ reach about to the posterior
margin of the third thoracic segment. Second basal of mandible
with one hooked bristle. Inner ramus of maxilla with four
bristles. First, second and third (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.
& Division line between the first and second thoracic seg-
ments with a sharp median projection; genital segment nar-
rower than in O. plumifera. Proximal joint of the distal por-
tion of the anterior antennae without the round process. Third
joint of outer ramus of first to third feet with three outer mar-
ginal bristles, of the fourth with two (fig. 51c).
Coloration: Transparent, without pigment.
Length: Female, 0.7-0.8 mm.; male, 0.5-0.6.
Occurrence: Rather abundant in hauls taken from inside of
the kelp beds at Point Loma. Both sexes found. The tow in
which Oithona occurs most plentifully contains scarcely any
other genera than Oncaea, Euterpe and Corycaeus.
Fam. HARPACTICIDAE.
1. Genus Microsetella Brady and Robertson.
Microsetella Brady and Robertson, 1873, p. 130, pl. 9, figs. 11-16,
Harpacticus Dana, 1847, p. 152.
Canthocamptus Dana, 1852, p. 1187.
Ectinosoma Brady, 1883, p. 99.
Ectinosoma Mobius, 1887, p. 116.
Microsetella Giesbrecht, 1892, pp. 78, 549, 750.
2 Body eylindrical, smaller in front and behind, anterior
portion with four 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. 510).
& Posterior portion of body with five segments, anterior
antennae rather strong grasping organs; feet much smaller.
en alae a lt
Vou.2] Hsterly—Copepoda of the San Diego Region. 211
1. Microsetella rosea Dana.
Harpacticus roseus Dana, 1847, p. 153.
Canthocamptus roseus Dana, 1852, p. 1189; 1855, pl. 83, figs. 1-10.
Microsetella rosea Giesbrecht, 1892, pp. 550, 554, pl. 44, figs. 32,
So ol 054 21, 45, toa.
Fig. 52.—Microsetella rosea. Female. (a) Lateral X83. (b) Fifth foot
<195. Re., outer ramus. (c) Furea, ventral, 195.
2 Longest bristle of furea 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.
& Unknown.
Coloration: Rather transparent, region of mouth red, and
also long fureal 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 Claus.
Harpacticus Dana, 1847, p. 152; 1852, p. 1189.
Euterpe Claus, 1863, p. 109.
Euterpe 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-jointed, outer ramus 1-jointed.
Rami of swimming feet 2-jointed in the first pair, 3-joited in
second to fourth pairs; fifth pair rudimentary (fig. 53d).
4 First and second abdominal segments not fused (fig. 53a) :
anterior antennae (fig. 53¢) are powerful grasping organs,
fourth and fifth joints fused and much thickened, geniculat-
ing with 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, 1847, p. 153; 1852, p. 1192; 1855,
pl. 83, fig. lla, b.
Euterpe gracilis Claus, 1863, p. 109, pl. 14, figs. 1-13.
Euterpe acutifrons Giesbrecht, 1892, p. 555, pl. 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, .73 mm.; females slightly smaller.
Oceurrence: Abundant in catches with Oithona, Oncaea and
Corycaeus.
tle i a ae Ss ay
ee
Vot.2] Esterly—Copepoda of the San Diego Region. 213
Fig. 53.—Euterpe acutifrons. (a) Male, lateral, 175. Ab.1, first
abdominal segment. Th.5, fifth thoracic segment. (b) 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 195.
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.
2 Anterior part of body composed of four segments, pos-
terior part of five; furea 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.
214 University of California Publications. [ ZooLocy
4 Posterior portion of body with six segments, fureal bristles
sometimes lengthened. The anterior antennae are grasping or-
gans, geniculating between the last two joints; posterior maxil-
lipeds longer, with thicker second joint and longer termina!
hook.
1. Clytemnestra rostrata Brady.
Clytemnestra tenuis Lubbock, 1860, p. 160, pl. 29, figs. 6-7.
Goniopsyllus rostratus Brady, 1883, p. 107, pl. 42, figs. 9-16.
Clytemnestra rostrata Giesbrecht, 1892, pp. 566, 572, pl. 45, figs.
19, 20, 22, 25, 26, 31, 33.
Clytemnestra rostrata Wheeler, 1899, p. 189, fig. 26.
=
—
—
om
ee
=,
is
See, caaicadae
or
if
VI
Fig. 54.—Clytemnestra rostrata. Female. (a) Dorsal, X45. (b) Furea,
dorsal, 265.
Furea at most as long as broad, bristles not plumose, equal
in length in both sexes. Anterior antennae in each sex seven-
jointed, last joint in female five times as long as the preceding
one: lancet-shaped bristle lacking in the male. Outer ramus
of posterior antenna replaced by one bristle; second basal of
first 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.
Vou.2] Esterly—Copepoda of the San Diego Region. PAYS)
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.
Fam. ONCAEIDAE.
Oncaeidae Giesbrecht, 1892, p. 81.
Paired eyes with cuticular lenses and pigment bodies not
developed.
2 Form of body in general like the Cyclopidae. Each por-
tion of the furea 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
outer.
& 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, 18438, p. 62.
Antaria Dana, 1852, p. 1227.
Antaria Claus, 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 Otthona. 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.
2 Both portions of body with five segments (figs. 55a, 56a).
Anterior antennae 6-jointed, posterior 3-jointed, hooked bristles
of medium leneth(c/. 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
joint.
4 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, 1863, p. 159, pl. 30, figs. 1-7.
Oncaea conifera Giesbrecht, 1892, pp. 591, 603, pl.°2, fig. 10; pl.
47, figs. 4, 16, 21, 28, 34, 38, 42, 55, 56.
Fig. 55.—Oncaea conifera. Female. (a) Lateral X45. (b) Posterior
maxilliped 265.
2 Median portion of second thoracic segment 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. Furea 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. 555). Processes at end of third joint
of inner ramus of swimming feet very large, present even in the
fourth pair, the adjacent lancet-like bristle shortened. Fifth pair
of feet elongated, with thickened terminal bristle.
& Lips of genital orifice long, furea short and broad.
Vou.2] Hsterly—Copepoda of the San Diego Region. 217
Coloration: Often distinetly green-yellow tint to body, which
is not very transparent.
Leneth: Female, 1.2 mm.; male, about 0.8 mm.
Oceurrence: 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, pl. 47, figs. 3, 6, 26, 46, 59.
Oncaea minuta (male) Aurivillius, 1899, p. 29, figs. 1-3.
Fig. 56.—Oncaca minuta. Female. (a) Lateral 140. (0b) Posterior
maxilliped X265. (c¢) Posterior antenna X 265.
? Genital segment longer than the rest of the abdomen, the
following segments broader than long; furea 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.
4 Unknown (?)
Coloration: Reddish throughout body; eggs red.
Length: Female, 0.46-0.5 mm.
Occurence: Rather uncommon; a few come during the sum-
mer.
218 University of California Publications. [ ZooLocy
Fam. CoRYCAEIDAE.
Corycaeidae Giesbrecht, 1892, p. 83.
Paired eyes highly developed in both sexes or in females,
with large cuticular lenses and pigment bodies.
2 The broad front and the two chitin lenses, sometimes con-
tiguous and sometimes separated, are characteristic of the body
form (fig. 57b; 62.). Anterior portion of the body may be coni-
eal (Corycaeus fig. 62) or eubieal (Copilia) or oval and de-
pressed (Sapphirina fig. 58a, b; 59a. Corina). The number of
segments may be 10 (Sapphirina), 8 (Corina, Copilia) or 7 to 4
(Corycaeus); each part of the furea with only four or five
bristles. Anterior antennae 3- to 6-jointed, posterior antennae
(fig. 8c) with at least a heavy terminal hook; mandibles reduced
to blade; maxillae oval or elongate platelets, with 3 to 5 bristles;
anterior maxillipeds as in the Oncaeidae, posterior 3-jointed, ter-
minal hook strong. Rami of swimming feet 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 rudiment consisting of
a single bristle.
4 Sexual peculiarities in form of body and posterior maxil-
lipeds, also in the other appendages and more striking than in
the Oncaeidae.
1. Genus Sapphirina J. V. Thompson.
Sapphirina J. VY. Thompson, 1829.
Sapphirina Templeton, 1836.
Sapphirina Dana, 1848, p. 41; 1849, p. 281; 1852, p. 1234.
Sapphirina Claus, 1863, p. 149.
Sapphirina Haeckel, 1864, p. 102.
Sapphirina Brady, 1883, p. 121.
Sapphirina Giesbrecht, 1892, 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. Furea leaf-like, with five bristles. Rami 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.
(ee GS te i ll i gg ele A OE A i ii
ie
Vot.2] Esterly.—Copepoda of the San Diego Region. 219
2 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).
& 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, pl. 87,
figs. 1 a-d.
Sapphirina salpae Claus, 1863, p. 152.
Sapphirina gemma Brady, 1883, p. 127; pl. 48, fig. 6-8.
Sapphirina salpae Giesbrecht, 1892, pp. 618, 641; pl. 2, fig. 9; pl.
52, figs. 1, 2, 18, 19, 27, 45, 51; pl. 53, figs. 7, 23, 24, 60;
pl. 54, figs. 9, 13, 15, 16, 19, 57.
Sapphirina iris, Giesbrecht, 1895, p. 261.
2 Furea more than 21% 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 114 times as long as the three terminal
joints. Inner ramus of posterior antennae about 44 the length
of the second basal, end hook half as long as the second joint of
the inner ramus. Inner ramus of fourth foot lttle 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.
220 University of California Publications. | ZooLoGy
& Body about 21% times as long as its greatest width (fig.
57a). Eye lenses ventral, overhung by the margin of the front
of the head. Furea, anterior antennae, fourth pair of feet like
same parts in the female, posterior antennae, mandible, maxilla,
Fig. 57.—Sapphirina iris. (a) Male, dorsal, X9. (b) Female, dorsal,
x9. (ce) One ramus of furea of female, dorsal, X83. (d)
Outer ramus of fourth foot of male 140.
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.
Vot.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, pl.
87, figs. 3a, bD.
Sapphirina danae Lubbock, 1856, p. 33, pl. 12, figs. 9-11.
Sapphirina clausii Haeckel, 1864, p. 104, pl. 2, figs. 21-25.
Sapphirina angusta Giesbrecht, 1892, pp. 619, 641; pl. 52, figs. 5,
Gebs, 08, G65 pl..o3, fies. 6, 17, 29; 30; pl o4, figs. 2; 8, 17,
20, 60, 61.
aS riQ | be
Fig. 58.—Sapphirina angusta. (a) Female, dorsal, furca not shown, x14.
(b) Fourth foot of female X160. Ke., outer ramus. (¢)
Posterior antenna, female, 160. B.2, second basal joint.
Ri.2, second joint of inner ramus. (d) Furea of male,
dorsal, X60.
2 Head longer than broad; furea 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
222 University of California Publications. | ZcoLocy
of posterior antennae 5/7 as long as the second basal joint ter-
minal hook 24 as 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 34 the length of
the first and second joints together, with two bristles on the end
(fig. 58b).
& Length of trunk 214 as much as its greatest diameter. Eye
lenses as in iris. Furea, 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: Ege cases blue, otherwise as SN. iris; the males
are brilliantly iridescent.
Leneth: Female, 2.5-5 mm; male, 3-5 mm.
Occurrence: Both sexes oceur frequently m summer and
winter.
3. Sapphirina scarlata Giesbrecht.
Sapphirina scarlata Giesbrecht, 1892, p. 642; pl. 52, figs. 42, 60,
61; pl. 53, figs. 12, 39; pl. 54, figs. 25, 31, 72.
Fig. 59.—Sapphirina scarlata. Female. (a) Dorsal, X18. (b) Fourth foot,
_ ™§140. &i., inner ramus.
2 Head broad, furea 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 114 times as long as the terminal joint.
—— a a RE Ge eh
Vout.2] Ksterly.—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. 59d),
terminal joint of inner ramus as long as first or second joints,
with two bristles at the end.
4 Length of trunk not quite twice its greatest breadth, eye
lenses set back almost on the margin of the front. Furea, 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.
\ DS SNS
\ . S
\M, Mf SS \
Y Z Ses
I as
Wi \\
a \
XY
\\
Cc
d
Fig. 60.—Sapphirina lomae, n. sp. Male. (a) Posterior maxilliped X83. (Db)
Anterior antenna, bristles omitted, X83. (¢) Third joint
of inner ramus of second foot 195. (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% 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.
S. lomae 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, 1904, but
are so badly mutilated that it is impossible to make a drawing of
the entire animal.
Length: 3.2 mm.
Occurrence: San Diego, May 31, 1904, 2 males.
Nore.—The species of Sapphirina are separated into two general groups,
according as the inner ramus of the fourth foot is very small and narrow
(fig. 59b), compared with the outer ramus, or at least 34 as long as the
outer. S. iris and S. angusta belong in the latter group, and S. scarlata and
S. lomae in the former. S. iris is distinct from any other species in having
3 bristles on the outer margin of the third joint of the outer ramus of the
fourth foot; S. angusta may be recognized by the shape of the furcal rami.
2. Genus Corycaeus Dana.
Corycaeus Dana, 1848, p. 35; 1849, p. 280; 1852, p. 1203.
Corycaeus Lubbock, 1856, p. 32; 1857, p. 409; 1860, p. 182.
Corycaeus Claus, 1863, p. 154.
Corycaeus Giesbrecht, 1891, p. 480; 1892, pp. 85, 659, 735.
Corycaeus Dahl, 1894b, p. 67.
Corycaeus Wheeler, 1899, p. 191.
2 Eye lenses close together, in some cases contiguous; fifth
thoracic segment very short. Anterior antennae 6-jointed,
bristles not plumose; second basal of posterior antennae large,
first basal very short, each provided with a long, heavy bristle,
inner ramus with a thick, strongly curved hooked bristle. Ter-
mina! 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
ramus 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.
ee ee ee
Vou.2] Ksterly—Copepoda of the San Diego Region. 225
4 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, pl. 86,
fig. 4a.
Corycaeus limbatus Brady, 1883, p. 114, pl. 49, figs. 18-22.
Corycaeus venustus Giesbrecht, 1892, pp. 659, 674, pl. 51, figs. 32,
33, 34, 47.
Fig. 61.—Corycaeus venustus. (a) Female, dorsal, X83. (b) Fourth foot,
female, X140.
2 Cephalothorax with 4 segments, abdomen with 2, ventral
keel rounded, furea almost 5 times as long as broad (Genital seg-
ment: anal segment: furea:: 3:2:2).
226 University of California Publications. | ZOOLOGY
4 Genital segment about 34 as long as the anal segment and
furea 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 taken June 16, 1904.
2. Corycaeus carinatus Giesbrecht.
Corycaeus carinatus Giesbrecht, 1892, pp. 661, 675; pl. 51, figs.
20, 26.
Corycaeus carinatus Wheeler, 1899, p. 192, fig. 30.
Fig. 62.—Corycaeus carinatus. (a) Female, dorsal, X40. (b) Female,
lateral, 40.
2 Cephalothorax with 2 segments, abdomen with 1; ventral
keel beak-like and pointing back; the abdomen tapers toward the
posterior end, furea half as long as the rest of the abdomen,
about 4 times as long as broad.
& Unknown.
Coloration: Red or yellowish red pigment in region of mouth,
extensions of thoracic segments, and in the genital segment; eye
red.
Length: 0.86 mm. to 0.92 mm.
Occurrence: A few specimens taken at San Diego, Dec. 30,
1903, and Jan. 4, 1904.
Cambridge, Mass.,
January 4, 1905.
es
I
=~!
Vou. 2]... Hsterly—Copepoda of the.San Diego Region.
BIBLIOGRAPHY.
Aurivillius, Carl W. S.
1899. Animalisches Plankton aus dem Meere zwischen Jan Magen,
Spitzbergen, K. Karls Land und der Nord Kuste Nor-
wegens. Kongl. Svensk. Vet.-Akad. Hand., Bd. 32, No. 6, pp.
1-71, figs. 1-5. (Heterorhabdus angulata, new species).
Baird, W.
1843. Notes on British Entomostraca. Zoologist (Newman), Vol. 1, pp.
193-197. (Deseription of Oithona and O. plumifera.)
Boeck, Axel.
1864. Oversigt over de ved Norges Kyster jagtagne Copepoder hen-
hérende 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. Report on the pelagic Copepoda collected at Plymouth in 1888-89.
Jour. Marine Biol. Assoc., London (2), Vol. 1, pp. 144-152,
pls. 11-12. (Deseription of Gymnoplea found and Oithona,
Euterpe, Corycaeus, Oncaea).
Brady, G. Stewardson.
1878-1880. A monograph of the free and semi-parasitic Copepoda of
the British Islands. Ray Society, London.
1878. Vol. 1, pp. 1-148, pls. 1-33.
1880a. Vol. 2, pp. 1-182, pls. 34-82.
1880b. Vol. 3, pp. 1-83, pls. 83-93,
1883. Report on the Copepoda collected by H.M.S. ‘‘Challenger’’ dur-
ing the years 1873-1876. Challenger Rep., Vol. 8, 142 pp.,
do pls.
1899. On the Marine Copepoda of New Zealand. Trans. Zool. Soec., Lon-
don, Vol. 15, part 2, pp. 31-54, pls. 9-13. (New family Ento-
molepidae; 4 new genera; new species in Acartia, Temora,
Centropages (2), Ectinosoma).
Brady, G. S. and David Robertson.
1873. Coutributions 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, pls. 8-9.
Canu, E.
1896. Copepodes. Result scient. Camp. Caudan, pp. 421-437. (Abstract
in Jour. Roy. Mic. Soe. London, 1897, p. 124). [Neoscole-
cithrix new genus = Scolecithria Koehleri.|
228 University of California Publications. [| ZooLoGy
Car, L.
1884. Ein Beitrag zur Copepoden-fauna des Adriatischen Meeres. Arch.
Naturg., Jahrg. 50, Bd. 1, pp. 236-256, pl. 17. (Descriptive
notes; synonymy, distribution. Good literature list and re-
views).
Claus, C.
1863. Die freilebenden Copepoden mit besonderer Beriicksichtigung der
Fauna Deutschlands, der Nordsee und des Mittelmeeres. Leip-
zig, Verl. W. Engelmann., 230 pp., 37 pls. (Monograph deal-
ing with morphology very largely; descriptions of genera and
species).
1866. Die Copepoden-Fauna von Nizza. Ein Beitrag zur Charakteristik
der Formen und deren Abiainderungen ‘‘im Sinne Darwin’s.’’
Schriften Ges. Naturw. Marburg, 1. Supplem.-heft, 35 pp., 5
pls. (Discussion of variability, and completion of some de-
scriptions from 1863.)
1893. Ueber die Entwicklung und das System der Pontelliden. Arb.
Zool. Inst. Wien u. Zool. St. Triest, Vol. 10, pp. 233-282,
pls. 1-5.
Cleve, P. T.
1900. Notes on some Atlantic plankton Organisms. Kong]. Svensk. Vet-
ensk.-Akad. Handl., Bd. 34, No. 1, pp. 1-22, pls. 1-8. (New
species in Acartia; male of Euchirella rostrata).
1901. Plankton from the Indian Ocean and Malay Archipelago. Kong].
Svensk. Vetensk.-Akad. Handl., Bd. 35, No. 5, pp. 1-58, pls. 1-8.
(New species in Acrocalanus, Calanopia, Tortanus, Pseudodi-
aptomus ; Metacalanus new genus.)
Dahl, F.
1893. Pleuromma, ein Krebs mit Leuchtorgan. Zool. Anz., Vol. 16, pp.
104-109. (Addition of several species to the genus; consid-
eration of lateral pigmented body as a phosphorescent organ).
1894a. Leuchtende Copepoden. Zool. Anz., Vol. 17, pp. 10-13.
1894b. Ueber die horizontale und verticale Verbreitung der Copepoden
im Ocean. Verh. Deutsch. Zool. Gesell. auf vierten Jahresvers.,
pp. 61-80, 4 figs. (Distribution, diagrams and keys for
Corycaeus, Calanus, and Heterochaeta. Five new species in
Corycaeus, 4 in Heterochaeta).
1894c. Die Copepodenfauna des unteren Amazonas. Ber. naturf.
Gesell. Freib., n. ser., Vol. 8, pp. 1-14, pl. 1. (Acartia
giesbrechti new, Labidocera fluviatilis new, Paracalanus crass-
irostris new.)
Dana, J. D.
1845. Description of a new genus of Cyclopidae. Proce. Acad. Nat. Sci.
Phil., Vol. 2, pp. 285-286. (Original description of Corycaeus ;
brief descriptions of other genera and species).
1846. Notice of some new genera of Cyclopacea. Am. Jour. Sei. and
Arts, ser. 2, Vol. 1, pp. 225-230; Ann. Mag. Nat. Hist., Vol. 18,
pp. 181-185.
a
ed
Vou.2] Esterly—Copepoda of the San Diego Region. 229
1847. Conspectus Crustaceorum, in orbis terrarum circumnavigatione,
C. Wilkes e Classe Reipublicae Federatae duce, collectorum
auctore J. D. Dana. Part 1. Proc. Am. Acad. Arts and Sci.,
Vol. 1, pp. 149-155. (Cyclopidae, Tarpacticidae).
1848. Conspectus Crustaceorum quae in orbis terrarum circum-
navigatione, Carolo Wilkes e Classe Reipublicae Federatae
duce, lexit et descripsit Jacobus D. Dana, Part IJ. Proc. Am.
Acad. Arts and Sci., Vol. 2, pp. 9-55. (Calanus, Scribella,
Huchaeta, Undina, Candace, Cyclopina, Catopia, Acartia, Pon-
tella, Corycaeus, Antaria, Copilia, Sapphirina, 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 pls.; Copepoda, pls. 70-78. (Monograph; system of
classification here, adopted by later writers; descriptions very
brief and unsatisfactory, figures not from high magnifica-
tions).
Foster. E.
1904. Notes on the free-swimming Copepods of the waters in the vicinity
of the Gulf Biologic Station, Louisiana. 2nd Rep. 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 R. Corvetto ‘‘ Vettor
Pisani,’’ negli anni 1882-85 dal tenente di vascello Fran-
cesco Orsini nel Mar Rosso, nel 1884. Atti Accad. Lincei,
Roma (4) Rend.
1888. Vol. 4, 2-Sem., fase. 10, pp. 285-338.
TSSON Volo Semen tase. eles pps ll:
189la. Vol. 7, 1 Sem., fase. 10, pp. 474-481.
1891b. Vol. 7, 2 Sem., fase. 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 pls. (Monograph; descriptions, dis-
tribution, cause of migration, 59 genera, 299 species).
1895. Reports 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, pls. 1-4.
230 University of California Publications. [ ZooLouy
1896. Ueber pelagische Copepoden des Rothes Meeres gesammelt vom
Marinestabartz Dr. Augustin Kramer. Zool. Jahrb., Abth.
System., Bd. 9, Hf. 2, pp. 315-327, pls. 5-6. (Five new species
in Scolecithriz, Centropages, Monops, Oithona, Schmackeria).
1897. Notizen zur Systematik der Copepoden. Zool. Anz., Bd. 29, pp.
253-255. (Female of Arietellus setosus, Scolecithrix similis).
Giesbrecht, W. and O. Schmeil.
1898. Copepoda. I. Gymnoplea. Das Tierreich (Schulze), Lief. 6, Berlin,
1898, pp. XVI, 169, 31 figs. (5 families, 65 genera, 370 estab-
lished and 105 doubtful species, 3 varieties. Literature com-
plete to end of 1897).
Gunner, J. E.
1765. Nogle smaa rare, mestendelen nye norske Sodyr, beskrevene.
Skrifter, Kjébenhavnske Selsk. (Acta Havn.), Bd. 10, p. 175.
(Citations from Giesbrecht, 1892, 1898). [Description of
Monoculus finmarchicus-Calanus finmarchicus. |
Haeckel, E.
1864. Beitriige zur Kenntnis der Coryeaiden. Jena. Zeits. Med. Naturw.,
Bd. 1, pp. 61-112, pls. 1-3. (Systematic; anatomy of male of
Copilia and Sapphirina).
Herdman, W. A.
1891. The biological results of the cruise of the 8. Y. ‘‘Argo’’ round
the west coast of lreland in August, 1890. Trans. Biol. Soe.
Liverpool, Vol. 5, pp. 181-212. (32 species diagnosed by I.
C. Thompson).
Herdman, W. A., I. C. Thompson, A. Scott.
1897. On the plankton collected continuously during two transverses
of the North Atlantic in the summer of 1897, with descrip-
tions of new species of Copepoda and an appendix on dredg-
ing in Puget Sound. Trans. Biol. Soc. Liverpool, Vol. 12,
pp. 33-90, pls. 1-4, 4 figs. (89 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,
pls. 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 Proe. N. Zeal. Inst., Vol. 27,
pp. 214-233, pls. 15-23. (New species in Corycaeus and
Labidocera.)
1896. Zwei neve Pontella-Arten aus Neu-Siid-Wales. Zool. Jahrb.
Abth. Syst., Bd. 9, pp. 720-724, 4 figs.
Vout.2] Esterly—Copepoda of the San Diego Region. 931
Kroyer, Henrik.
1842-1845. Crustacés. Voyages de la commission scientifique du Nord
en Scandinavie . . . pendant les Années 1838, 1839,
1840, sur la corvette ‘‘La Recherche.’’ Publ. by Paul
Gaimard, Paris. Atlas, pls. 41-43. (Citations from Giesbrecht,
1892, 1895.)
Leach, W. E.
1819. Article: Entomostracés. Dict. Se. Nat. Strasbourg and Paris,
T. 14, p. 524. (Description of genus Calanus.)
Leuckart, R.
1859. Carcinologisches. Arch. Naturg., Jahrg. 25, Bd. 1, pp. 282-
262, pls. 6-7.
Lubbock, John.
1853a. Description of a new genus of Calanidae. Ann. Mag. Nat.
Hist. (2), Vol. 11, pp. 25-29, pl. 1. (Labidocera darwinn,
new genus, new species; first description of Labidocera.)
1853b. On two new subgenera. of Labidocera. Ann. Mag. Nat. Hist.
(2), Vol. 11, pp. 25-29, pl. 10.
1856. On some Entomostraca collected by Dr. Sutherland in the
Atlantic Ocean. Trans. Entom. Soc. London (2), Vol. 4,
part 2, pp. 8-39, pls. 2-12.
1857. Description of eight new species of Entomostraca found at
Weymouth. Ann. Mag. Nat. Hist., Vol. 20, pp. 401-410, pls.
10-11.
1860. On some oceanic Entomostraca collected by Captain Toynbee.
Trans. Linn. Soc. London, Vol. 13, pp. 173, 192; pl. 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. Komm.
Unters. Deutsch. Meere, 5 Ber., 12-16 Jahrg., pp. 109-126;
pl. 7-8. (19 pelagic species; tables of distribution, etc.)
Norman, A. M.
1903. Copepoda Calanoida, chiefly abyssal from the Faroé 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, pls. 3-4. (Original
descriptions of Huchaeta, Oncaea.)
Sars, G. O.
1900. Crustacea. The Norweg. North Polar Exped., 1893-1896. Edited
by Fridtjof Nansen, Vol. 1; Crustacea, V; Copepoda, pp. 35-
126, pls. 7-35. (3 new genera, 12 new species. )
232 University of California Publications. [ ZooLoGy
Scott, Andrew.
1896a. Description of new and rare Copepoda. (1) Rep. Sea-Fish.
Lab., Liverpool, pp. 32-56, 5 pls. (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.)
1896b. On Scolecithrix hibernica, a new species of Copepod, with some
remarks on the distribution of the Crustacea. Ann. Mag.
Nat. Hist. (6), Vol. 18, pp. 362-367.
190%. On some Red Sea and Indian Ocean Copepods. Trans. Liv.
Biol. Soc., Vol. 16, pp. 397-428, pls. 1-3. (New species in
Candacia, Calanopia, Stenhelia, Delavalia.)
Scott, Thomas.
1894. Report on Entomostraca from the Gulf of Guinea. Trans. Linn.
Soe. London, Ser. 2, Vol. 6, pp. 1-16, pls. 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. 419-446, 4 pls. (Six species
in Bradya, 5 new; 14 in Ectinosoma, 9 new.)
Steuer, Adolf.
1904. Copepoden der Valdivia-Expedition. Zool. Anz., Bd. 27, pp.
=
593-598, 4 figs. (Valdiviella new genus; new species in
Lucicutia and Augaptilus.)
Streets, T. H.
1877. Contributions to the Natural History of the Hawaiian and
Fanning Islands and Lower California, made in connection
with the U. S. North Pacific Surveying Expedition,
1873-1875. Smithson. Mise. Coll., Vol. 13, No. 7, pp. 138-141.
Thompson, I. C.
1888a. Copepoda of Madeira and the Canary Islands. Jour. Linn, Soe.
London, Vol. 20 (1890), pp. 145-156, pls. 10-13. (64 species;
6 new, 3 new genera including Mecynocera.)
1888b. Report on Copepoda collected in Maltese seas. Proc. Biol. Soc.
Liverpool, Vol. 2, pp. 137-151, pls. 6-9.
1888¢. Second report on the Copepoda of Liverpool Bay. Proc. Biol.
Soe. Liverpool, Vol. 2, pp. 63-71, pls. 1-2.
189Ga. Revised report on the Copepoda of Liverpool Bay. Rep. Fauna
Liverpool Bay, Vol. 4, pp. 81-136, 21 pls.
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. No. IV. Report on the Copepoda collected by
Dr. G. H. Fowler. Proce. Zool. Soc. London, 1898, pp. 540-
544. (Table; distribution.)
Vou.2] Hsterly.—Copepoda of the San Diego Region. 233
1900. Report on two collections of tropical and more northerly plank-
ton. Trans. Liv. Biol. Soc., Vol. 14, pp. 262-294, 1 pl., 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.
EGISta i) WOly 12s spp. l-sonepls; 1-7-5 (Hour enew, species
in Huchaeta, 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.
1908. 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,
pls. 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., pls. 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 Faroé Channel and Shetlands. Preliminary
note on some Radiolaria and Copepoda. Jour. Marine Biol.
Assoc., Vol. 6 (n. s.), pp. 344-371, pls. 1-4. (New species
in Aegisthus, 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. Assoe. 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 Faroé
Channel. Jour. Mar. Biol. Assoc. N. S., Vol. 7, pp. 110-146,
1 pl. 1 fig. (21 new species in Megacalanus, new genus,
ELucalanus, Gaetanus, Pseudaetideus, new genus, Aetideus,
Faroella new genus, Chiridius, Candacia, Spinocalanus, Xanth-
ocalanus 3, Scolecithria 2, Lophothrix, Heterorhabdus, Luci-
cutia 2, Augaptilus 2, Paraugaptilus new genus, Euchirella
new variety.
ah
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NON-INCRUSTING CHILOSTOMATOUS
BRYOZOA OF THE WEST COAST
OF NORTH AMERICA.
BY
ALICE ROBERTSON.
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 (757), who deseribed and figured a
number of species from San Francisco Bay. Later Gabb and
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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
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comprising, in the main, those Chilostomes which grow as free,
flexible colonies. The inclusion of the adherent genera Aetea
236 — University of California Publications. | ZooLocy
and Eucratea is somewhat arbitrary since they are more or less
inerusting. 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 1904 by the Department
of Zoology of the University of Washington, and kindly placed
at my disposal by Professor Trevor Kincaid.
In the endeavor to make this work as useful as possible to
the genera! student as well as to the expert, original figures are
given of all the species mentioned. These include a habit sketch
which is a photograph wherever possible, and one or more other
figures giving details and variations. It is thought best 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, except in two or three
instances, of detached papers scattered through scientifie jour-
nals is not readily accessible to the general student. Second,
the number of cosmopolitan species, or of species easily identi-
fied, is surprisingly small, while the number 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 Pacifie 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-Incrusting 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 Hineks, further supplemented by Miss Jelly’s invaluable
Synonymie Catalogue (’&9). 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
collections.
It is needless here to enter into an exhaustive discussion of
the bryozoa as a class. Such can be found 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 (Pl. XVI). The units of which a
zoarium is composed consist of the zowcia 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 ealeareous and opaque, or semi-calecareous
or chitinous and transparent. The contents of a zowcium whose
walls are transparent are easily made out. Below is given a
figure of a zocecium of Beania mirabilis with its contained poly-
pide in a state of retraction folded within it. The zoccium
(z@.)is seen to be a sae or bag whose front or ventral face is
bordered with a number of spinous processes, some erect (é. sp.),
others curved (c. sp.). Within the zoccium is the polypide
(pd.) consisting of a bent tube, the intestinal canal, having
238 University of California Publications. [ ZooLocy
a cirelet 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—Beania mirabilis. A zowcium and its inclosed polypide shown in
profile view. c. sp. crossed spines; car. val. cardiac valve; é. sp.
erect spine; i. intestine; m. mouth; @s. wsophagus; 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; 2a.
zoccium.
as the pharynx (ph.) which is really a portion of the cesophagus
(ws.); 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 (7.), 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 zowcium. These are part of the
parieto-vaginal muscles which assist in retracting the tentacular
VoL. 2] Robertson.—Non-Incrusting Bryozoa. 239
sheath (par. mus.). Likewise, extending from the base of the
tentacles to the basal wall of the zocecium another band of re-
tractor muscles is visible whose contraction draws the polypide
within the zoccium (re. mus.). Fig. 2 represents a zocecium of
Fig. 2—Bowerbankia imbricata. A zoccium and inclosed polypide. a.
anus; giz. gizzard; int. intestine; m. mouth; ws. esophagus; py.
val. pyloric valve; st. stomach; tent. tentacle.
Bowerbankia imbricata, in which the polypide is expanded.
Here the cesophagus (@s.) is stretched out to its fullest extent,
the mouth (m.) being at the upper margin, and the tentacles
outside the zoecium. This polypide possesses a region between
the wsophagus 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 caleareous, their branches frequently possess flexible joints at
definite points along the length (PI. V, fig. 14, j.). The zocia
included between these joints form an internode, the number
of zocecia in an internode being rather definite for any given
species. In Menipea ternata e.g., there are, as a rule, three
zocecia in an internode (fig. 14). From the lower zocecia smal]
fibres are given off known as rootlets, because they serve to
anchor the colony (fig. 14, 7.). Zocecia, 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 zoecium. This may oceupy
a part of the front wall only as in Pl. IV, fig. 2, ap.; or almost
the whole of it as in Pl. X, fig. 50, ap.).
Area—the caleareous wall inclosing the whole or part of the aperture.
(Pl. XV, fig. 88, a.).
Avicularium—an appendage of the zowcium more or less resembling a
bird’s head. This may be sessile as in Pl. V, fig. 14, av., or
pedunculated as in Pl. II, fig. 60, av.
Epistome—a ciliated lobe which overhangs the mouth, and is present
only in the Phylactolemata or fresh-water bryozoa.
Internode—the zoecia included between the flexible joints of a branch
or stem (PI. V, fig. 14).
Introvert—the thin cuticle at the anterior end of the polypide which
may be retracted into the interior of the zowcium.
Joint, or articulation—a non-calcified portion of the wall of the zoe-
cium permitting more or less flexibility in the stem or branch
(EE Vi, fie. mla9>).
Lophophore—the membranous floor or rim surrounding the mouth of
the polypide from which the tentacles spring.
Oecia or Ovicell—synonymous terms for the chamber above the zoe-
cium in which the embryo develops (Pl. V, fig. 14, @.).
Operculum—the chitinous lip by which the orifice of the zowcium is
closed (Pl. IV, fig. 2, and Pl. XIV, fig. 86, op.). When the
aperture is entirely membranous, the operculum is inconspicuous.
(RIES fio 50)
Orifice—the opening at the summit of the aperture through which the
polypide emerges. (Text fig. 1, or.).
Scutum—a modified spine overhanging the aperture (fig. 14, sc.), often
being large and flabellate (Pl. 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.
Vor. 2] Robertson.—Non-Incrusting 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.).
S
Vibracular chamber—a chamber on the dorsal side of the zocwcium in
some of the Cellulariide, resembling an avicularium, from which
extend a hairlike process called the vibraculum, and the rootlet
(Pl. 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.
LIST OF SPECIES TREATED.
Aetea anguina Linneus. Bugula pacifica Robertson.
Aetea truncata Landsborough. Bugula flabellata Thompson.
Eucratea chelata Linneus. Bugula pugeti sp. nov.
Gemellaria loricata Linnveus. 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 Robertson. Stirparia occidentalis sp. nov.
Menipea pribilofi sp. nov. Stirparia californica sp. nov.
Scrupocellaria californica Trask. Corynoporella spinosa sp. nov.
Scrupocellaria varians Hinecks. Synnotum aviculare Pieper.
Scrupocellaria diegensis sp. nov. Cellaria borealis Busk.
Caberea ellisi Fleming. Cellaria mandibulata Hincks.,
Bugula neritina Linneus. Cellaria diffusa sp. nov.
Bugula murrayana Johnston. Tlustra lichenoides Robertson.
Bugula californica sp. nov. Flustra membranacea-truncata Smitt.
242 University of California Publications. [ ZooLocy
PHoyLtuM Mo.uuuscoirpa Milne-Edwards.
Class BRYOZOA Ehrenberg.
Sub-Class Ectoprocta Nitsche.
Colonial bryozoa with anal orifice outside the lophophore;
a well developed introvert, and a spacious eclome.
Order GYMNOLASMATA Allman.
Polypide destitute of an epistome; lophophore circular.
Sub-Order I. CuHmostomata Busk.
Gymnolemata with caleareous or chitinous zocecia whose
orifice is closed by a movable chitinous lip or ‘‘operculum’’;
ova usually matured in globular omcia situated above the orifice
of some of the zowcia; vibracula or avicularia, or both, fre-
quently present.
Of the fifteen or more families included by Hincks in the
Chilostomata, all but the first seven, viz., the Membraniporide,
Microporide, Cribrilinide, Microporellide, Porinida, Myrio-
zoidw, Escharide, are omitted here. These are incrusting, or
if erect, are for the most part non-flexible in habit of growth.
Below is given a key to the first seven families of the Chilostomes
which are, for the most part, erect and flexible in habit of
growth.
KEY TO THE FIRST SEVEN FAMILIES OF THE CHILOSTOMATA.
1, Golemy \eree pings (£2 a. Faye bc eee ee ee ee ee ee oe 2
i Colonyaerect: =e er er or Pera ees eee! Se 3
2. Colony sending up erect branches from a pane just below the
2M O =) al AF < eee RoR ES ee SO oe a ee ent Eucratiide
2. Colony not sending up erect branches ee Be ne se eee eee Aeteidz
3. Colony articulated: 3 j. 6 ea ee er ees 4
3s. Colony mot ventiieulate diye: 05 ee ie oe es ene ee ae eee 5
4. Zoecia multiserial, arranged round an imaginary axis....... Cellariide
A$ ZO CCL A OVS CY NRL ace eee eee es ee ee Cellulariide
Shey Aclenoh Mirna dsweeegeoh lovee stor lop Kc eee: ee ee 6
5.) AOWGIa NO’ Wack GO: eC kre oe eee eae 7
6.: Zoweia, with: aviculariay 3-23 ee eee Notamiide
6:5 ZO C2 Glah swat hob you vil Gel ret ye eee Eucratiide
7. Zoecia biserial, avicularia pedunculated —.............22...... Bicellariide
7. Zowcia multiserial, avicularia sessile —........2.2 Flustride
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. Zowcia uniserial, arising from each
other in a tubular prolongation of greater or less length.
Aperture terminal, orifice at its summit.
The definitions of family Aeteide, of genus Aetea, and of
species Aetea anguina 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 zoecium. The adherent portion of the zocecium he ©
regarded as a stolon which together with the circle of sete 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
99
‘*stolonie portion’’ so-called, is not a stolon, but an important
part of the zocecium.
Aetea Lamouroux.
Aetea Lamouroux, 1812.
Anguinaria, Johnston, 1847.
Aetea, Hincks, 1880.
Aetea, Jullien, 1888.
Zoarium adherent to the substratum. Each zow@ciuwm par-
tially adherent, partially erect, the erect portion carrying at its
distal extremity the membranous aperture with its operculum.
KEY TO THE SPECIES OF AETEA.
1. Tubular portion of zoecium ringed, upper third spatulate, bent........
SEN 5 sos Sa eas Os SOT pa Me eR RD SB EE, ERS A. anguina
1. Tubular portion of zowcium not ringed, upper extremity not bent,
Dovite exci cl Cla iY, UlTy Cal ie eee cere eee eeee eC ILE Leann es A. truncata
244 University of California Publications. [ZooLocy
1. Aetea anguina (Linneus) Lamouroux.
Pl. IV. Figs. 1, 2, 3, and 4.
Sertularia anguina Linn., 1758, ed. 10, p. 816.
Cellularia anguina, Pallas, 1766, p. 78.
Corallina anguiformis Ellis, 1767, Ger. ed., p. 50, Pl. XXII, figs.
(as One DE
Cellularia anguina, Ellis, 1767, p. 434, Pl. 19, fig. 10.
Cellaria anguina, Ell. and Solander, 1786, p. 26.
Aetea anguina, Lamouroux, 1812, Vol. III, p. 184.
Anguinaria spatulata, Johnston, 1847, ed. II, p. 290, Pl. L, figs.
iy. Be ;
Aetea anguina, Busk, 1852, pt. I, Pl. XV, fig. 1.
Aetea anguina forma spatulata, Smitt, 1867, p. 280, Pl. XVI,
figs. 2, 4. a
Aetea anguina, Hincks, 1880, p. 4, Pl. I, figs. 4, 5.
Zoarium composed of creeping branches consisting of a
single series of zoccia growing irregularly over a stem, frond,
or other substratum (PI. IV, figs. 1, 2). Branches arising at
right angles to the zowcia from each side of the creeping por-
tion. Zoacia composed of a creeping posterior portion (fig. 2,
ad.), and an anterior, erect, tubular portion (tu.). The pos-
terior creeping portion of the zowcium narrow and tubular
where it arises from the neighboring zocecium (ad.) gradually
widening anteriorly or toward the growing point (gr.), then
turning upward almost at a right angle and becoming erect
(tu.), the remainder creeping on to give rise to a new zocecium.
The erect portion is tubular, the lower two-thirds being minutely
ringed, the upper one-third being somewhat inflated, and in
many cases bent forward thus forming the so-called snake’s
head (s. h.). At the base of the tubular portion and anterior
to it, a septum (sep.) separates the zocecium 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 zoccium (fig. 3, pd.). The
tentacle-sheath terminates above in a circle of sete which are
od
VoL. 2] Robertson.—Non-Incrusting Bryozoa. 245
everted during the expansion of the polypide. Owcia globular,
membranous, situated at the upper extremity on the ventral side
of the zocecium just below the operculum (fig. 4, @.).
As Smitt (’67), Waters (’79), and Jullien (’88) have
already shown, there has been much misconception in regard to
the zocecium of Actea. Busk (749) considered the erect, tubular
portion alone to be the zowcium, and in this error was followed
by Hineks (’80) who regards the erect tube as the zoccium,
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-ealled ‘‘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 (ov.) 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. [ ZooLocy
the ventral side of the zocecium below the operculum but exterior
to the aperture, and contains an ovum in the early cleavage
stage (@.). In all cases in which the ocecium is present the
tubular part of the zocecium 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 occium of Aetea anguina; Waters has figured
the owcium 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-
described one peculiar to this coast, but the owcium of Aetea
anguina not having been hitherto known, and no other distine-
tions being apparent between our form and that found in for-
eign localities, this 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
form.
This species is abundant at San Pedro and San Diego, grow-
ing over kelp, hydroid stems, shells, and other bryozoa.
2. Aetea truncata (Landsborough) Busk.
Pen tissi to, 6:
Anguinaria truncata Landsborough, 1852, p. 228, Pl. XVI, figs.
57, 57*.
Salpigia Hassall Coppin, 1848, p. 273, Pl. X, fig. 3.
Aetea truncata, Busk, 1852, p. 31.
Aetea truncata, Smitt, 1865, Pl. II, figs. 5-14; Pl. ILI, figs. 1-8.
Aetea truncata, Smitt, 1867, 279 and 295, Pl. XVI, fig. 1.
Aetea truncata, Hincks, 1880, p. 8, Pl. I, figs. 8-11; Pl. II, fig. 3.
Zoarium growing irregularly over the substratum. Zowcia
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 (tw.), 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 Bryozoa. 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 zocecium of the preceding species. Oacia not
known.
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 zowcium 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
ereeping 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. Zowcia
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. Ocacia globose.
KEY TO GENERA AND SPECIES.
IE MColomyaCRCe Pill Oyen ee i eee Eucratea chelata
AEC OLOT AOE CI UN cna got eee tes ee ee dee ae an eee 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 zocecia arranged uniserially, the erect branches arising from
the ventral wall of the zowcia. Zowcia prolonged into a tube
of greater or less length. Aperture large.
248 University of California Publications. - |ZooLocy
3. Eucratea chelata (Linneus) Lamouroux.
PL Vy figs: 7 859:
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.
Catenaria chelata, d’Orbigny, 1850, Vol. V, p. 43.
Scruparia chelata, Busk, 1852, pt. I, p. 29, Pl. XVII, fig. 2.
Eucratea chelata, Smitt, 1867, pp. 281 and 301, Pl. XVI, figs. 7-9.
Eucratea chelata, Hincks, 1880, p. 11, Pl. ILI, figs. 9, 10.
Zoarium ‘composed of ereeping branches consisting of a
single series of zocecia growing irregularly over the substratum
and adherent to it, and of erect branches (PI. V, figs. 7 and 8).
Creeping branches (cr. br.) arising from the sides of the ad-
herent zocecia; the erect, free branches (e. br.), from a point on
the zocecial wall just below the aperture. Zoawcia narrowed pos-
teriorly into a tube, widening gradually anteriorly; the polypide
even in contraction remaining in the anterior portion of the
zocecium. Aperture (ap.) oval, surrounded by a thin, raised,
unarmed margin; operculum at its summit (op.). Oacia
mitriform, somewhat pointed above, with a keel down the center,
borne on an imperfectly developed zowcium arising just below
the aperture (fig. 9, w.). Tentacle-sheath terminating above in
a circle of setz which are everted during expansion of the poly-
pide.
The presence of sete both in Eucratea and Aetea is given
only on the authority of others. Few of our specimens of
Eucratca possessed ocecia, and none perfect ones. Figure 9 is
taken from Hincks.
Although not abundant at any one point, Hucratea chelata
has been found at several localities on the coast of southern
California.
Gemellaria Savigny.
Gemellaria Savigny, 1811.
Notamia Farre, 1837.
Gemellaria, Hineks, 1880.
Zoarium erect, branching dichotomous, each branch given
aff trom the sides of the zowcia close to their upper extremity.
Vo. 2] Robertson.—Non-Incrusting Bryozoa. 249
Zoecia joined back to back, each pair arising from the anterior
extremity of the preceding pair. Aperture large, sloping
slightly upward. Oacia?
4. Gemellaria loricata (Linneus) Savigny.
Pl. 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, Pl. XXI, figs. b, B.
Cellaria loricata, Ell. & Sol., 1786, p. 24.
Notamia loriculata, Farre, 1837, p. 413, Pl. XX VII, figs. 6-9.
Gemellaria loriculata, Savigny, 1811, Egypt. Polyp.
Gemellaria loriculata, Van Beneden, 1844, p. 33, Pl. V, figs. 1-7.
Gemellaria loricata, Johnston, 1847, ed. II, p. 293, Pl XLVI,
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, Pl. XVII, fig. 54.
Gemellaria loricata, Hineks, 1880, p. 18, Pl. ITI, 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). Zoacia narrowed below, widen-
ing above; aperture (fig. 11, ap.) 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 Orea, Prince Willams Sound, where it was taken
from shore rocks at low tide; at Juneau, it was dredged at 20
fathoms. Hinecks reports it from Queen Charlotte Islands.
Cellulariidae Busk.
Escharide (part) Johnston, 1847.
Cabereade Busk, 1852.
Cellulariade Busk, 1852.
Cellularie Smitt, 1867.
Cellulartide, Hincks, 1880.
Zoarium erect, branching dichotomous. Zowcia in two or
more series, closely united in the same plane; avicularia and
250 University of California Publications. [ ZooLocy
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 zoccia 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 owcia. 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. Caberea is represented by
but one species, the northern C. ellisi.
KEY TO THE GENERA OF CELLARIID.
Zocecia three or more in an internode without vibracula........ Menipea
Zowcia many in an internode, vibracula present ....................------..--------
Vibracular chamber not covering dorsal surface of zoccium .......... ...
ce he A To ac St a ec eg Re ee as CR Scrupocellaria
Vibracular chamber directed obliquely and covering the dorsal sur-
face of the zocecium ................. Baa Re bree «EES ed re Caberea
De
bo
bo
Menipea Lamouroux.
Pl. V, fig. 16.
Menipea Lamouroux, 1816.
Cellularia, Johnston, 1847.
Cellarina, Van Beneden, 1849.
Emma, Busk, 1852.
Menipea, Busk, 1852.
Cellularia, Smitt, 1867.
Menipea, Hincks, 1880.
Zoacia oblong, widest above, attenuated below, often elon-
gated downward; lateral avicularia generally present, sometimes
wanting; frontal avicularia generally present on the zocecium
at the bifureation of a branch. No vibracula. Ow@cia 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, 7. d.
and r.) ; second, in a definite dorso-lateral chamber situated just
above the lateral avicularium (7. 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-Incrusting Bryozoa. 251
colony (7.); those arising in the dorso-lateral chambers may
assist In anchoring the colony when they occur on zoccia in the
lower part of the colony; usually, however, they turn upward,
and twining about the other portions of the colony, function as
tendrils (uw. 7.).
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.
KEY TO THE SPECIES OF MENIPHA.
ibe Zomcia commonly: threes am ameinGermoc eyes see eee eee 2
Te, PACs ON, waneet ay? Tira Ewa, TAKEN) eee ee Serer seee ne eeeeroreeees M. erecta
Pee SPUMCS a tWiOmOlM GNC Cg arate ee er seer ee cect 2 ee ee een asevacan te eoee 3
PA STOMMES MIO ME mt ele tai OW O Ty GEC Cee sees ree oe eee ees nee 5
3. Scutum half way down on the margin of the aperture —__................. a
3. Scutum two-thirds of the way down on the margin of the aperture...... 6
AS CUE TITS 10 eas PO = ly eeee ee eae ee score ees eee M. ternata
A Seutumelaraesy tabelabe 22s: anne OMA kes ee M. gracilis
Oe MBS CUE ULI STAM @ eee ne crete 2k se ee Ne PEL Aes ani oned Meda USSF 6
DOSS G Ul UT aa CLUaVAliGl © Cl lapeseee ace een eee ee een et eee cr eee ee ee if
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......
AS Pirie eae ne ae RR cena Ee Se BR er aes Na Se eoe ke M. occidentalis
7. Seutum two-thirds of the way down on the margin of the aperture......
BEI oe _--.------...---... MM. occidentalis subspecies catalinensis
5. Menipea ternata (Ellis and Solander) Busk.
TEI We akegss WU, dG), eS als) WOR ABI NOLS sites, alyie
Cellaria ternata Ell. and Sol., 1786, p. 30.
Cellularia ternata, Johnston, 1847, p. 335, Pl. LIX, figs. 1, 2.
Cellarina gracilis, Van Beneden, 1849, p. 67, Pl. X, figs. 1, 2.
Menipea ternata, Busk, 1852, pt. I, p. 21, Pl. XX, figs. 3-5.
Cellularia ternata, Smitt, 1867, p. 282, Pl. XVI, figs. 10-26.
Menipea ternata, Hincks, 1880, p. 26, Pl. VI, figs. 1-4.
Zoarium consisting of loosely spreading, straggling branches,
‘or sometimes of rather large dense masses, frequently 30
bo
Ol
bo
University of California Publications. [ ZooLocy
to 35 mm. high and from 60 to 80 mm. broad (Pl. V,
figs. 12, 13). Branching dichotomous. Internodes consisting of
three zocecia, as a rule, although there are many instances of
five or seven zoccia to an internode. The longer internodes are
generally terminal and bear the ovicells. Joints light colored,
arising in distinct tubes (fig. 14, j.). Zoacia elongated, narrowed
below; aperture oval (ap.), 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; zoccium
at bifureation of branch having sometimes one, sometimes two
rather long spines at its summit. Towards extremity of branches
the spines increase in length, often very considerably. Scutwm
varying in shape from a mere spine to a somewhat broad flabel-
late process (sc.). Lateral avicularia (av.) large, and promi-
nent, generally developed on all the zocecia of an internode
except the one at 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. Oacia smooth, somewhat
elobose, reaching not quite to the lower margin of the next upper
zocecium (oe.). From a flat disk in front or to one side of the
aperture of many of the lower zoccia in a colony, root fibers (7.)
extend downward, assisting in anchoring the colony to the sub-
stratum; from zocecia higher in the colony other upward ex-
tending fibres arise in a dorso-lateral chamber (7. ch.) which is
slightly raised and which projects from the zocecial 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
Islands.
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 zocecia 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 p», and that of the species found
VoL. 2 | Robertson.—Non-Incrusting 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
spring.
6. Menipea gracilis Busk.
Pl. 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 zoceeia;
internodes of three zowcia only, often very long, ranging from
1,600 to 1,900 » in length. Zoawcia enlongated, aperture large.
(fig. 19). Zocecia with two or three spines, depending upon
the presence or absence of the lateral avicularia. 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 seutum. Zocecium
at bifurcation possesses one or two spines, more often one, at
its summit. Scutwm (sc.) large, flabellate, projecting outward
and arching over the aperture in well developed specimens, pro-
jecting beyond the outer edge of the zoecium. Lateral avicularia
frequently absent, when present often minute. Frontal avi-
cularia (fr. av.) oceur on the zocecium at the bifurcation of the
branch, but these too, are frequently absent; on ocecial inter-
nodes,. however, they are often found on each zocecium, situated
a little to one side of the aperture and so close to the owcium
as to seem perched upon its upper margin (fig. 20, av.).
Oacia (oe.) high, rounded, with striations radiating from a
thin place at the base of the owcial 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 Califorma 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.
PA Gh):
This species is abundant at Orea, Prince Williams Sound;
reported by Hineks from Cumshewa Harbor, Queen Charlotte
Island.
7. Menipea occidentalis Trask.
Pl. VI, figs. 22, 23, 24, 25.
Menipea occidentalis Trask, 1857, p. 113, PISA, hip.
Menipea compacta Hincks, 1882, vol. 10, p. 461.
Menipea compacta Hincks, 1884, vol. 13, p. 208, Pl. LX, fig. 8.
Zoarium forming bushy tufts from 15 to 50 mm. in height,
attached by a large number of root fibres. Root fibres not de-
veloped throughout the colony, this species being in no sense a
climber. 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.). Internodes consisting of three zocecia (fig. 23). Joints (j.)
yellow or brown arising from definite chambers on each side of
the most anterior zocecium of an internode. Zoawcia elongated,
narrowed below; aperture (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
(u. sp.), and between these two and the two crossed spines, two
other opposite flaring spines (fl. sp.) ; on the terminal zocecia
these flaring spines often grow very long so that each zocecium
is then bordered anteriorly with a bristlng array of four long
spines. Scutwm (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 zocecia except the one at the bifurcation of a branch.
Frontal avicularia wanting. Oa@cia usually developed on the
tertiary branches, large, globose, covering the zocecial wall below
VoL. 2] Robertson.—Non-Incrusting Bryozoa. van
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 zocecia 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 (757),
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
searce on the shore and in dredgings, and the colonies are small
and delicate.
8. Menipea occidentalis catalinensis. Subsp. nov.
Pl. 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 seutum 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 zoccia of the subspecies catalinensis are larger
than those of the type M. occidentalis, but it was unintentionally
made on a larger scale of magnification than was that of figure 23.
9. Menipea erecta Robertson.
PI Val, fies..28) 29573031"
Menipea erecta Robertson, 1900, p. 317, Pl. XTX, 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, five, or seven zocecia, the number in-
ereasing toward the extremities of the branches; articulations
extending through the zoccia just below the aperture, some-
times including its lower border (fig. 29, art.). Zoacia biserial,
alternate, narrowed below ; aperture oblong, occupying more than
half the front; margin raised, crenulate, with one or two blunt
spines at the upper outer angle; scutuwm (sc.) a flattened spine,
sometimes growing broad and bifid. Lateral avicularia fre-
quently wanting (fig. 30), sometimes feebly developed (fig. 31),
and again rather large (fig. 29). When avicularia are present
there is usually but one spine at the upper, outer angle, although
this is not the invariable rule. Frontal avicularia (fr. av.) 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. Oacia large, glo-
bose, more or less striated. Root fibres mainly upon the lower
zocecia. springing directly from the zocecial wall and extending
downward (fig. 30, 7.).
Examination of material from several localities, especially
of that from Puget Sound, leads me to unite the species for-
VoL. 2 | Robertson.—Non-Incrusting Bryozoa. 257
merly identified as Scrupocellaria scabra Van Beneden with
Menipea erecta Robertson (700).
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 zoecia 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. nov.
Pl. VII, figs. 32, 33; Pl. VIII, fig. 34.
Zoartum 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 (Pl. VII, fig. 32.). Branching
dichotomous, main branches long, possessing a number of shorter
inward curving secondary branches. Jnternodes consisting of
three zocecia except the ocecial internodes which generally con-
sist of five; separated by dark colored chitinous joints. Zowcia
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 bifureating zocecium may have
but two spines, but it is frequently found bristling with three
or four. Scutum (Pl. VIII, fig. 34, sc.) simple, often a mere
spinous process; when better developed, it broadens at the free
extremity. Spines and secuta distinctly jointed. Lateral avi-
cularia generally present, large (lat. av.). Frontal avicularia
(fr. av.) few, when present found only on the zocecium at the
bifureation of a branch, large, raised, beak set obliquely, some-
times transversely to the length of the zocecium. Owcia globose,
smooth (Pl. VII, fig. 33, 0e.). 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 (Pl. VIII, fig. 34, r.).
Often on a zocecium in the same internode, and on zocecia at
higher levels, just above the lateral avicularia, other rootlets
eurving upward, develop from a circular chamber slightly raised
above the zocecial wall (wv. 7.).. These upward growing fibres are
of much less frequent occurrence than in M. ternata 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 M. occidentalis rather than M. ternata.
Menipea pribilofi 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, Unalaska, and Yakutat, Alaska.
Serupocellaria Van Beneden.
Cellularia Pallas, 1766.
Cellaria, Ell. and Sol., 1786.
Scrupocellaria Van Beneden, 1844.
Cellularia, Johnston, 1847.
Canda, Busk, 1852.
Scrupocellaria, Hincks, 1880.
Zoarium jointed. Zowcia numerous in each internode, rhom-
boid; aperture with or without scutum; a sessile aviculariwm 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.
KEY TO THE SPECIES OF SCRUPOCELLARIA.
i. Vibracular chamber tonpey ery) Z0ccerimae serene See eee 2
1. Vibracular chamber not on every z0@¢ium <2 2 3
2. Vibraculum slightly longer than a zoccium ...........-..--.--...-- S. varians
2. Vibraculum as long\as three: zomeia ee S. diegensis
3. Vibraculum not as long as a zocecium ..............2--.2-20------ S. californica
VoL. 2] Robertson.—Non-Incrusting Bryozoa. 259
11. Serupocellaria californica Trask.
Pl. VIII, figs. 35, 36, 36a, 37.
Scrupocellaria californica Trask, 1857, p. 114, Pl. 4, fig. 2.
Scrupocellaria brevisetis (?) Hincks, 1882, p. 462.
Zoarium growing in large compact tufts, somewhat coarse in
appearance. (PI. VIII, fig. 36a.) Branching dichotomous, inter-
nodes consisting of five, seven, or nine zowecia. Zowcia shghtly
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 seutum. Zocecium at the
bifureation (z@. bi.) 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 secutum. Scutwm (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 zocecium just below the
aperture. Vibracular cell frequently lacking, and not visible
from the front surface except on the zocecium at the bifurcation
of a branch (v. c.). Vibracula, when present, found only on
the zocecia of the lower part of the colony where root fibres are
given off (r.). This is usually true except of the zocecium at
the bifurcation of a branch where vibracular cells are generally
present irrespective of the formation of a root fibre. Vibracu-
lum shorter than a zocecium; 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 zocecium
(gr.), the short stout vibraculum much like the elongated upper
mandible of an avicularium. Oacia 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, 7.).
260 University of Californa Publications. | ZooLocy
This species first figured and described by Trask (757) from
San Francisco Bay is probably the S. brevisetis, 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. californica 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. Serupocellaria varians Hincks.
Pl VOW, fies: 38,39- Blo XV1, fig. 95:
Scrupocellaria varians Hincks, 1882, p. 461, Pl. XTX, figs. 1-le.
Zoarium forming bushy tufts 12-25 mm. in height. (PI.
XVI, fig. 95.) Branching dichotorhous, internodes consist-
ing of a variable number of zocecia, those in the lower part of
the colony containing usually five or seven zoccia, those at a
higher level, nine or eleven. Joints yellow. Zowcia biserial,
alternate, slightly narrower below; aperture oval, occupying
more than half the front; scutwm sometimes a mere spine, some-
times bifid or trifid, usually inclined downward (Pl. VIII, fig.
38) ; two spines on the upper margin, one of them just above the
seutum, 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. av.) ; 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.). Owcia (fig. 38, oe.), smooth, developed on each
zocecium of the terminal internodes. Rootlets springing from the
base of the vibracular chamber (fig. 39, 7.).
VoL. 2 | Robertson.—Non-Incrusting 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 (mus.) ; thus, in fig. 39 (¢. 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. cali-
fornica, but as has been said, the increase of size of the avicularia
in the two eases 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 seuta, 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 Hineks 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. Serupocellaria diegensis, sp. nov.
Pl. IX, figs. 41, 42, 43, 44; Pl. XVI, fig. 96.
Zoarium forming a coarse bushy mass often 50 mm. in
height (Pl. XVI, fig. 96). Branching dichotomous, internodes
consisting of a variable number of zocecia, nine. thirteen,
seventeen or more, especially in the terminal internodes. Joints
somewhat inconspicuous, often occurring as high on the zocecium
as the lower margin of the aperture. Zoacia biserial, the two
rows inclined at an angle so that the internode is keeled, and
in cross section is triangular; rather short and broad, shghtly
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 zocecia large, covering more than half the
aperture, thickened on the inner surface and raised on a ped-
unele. Spines three, four, or five, the one immediately above
the seutum, and frequently the one opposite, bifid (67. sp.). In
262 University of California Publications. [ ZOOLOGY
well developed zocecia there are three spines on the outer mar-
gin, 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 avicularia
of moderate size, usually found on each zoccium. A series of
sessile, frontal avicularia (f7. av.) extends between the zocecia,
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 zoccia
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 zocecium at the bifureation
of a branch usually bears a raised avicularium of extraordinary
size, with elongated beak directed obliquely, sometimes to the
right, sometimes to the left (67. av.). Vibracular chamber dorsal,
large, its length equalling half that of the zocecium on which it
is placed (fig. 42, v. ch.) ; vibraculum long, two and a half times
as long as the individual zowcia (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 zoccia the place of
the rootlet is marked by a pore (fig. 42, p.) Oacia 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 calear-
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 Caberea, 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 (752), but its habit
is very different and it does not possess the peculiar scutum for
which the latter is named.
S. 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
locality.
Caberea Lamouroux.
Caberea Lamouroux, 1816.
Flustra, (part), Johnston, 1847.
Cellularia (part), Johnston, 1847.
Caberea, Busk, 1852.
Canda, d’Orbigny, 1850.
Caberea, Smitt, 1867.
Caberea, Hincks, 1880.
Zoarium not articulated. Zoacia 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 lne, 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; Pl. IX, figs. 45, 46.
Flustra ellisii Fleming, Mem. Wern. Soe. ii, p. 251, Pl. 17, figs. 1-3.
Flustra setacea Johnston, 1847, ed. Il, p. 346.
Cellularia hookeri, Busk, 1852, pt. I, p. 37, Pl. XX XVIII, 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). Zowcia in two or four rows,
short, subquadrangular (Pl. IX, fig. 45); aperture elliptical,
occupying nearly the whole of the front, with a broad minutely
eranular 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 (lat. av.) small and inconspicuous, placed a little below
the top of the zowcium, with a rounded mandible; frontal avicu-
laria (fr. av.) raised, two below the aperture, placed one on
each side, or sometimes only one; mandible rounded, directed
downwards. Vibracula very long, serrate (v.). Owcia flattened,
frequently with a depressed, smooth, semicircular space in front
from which fine striae radiate to the margin. footlets arise
from the side of the vibracular chamber, the two on opposite
sides uniting in the median dorsal line with each other and with
those below, forming a thick keel-like cable along the dorsal sur-
face of the branches (fig. 46, 7.). 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.
Bicellariade Busk, 1852.
Bicellariee Smitt, 1867.
Bicellariide, Hincks, 1880.
Zoarium erect, with or without articulated stem or pedunele,
or composed of a number of zocecia connected by tubular pro-
cesses. Zowcia rather loosely united in one, two, or more series,
or disjunct, boat-shaped or sub-tubular; aperture occupying a
large proportion of the front. Avicularia, when present, capi-
tate, pedunculated, and articulated. Owcia either pedunculated
or sessile.
The diagnosis of the family Bicellartidw as given by Hincks
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 zoccia.
KEY TO THE GENERA OF BICELLARIIDAS.
COW O ray ae LCC ex os reese eee ee ce ee
Goloniy:. ereeprin ce 526 eee a i ence eee
Gollomy,-wumsiialle dy Bee ac ee
Gollomiy; stevie aa a ee eee
FioceGlan HUIS err | cone sears 5 eee en ee ee ee Corynoporella
Fomeia. i= wOr Mla SO rey eee cease eee ree ee eae nate ee Bugula
PO CGA WWUSCTUE UN See aN rer ee ee See ee Stirparia
Zoewcia uni- or multiserial, with tubular connections ................ Beania
H® Co O1 bo
OUR 99 69 PO DO
~ l
—
Vou. 2] Robertson.—Non-Incrusting 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.
Ornithoporina d’Orbigny, 1851.
Bugula, Busk, 1852.
Bugula, Hincks, 1880.
Zoarivum erect, branched. Zoacia 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 zowcium. Oacia sessile
at summit of aperture, or pedunculate upon the side of the
aperture.
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
murrayand, is circumpolar, and another, B. neritina, is ecosmo-
politan in range.
KEY TO SPECIES OF BUGULA.
Ji SUT OSCOMEY So ONISK Sta IES SES ew Lo ane 8 ae oe Bc epee ae J Ne 2
The see OTE, ATO LES (ENE ESSA eA ae ee ed en Ue een 4
2. Avicularia, if present, at summit of aperture -............... B. neritina
Zee ASYM aT mete a SOMO Ap CL GUE Cyst ee rence ge ne ee B. pacifica
2. Avicularia between summit and base of aperture =... 0... 3
3. Avicularia much elongated, half as long as zowcium, slightly more
than half way between summit and base of aperture
Re ee os RE SN, Aon IRAE STi ee NE B. longirostrata
3. Avicularia not elongated, less than half the length of a zowcium, just
half way between base and summit of aperture....B. californica
Ape AVLGTU RABY ap TSE ier teens 8 eR he eet ged ea EAs a RD 5
cI a)a Nv CVU: Tose Bese] OYS(2) 01 ee ke, A a eee eo 2 B. laxa
DaeAWwAeUl aria at baser Oh aOerbiut esse eee eee ee -B. murrayana
5. Avicularia between base and summit of aperture _.............00...... 6
6. Avicularia less than half way from base and summit of aperture........
BS eee eas oa. eee a ase ee eee eb. flabellata
6. Avicularia near base of aperture, beak much curved _...
poe SSE At at Bnad AO Seema BA Soang Mares Pete EN (Oe. tS Oe B. curvirostrata
266 University of California Publications. [ ZOOLOGY
15. Bugula neritina (Linneus) Oken.
Pl. IX, fig. 47; Pl. XVI, fig. 97.
Sertularia neritina Linneus, 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.
Acamarchis 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, Busk, 1852, p. 44, Pl. XLITI, figs. 1-6.
Bugula neritina, Waters, 1887, p. 91, Pl. IV, figs. 3, 15.
Zoarium consisting of large bushy tufts 75 to 100 mm. in
height, brown or reddish brown, often tinged with purple.
(Pl. XVI, fig. 97.) Branching dichotomous. Zoecia biserial,
quadrangular, truneate above; aperture occupying more than
two-thirds of the front; a short spine or denticle at the summit
of the sides of the zowcium (PI. IX, fig. 47, den.). Owcia very
conspicuous and numerous (0e.); globose, attached to the inner
anterior angle of the zocwcium by a short peduncle. Avicularia
none. Rootlets 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,
ete., 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. Il, p. 347, Pl. LXITI,
figs. 5, 6.
Avicella multispina Van Beneden, 1848, Vol. 15, pt. I, p. 76, figs.
(he $e
? 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-Incrusting Bryozoa. 267
more mm. in diameter (Pl. XVI, fig. 98), or of much elon-
gated, narrow, strap-like branches (fig. 99). Zoawcia multi-
serial, the branches having from three or four, to twelve or four-
teen rows; alternate, oblong, slightly narrower below, truncate
above (Pl. 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 (ba. sp.). Avicularia pedunculate, those on
the marginal zowcia, when present, much larger than those on
the intermediate zoccia; often absent from both marginal and
intermediate zocecia. Oacia (oe.) large, subglobose, marked by
radiating lines. Rootlets given off from marginal zocecia near
base of colony.
B. murrayana is found at Kadiak, Orea, Pribilof Islands,
Alaska, and in Puget Sound, but does not extend farther south.
The specimens from Orea, Prince Williams Sound, are ro-
bust, often possessing a large number of zocecia 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 zocecia 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 californica sp. nov.
Pl. X, fig. 49; Pl. XVI, fig. 100.
Zoarium consisting of a number of masses whose branches
have a distinetly 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 zowcia, dichotomously divided to form a fan or
frond. Zowcia narrow below, elongated, bearing three spines at
the distal extremity, one stout and long (Pl. X, fig. 49, sp.),
extending in a direction parallel with the branch, longer than
the other two, a continuation of the wall of the zocwcium; 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 zowcium. Avicularia large, pedun-
culate, situated near the top or not more than half the length
of the zocecium below the top. Oacia large, high, globose, extend-
ine above the lower margin of the aperture of the next upper
zocecium. Rootlets developed on lower zocecia, springing from
front or ventral surface of zocecium below the aperture.
The zoccia 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 zocecia are somewhat
shorter and stouter; the avicularia are of much the same form
and size, but are placed lower down on the zocecium; 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 B. turbinata.
Colonies of large size oecur abundantly at Dillons Beach,
California; B. californica is also found, but of less luxuriant
erowth at Lands End, San Francisco Bay, and at Pacific Grove,
Monterey Bay.
18. Bugula pacifica Robertson.
Pl. X, fig. 50; Pl. XVI, fig. 101.
Bugula purpurotincta, Robertson, 00, p. 320.
Bugula pacifica Robertson, 700, p. 321.
Zoarium consisting of large bushy tufts having a somewhat
spiral growth, often 75 mm. in height (Pl. XVI, fig. 101) ; col-
onies frequently united by a sort of cable formed by union of
VoL, 2] Robertson.—Non-Incrusting Bryozoa. 269
the root fibres. More delicate in appearance than the preceding
species, often tinted purple. Zowcia elongated, biserial, armed
at the distal extremity with three spines, the outer one lone
(300 »), projecting outward (Pl. X, fig. 50, sp.) ; the other two
shorter (100 »), being continuations of the margin of the aper-
ture (sp.’); Aperture occupying almost the whole of the front
of the zoecium. Avicularia large, pedunculate, at the base and
to the outer side of the aperture. Oq@cia remarkably small, not
rising more than 60 or 80 » above the zoccium, while the em-
bryo (emb.), in many cases measuring 200 p, extends down-
ward into the upper part of the zoccium. Rootlets numerous.
extending from the lowest zocecia of a colony.
This species was formerly (’00) somewhat. tentatively iden-
tified as B. purpurotincta 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 zoccia, and in part in the degenerated poly-
* and is quickly lost after the material
pides or ‘‘brown bodies,’
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 omwcia 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 occium, 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
ocecia elevated above the mouth or orifice of the zowecium, func-
tional polypides and embryos cannot exist simultaneously in
270 University of California Publications. | ZooLocy
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 Orea, 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.
19. Bugula flabellata J. V. Thompson.
Pl) axe dies SoZ:
Bird’s Head Coralline Ellis, 1767, Ger. ed., p. 109, Pl. XXXVIII,
Olt.
Cellularia avicularia (part) Pallas, 1766, p. 68.
Flustra avicularis, Johnston, 1847, p. 346, Pl. LXIII, figs. 3, 4.
Avicularia flabellata Thompson, 1847, MS. Brit. Mus.; Gray, Brit.
Mus. Radiata, p. 106.
Avicella avicularia Van Beneden, 1848, p. 75.
Ornithoporina avicularia, d’Orbigny, 1850, p. 322.
Bugula flabellata, Busk, 1852, p. 44, Pls. LI, LILI.
Bugula avicularia forma 2, B. flabellata, Smitt, 1867, pp. 290, 345.
Bugula flabellata Thompson, 1868, Pl. 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 flustroides, Verrill, 1879, p. 52.
Bugula flabellata, Hincks, 1880, p. 80, Pl. XI, figs. 1-3.
not Bugula flabellata, Robertson, 1900, p. 431.
Zoarium from 12-25 mm. in height, consisting of a number
of fan-shaped fronds divided into narrow branches (fig. 51).
Branching dichotomous, occurring where a branch reaches a
width of six or seven rows of zoccia. Zowcia elongated, of about
equal length throughout, aperture 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-Incrusting Bryozoa. 271
way from the top of each zocecium; those on the intermediate
zocecia smaller than those on the margin; marginal avicularia
of medium size, point of mandible bent abruptly downward.
Oawcia small, globular, smooth, well elevated above the top of the
zocecium, 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 zowcium. It agrees,
however, with the figure given by Busk in which he represents
three spines on the outer side of the marginal zocecia. 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. nov.
PIX, figs. 53,54; Pl. Xl, fig: 55.
Bugula flabellata, Robertson, 1900, p. 321.
Zoarium composed of numerous small somewhat spiral tufts
from 10 to 25 mm. in height (fig. 53). Zowcia 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 zocecia,
two on the outer side (ou. sp.), one on the inner (in sp.) ; inter-
mediate zocecia with two spines only; the middle of the anterior
edge of each zocecium occupied by a round knob or process
whose significance is unknown (k.). Avicularia (av.) found
only on the marginal zoweia, large, beak somewhat curved, sit-
uated half way from the top, or just below the middle of the
zocecium. Oacia none. Embryos (Pl. XI, fig. 55 emb.) develop-
ing in the anterior part of the zocecium, and no additional ocecium
being formed. Embryos abundant in the older parts of the
colony, and those zocecia possessing well developed embryos con-
taining no polypide. Rootlets extending from the lower zocecia
of the colony, forming a cluster at its base and attaching it to
shells, rocks, or sea weed.
272 University of California Publications. [ ZooLocy
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 owcia are not 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. In the zoccia just posterior
to these, full grown polypides appear and the genital products
are abundant. In still older zowcia, a young embryo appears
close to the anterior edge of the orifice of each zocecium, and
the polypide in such a zocecium begins to degenerate. In older
zocecia, nothing is left of the polypide but a brown body, while
the embryos are well developed, almost ready to hatch (Pl. XI,
fig. 55, emb.; b. b.) ; in still older zowcia, regenerating polypides
(re. pd.) are found in zoccia 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
zocecium, are not the only characters which separate this form
from B. flabellata. The shape of the avicularia, and the number
and position of the spines are sufficient and constant specific
differences.
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. nov.
Pl. XI, figs. 56, 57, 58.
Zoarium consisting of numerous strap-lke, dichotomously
branched tufts, from 35 to 75 mm. in height (fig. 56); the
branches much tangled, due to the binding fibres (6. f.) given
oft from the different parts of the colony which unite adjacent
branches or portions of a branch. Zoewcia 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-Incrusting Bryozoa. 218
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.’). Ow@cia (fig. 58, oe.) globose, large,
marked by faint strie. 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 seale. 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.
University of California Publications. [ ZOOLOGY
bo
4
TS
22. Bugula longirostrata, sp. nov.
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. Zoawcia (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 zocecium. Avicularia (av.) extremely
slender and long, being more than half the length of a zocecium ;
the beak (bk.) alone being almost half as long as the whole
appendage; pedunculate, arising on the outer, lateral wall of
the zocecium about half way from its summit; when extended in
a direction parallel with the zowcium, the tip of the beak of the
avicularium extends beyond the summit. Owcia (o0e.) small and
shallow, just sufficiently high to inclose the ovum (0.), but cover-
ing only a portion of the embryo (emb.), the major portion of
which occupies the upper part of the zoecium. Rootlets arising
from the dorsal wall of the lower zoccia.
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 owcia, these being insufficient to inclose the
embryos when fully grown. The ocwcium of B. longirostrata is
shghtly larger than that of B. pacifica, but relatively to the
length of the zowecium 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-Incrusting Bryozoa. 275
23. Bugula laxa, sp. nov.
Pl. XII, figs. 61, 62.
Zoarium consisting of numerous fronds much divided into
segments possessing from two to fifteen rows of zoccia 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. Zowcia 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
zocecium, the outer, marginal zocecia usually possessing, besides
the stiff spines at the summit, nine spines at regular intervals
along its length; the intermediate zocecia having a smaller num-
ber; aperture occupying the whole of the front of the zocecium.
Avicularia lacking. Oa@cia (oe.) rounded, prominent, with
faintly radiating striw. Rootlets (fig. 62, r.) numerous, arising
from the right or left dorsal anterior angle of the marginal, and
of many of the intermediate zoccia.
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 zowcia, the looseness of their connection with
each other, and more especially the fact that each individual
zocecium arises from a bud formed independently from a parent
zocecium, 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 zocecium from which either one
or two buds arise forming branches which extend sometimes as
long narrow strips of one or two series of zocecia, sometimes as
broad fronds of many series. The adjacent rows of zocecia are
276 University of California Publications. [ ZooLoGY
so loosely connected that in the broader frond-lke portions.
lacune frequently oceur where zocecia 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
Grove.
Beania Johnston.
Beania Johnston, 1847, ed. II, p. 372.
Diachoris Busk, 1852, pt. I, p. 53.
Zoarium subecorneous or ecaleareous, erect or decumbent.
Zoccia sessile or suberect, scattered, or in loosely connected
groups, united to one another by slender tubes originating from
the dorsal or lateral surfaces; aperture 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, Hineks, Jullien, and others, following the
usage of MacGillivray and Waters. Hincks (785) regards Dia-
choris of Busk as an artificial division but considers it to be
more nearly related to Bugula because of its boat-shaped zocecia
and articulated avicularia. These characters do not, however,
distinguish Bugula alone, whereas the peculiar mode of connec-
tion of the zocecia 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.
Pl. XII, figs. 63, 64, and text figure 1.
Beania mirabilis Johnston, 1847, p. 372, text figs. 69, 70.
Beania mirabilis, Busk, 1852, pt. I, p. 32, Pl. XXIV, figs. 4, 5.
Beania mirabilis, Smitt, 1867, pp. 295 and 357.
Beania mirabilis, Hincks, 1880, p. 95, Pl. 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). Zoawcia
VoL. 2] Robertson.—Non-Incrusting Bryozoa. 20%
boat-shaped (fig. 64, zoe.), having two erect spines (e. sp.) above
the mouth, and a variable number of marginal spines, Six, seven,
or eight on each side of the aperture; tubes (con. t.) connecting
the zoocia arising near the base of each zocecium laterally or on
the dorsal surface, 7.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 zowcium (br.). Owcia? 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 mirabilis is obtained in considerable quantity growing
over Alcyonaria and Bowerbankia at Dead Mans Island and
Santa Catalina Island.
25. Beania longispinosa sp. nov.
Pl. XII, figs. 65, 66.
Zoarium sub-erect, consisting of groups of zocecia united by
tubular connections (fig. 65). Zowcia 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 zocecia,
usually two on each side, uniting lateral zocecia, one at each end
uniting with a zocecium laterally and proximally. No avicu-
laria. Oacia (oe.) small, situated at the summit of the zocecia,
marked by parallel striae extending longitudinally. Rootlets very
long, extending from the dorsal surface of many of the zocecia,
sometimes 50 or more mm. in length (fig. 65, 7.).
This species seems to be related to Diachoris hyadesi 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 zocecium.
Beania longispinosa was obtained in several fathoms of water
off La Jolla, California, growing upon rock.
Stirparia Goldstein.
Stirparia Goldstein, 1879 or 1880.
Stirparia, Hincks, 1883.
Bicellaria (part), Busk, 1884.
Zoarium consisting of erect segmented stalks, bearing on their
summits feathery tufts whose branches consist of zocecia bise-
rially arranged. Zowcia of the normal Bicellarian or Bugulan
type, v.e., turbinate, with aperture 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
zocecia not turned inward; lower portion tubular. <Avicularia
and owcia 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 occia, of which Hincks
makes no mention. Ocecia are abundant on all the specimens
and are of the Bicellarian type, being pedunculate upon the inner
side of the zocecium near the top.
KEY TO SPECIES OF STIRPARIA.
1; Zomeia with) apertometurnem Wye) eee sce ees eee eee 2
i. A0ceela, with’ aperture mot) bummed inp wed skeen eee eee ee 3
2. Aperture occupying less than half the front, and armed with five
OES SUS PGS ee eee ce 8 ee ee ee eee ee eee S. ciliata .
2». Aperture occupying half or more of the front wall .....................-1...----.-
3. Margin of zoecia armed with two, three, or four spines .....................-
Se SSR og NOR ne ee ee See Rae S. occidentalis
3. Margin of zoecia armed with two spines on the upper outer angle......
SoBe Res i ie eens a he eek eR er es ean een, Lm S. californica
Ww
Vou, 2] Robertson.—Non-Incrusting Bryozoa.
ho
-—l
ie)
26. Stirparia ciliata, sp. nov.
Pl. XII, figs. 67, 68, 69; Pl. 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 zoccia, 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 zocecium of the branching
tufts similar to the distal segment (dis. seg.) of the stalk, being
often but little shorter, and but shghtly modified in other re-
spects from the ordinary stalk segments. This first zocecium tubu-
lar, elongated, possessing a circular aperture occupying about
one-third the length of the zocecium, and surrounded by spines;
from its anterior extremity two zocecia arise, and as growth
proceeds the zowcia that subsequently appear in the branch ac-
quire the typical form. Zowcia of the Bicellarian type, broadly
ovate above, tubular below, the aperture looking upward, the
tubular portion inclined inward (Pl. XIII, fig. 70). Aperture
oceupying 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 (b7. 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. Owcia (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 zocecium 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. ciliata is re-
lated to S. 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 S. annulata and for which it is
named. Many evidences are afforded of the zoccial origin of
these segments. Instances are found where a segment assumes
the shape externally of an ordinary zocecium, 7.e., it becomes
wider at the distal end and possesses an aperture on whose mar-
ein 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 numerous and slightly larger near 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.
Pl. 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 zocecia 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 zocwcial tuft being very sudden. Zocecial tufts relatively
short, not half as long as the main stalk, branching dichotomous.
Zoecia biserial, alternate; at the formation of a branch, the
series of zocecia separated by the interpolation of two new zocecia
on the inner side so that no one zocecium ean be said to be at
Vou. 2] Robertson.—Non-Incrusting Bryozoa. 281
the bifureation (fig. 74, b7.), 7.e., the branches bifureate between
two series of zocecia forming the proximal branch and not upon
a single zoecium. Zowcia (z@.) 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.) peduneculate, situated below and to one side of the aper-
ture; present usually on every zoccium. Oacia (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 S. 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 Pl. XIV,
fig. 76, is twice the length and thickness of those of the upper
portion of the same stalk represented in Pl. XIII, fig. 73.
This species is found in several localities on the coast of Cali-
fornia and Puget Sound.
28. Stirparia californica sp. nov.
Pl. XIII, fig. 75; Pl. XIV, figs. 77, 78, 79, 80.
Zoarium consisting of tall stalks surmounted by large, feath-
ery, branching tufts of zocecia (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 zowcial tufts. Segments of the stalk stout, some very long,
282 University of California Publications. [| ZooLocy
others short, showing a kind of alternation sometimes, and grow-
ing decidedly shorter toward the distal end: zoccial 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. Zowcia resembling those of Bugula,
especially of B. pacifica; elongated, only slightly narrower below
than above; aperture 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 at
the summit of the zocecia where each angle terminates in a long
jointed spine; from the distal margin of a zocecium a second
longer spine (sp.) appears, making 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 zoccium
just below the aperture. Oacia (oe.) globose, pedunculate, on
the inner side at the summit of each zoecium. fRootlets spring-
ing from lower segments of the branches of the stalk, passing
downward close to stalk and attaching themselves to basal disk.
(Pl. XIII, fig. 75, r.) |
But two species of Stirparia have been hitherto described, S.
glabra, Hineks (’83), and S. annulata, 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 8S. 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 VS. californica we find the boat-shaped
zocecium characteristic of Bugula. In all, the avicularia and
Vou. 2] Robertson.—Non-Incrusting Bryozoa. 283
ocecia 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 ealled to the fact that, as Hincks (’83) observed, the seg-
ments of the stalk are probably aborted zocecia. Their zocecial
nature is most plainly shown in NS. ciliata, in which as shown in
Pl. 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
eounted. This zocecium-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,
S. 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 S. 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 ealled to the difference in size
between the lower and upper segments of the stalk of S. occi-
dentaus (El wid, fie. 73> Pl. XPV, fic. 76). A similar differ:
ence exists in the segments of the stalk of S. 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 NS. 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 (7b.) which form in the ecto-
eyst of the lower portion of the stalk and extend to the cireum-
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 Pl. XIII, fig. 75, the stalk which forms the foot disk may
284 University of California Publications. [ ZooLocy
divide into a number of secondary stalks (b7.) each surmounted
by its zocecial tuft and sending down rootlets which attach them-
selves to the base. The colonies of S. californica grow in isolated
groups, thus differing from S. ciliata and SN. occidentalis, which
form tangled masses of stems springing from many interwoven
rootlets.
S. ciliata and S. occidentalis are shore forms, being found
between tides growing on sea weed, other bryozoa, ete. S. cali-
fornica, 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 Hincks.
Corynoporella Hincks, 1888.
Zoarium forming delicate white tufts, consisting of zocecia
disposed in a single series, and facing one way; dichotomously
branched. Zoacia more or less clavate, each originating from the
dorsal surface of the one beneath it, immediately below the sum-
mit; elongated, attenuated, tubular; aperture occupying a half
or less of the front surface. Avicularia articulated, attached to
the side of the aperture. Oacia globose.
29. Corynoporella spinosa sp. nov.
Pl. XIV, figs. 81, 82, 83.
Zoarium erect, much branched, each internode consisting of
a single zoccium (fig. 81); branches (fig. 82, br.) arising on
the dorsal surface near the summit, singly or in pairs. Zowcia
sub-clavate. narrowed below, widened above, the distal margin
truneate or only shghtly rounded; aperture membranous, occu-
pying a third or more often a half of the front surface; sur-
rounded by 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 (av.) 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. Owcia (fig. 83, oe.) globose, prominent, as.
VoL. 2] Robertson.—Non-Incrusting Bryozoa. 285
wide as the zowcial aperture, with lines radiating from the front
rim; few in number. Rootlets (fig. 82, r.) springing from a disk
on the side of most of the zocecia, just below where the branches
arise.
But one other species of this genus is described, viz., C. tenuis,
Hineks (’88) from the St. Lawrence. As Hincks remarks, ‘‘the
zocecia bear a strong resemblance to those of the genus Prettia,
but the articulated avicularium is a link connecting it with
Bugula.’’ The species C. spinosa has a decided resemblance to
Bugula in that its zoccia approximate the boat shape typical for
that genus. It seems also to bear a resemblance to Bugulella
Vernll (779), but in Corynoporella the method of branching as
well as the mode of union of the zocecia are distinctly different
from that in this genus.
Obtained from material from Alaska growing on a erab.
Exact locality unknown.
Notamiidae Hincks.
Gemellariade (part) Busk, 1852.
Notamiide Hincks, 1880.
Zoecia 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.
Notamia, Waters, 1885.
Synnotum Hincks, 1886.
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 zoecium. (Fig. 93.) Zowcia 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 zowcia in each pair
at the summit. Owcia?
286 University of California Publications. | ZOOLOGY
30. Synnotum aviculare (Pieper) Hincks.
Pl. XIV, figs. 84, 85.
Gemellaria avicularis Pieper, 1881, Vol. 1X, p. 43, Pl. I, 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. 6, 7.
Zoarium minute, slender, of delicate texture, glossy, simple
in habit, consisting of long branches which bifurcate at rather
distant intervals (fig. 84). Zoawcia attenuated below, expanded
above (fig. 85); aperture occupying more than two-thirds of the
front, narrowing below, the margin thin and unarmed; lateral
avicularia (lat. av.) small, sessile placed at the top of the zoc-
cium on the inner side, developed on every pair of zocecia, widen-
ing from the base upward with a minute beak; median avicularia
(me. av.) pedunculate, capitate, placed at the top of each pair
of zoccia in the median plane, subglobular, smooth, the beak
very slightly produced, with a sharp spike-like extremity.
The sessile avicularia in the. California specimens are not
alternate as Hincks (786) describes for those from the Adriatic.
but appear rather on every pair. he capitate avicularia are fre-
quently lacking in our colonies but this may be due to mutilation.
The rootlets (fig. 85, r.) frequently arise quite high up on a
branch from the front surface on the line midway between the
top and base of a pair of zocecia, sometimes even upon the aper-
ture of a zocecium.
S. aviculare occurs with other minute forms of bryozoa at
Dead Mans Island, San Pedro, and Ballast Point, San Diego,
California.
Cellariidae Hincks.
Escharide (part) Johnston, 1847.
Salicornariade Busk, 1852.
Ceilariew Smitt, 1867.
Cellartide Hincks, 1880.
Zoarium erect, calcareous, articulated; branching dichoto-
mous or non-dichotomous. Zoacia usually rhomboidal or hexag-
onal, disposed in series around an imaginary axis so as to form
eylindrical shoots.
Vot. 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. Zoawcia alternate, depressed in front, and surrounded by
a raised border. Avicularia immersed, irregularly distributed,
situated above a zocecium, or in place of one. Oacia immersed.
KEY TO SPECIES OF CELLARIA.
1. Branching dichotomous; avicularia with triangular mandible directed
LO syyamniyy Ts Cl ieee ys tec ee eae ae ee Ben ee ee eee C. borealis
OLEATE rear Lana LT Ca ETT U1 2
2. Internodes small, avicularia large, twice the size of an ordinary
zocecium, mandible rounded, directed upward........ Cc. mandibulata
2. Internodes long, few, avicularia small, mandible rounded, directed
UND EUR INGOT OMG WED Gl scorer re eee eee ee C. diffusa
831. Cellaria borealis (Busk) Smitt.
BI PXhVs fic. 86 BI xXeval fics 102:
Salicornaria borealis Busk, 1855, p. 254, Pl. I, figs. 1, 2, 3.
Cellaria borealis, Smitt, 1867, pp. 383 and 361, Pl. XX, fig. 17.
Zoarium growing in luxuriant masses 75-100 mm. in height
(Pl. XVI, fig. 102), branching dichotomous, except at the ex-
tremities where the internodes are frequently tipped with three
or four very small terminal internodes. IJnternodes of varying
length, about a centimeter on an average, club-shaped; joints
not black; young actively growing portions of the colony bright
flesh pink. Zowcia (Pl. 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 zoccia sepa-
rated by sinuous lines; front wall caleareous, and except upon
the operculum which is large and semicircular on the distal
edge, is marked by rather coarse punctures; orifice at summit
of zoecium. Avicularia (av.) above the zocecia distributed at
288 University of California Publications. [ ZooLocy
irregular intervals; mandible triangular, pointing downward.
Oawcia? 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. It is
abundant on the rocks at Orea, Prinee Williams Sound. It
seems to be a strictly northern species not being reported south
of Queen Charlotte Islands.
832. Cellaria mandibulata Hincks.
Pl. XV, figs. 87, 88, 89; Pl. XVI, fig. 103.
Cellaria mandibulata Hineks, 1882, p. 462.
Cellaria mandibulata Hincks, 1884, p. 203, Pl. TX, fig. 7.
Zoarium forming masses of delicate branches often growing
from 75-90 mm. in height (Pl. XVI, fig. 103). Internodes
slender. short, the longest from 6 to 8 mm. in length. Branch-
ing irregular, branches usually arising near the top of an inter-
node, but often from the middle, or from any point on its sur-
face; when arising from the top of an internode, two branches
are seldom opposite, the one usually being higher than the other.
Joints black. Zoacia 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 (Pl. XV,
fig. 87) ; in older stages they are somewhat wider, truncate above
and below (fig. 88); in the oldest stage, zocecia 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
ecrenulate 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 zoccia
(fig. 87); in older stages, the sinuous border becomes united
above and below the zocecia, and the crenulations are less marked
(fig. 89). Zocecial 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
Vor. 2] Robertson.—Non-Incrusting 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 lke
extra large zocecia; much broader than ordinary zocecia, 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 occium (or.) Oa@cia not
found in the youngest zocecial stage described above. In older
stages the orifice of the ocecium (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
zocecia of the colony.
Without doubt this is the species described by Hinecks 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. nov.
Pl. XV, fig. 90; Pl. XVI, fig. 104.
Zoarium consisting of a relatively small number of rather
long cylindrical internodes connected by dark chitinous joints;
imternodes varying in length, the longest attaining a leneth of
39 or 40 mm., the whole colony often 75 mm. high, and having
a strageling 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 zoccium; two branches, some-
times three or four arising at the distal end of the internode
290 University of California Publications. [ ZooLocy
(Pl. XVI, fig. 104). Joints black. Internodes club-shaped, stout.
Zoecia depressed, narrower below and above than in the middle,
truneate at each end; front wall caleareous, covered with minute
protuberances. (Pl. XV, fig. 90.) Operculum (op.) some dis-
tance below the top of the zocecium, lower edge slightly curved,
bearing a denticle (d.) near each corner, upper edge semicir-
cular. Oacial orifice (oe. or.) situated just above the operculum
and resembling it in shape; a broad mucro (m.), a continuation
of the caleareous zocecial wall in the middle of its lower edge.
Avicularium (av.) in place of a zocecium, almost square. Root-
lets springing from the lower zoccia of the proximal internode,
passing down close to the wall of the internode, then spreading
out disk-like around the base of the colony.
In its habit 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. australis and C. mandibulata which it
resembles in habit of growth and method of branching, always
from the middle of a zoccium. The zowcia resemble those of
C. australis in shape, but the position and shape of opercula
and occia 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 San Juan Island, and in
Port Orchard Channel, Puget Sound. It has also been dredged
both at San Pedro and San Diego.
Flustridae Smitt.
Escharide (part), Johnston, 1847.
Flustride (part), d’Orbigny, 1850.
Flustrade (part), Busk, 1852.
Flustride Smitt, 1867.
Flustride, Hincks, 1880.
Zoarium corneous and flexible, expanded, foliaceous, erect
or sub-erect. Zowcia contiguous, multiserial. Avicularia usually
of a simple type.
Vou. 2] Robertson.—Non-Incrusting Bryozoa. 291
Flustra Linneus.
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. Zowcia 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. Oacia immersed or raised.
34. Flustra lichenoides Robertson.
Pi XV, figs. 91, O25-Pi VE, figs. 105.
Flustra lichenoides Robertson, 1900, p. 322, Pl. XX, figs. 7, 7a, 8.
Zoarium unilaminar, consisting of broad foliaceous green or
brownish green fronds. (Pl. XVI, fig. 105.) Zowcia 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 eurving
inward, often meeting in the middle above the aperture ; aperture
occupying the whole front. Oacia (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
masses.
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
lamine 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-truncata Smitt, 1867, p. 358, Pl. XX,
figs. 1-5.
Zoarium composed of a number of erect, unilaminate fronds
(fig. 93). Zoecia irregularly quadrangular, truncate above and
below, often narrowed below (fig. 94); armed with a delicate
spine (sp.) at the distal corners; aperture occupying the whole
of the front; operculum (op.) semicireular. Avicularia (av.)
sparingly developed, in place of a zocecium; mandible semicir-
cular, directed upward. Oacia (oe.) small, immersed, not quite
as wide as the zocwcium against which each projects.
This appears to be the species which Smitt (67) describes
and figures, and which he finds in Arctic Seas growing on ascid-
ians, sertularians, ete., either creeping or erect. The material
in this collection consists of a few small erect fronds obtained at
Pribilof Islands, Bering Sea.
University of California,
Berkeley, February 15, 1905.
VoL. 2] Robertson.—Non-Incrusting Bryozoa. 293
LITERATURE CITED.
Allman, J. G.
1856. A Monograph on British Fresh-Water Polyzoa. Ray Society, Lon-
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-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, Pls. I-LX VIII.
Busk, G.
1855. Zoophytology. Quart. Journ. Micr. Science, Vol. III, pp. 253-256
and 320-322, Pls. I-IV.
Busk, G. ~
1881. List of Polyzoa collected by Capt. H. W. Feilden in the North-
Polar Expedition, &. Journ. Linn. Soec., Vol. XV, pp. 231-
241, Pl. I.
Busk, G.
1884. Report on the Polyzoa collected by H. M. S. ‘‘Challenger,’’ during
the years 1873-76. The Chetlostomata, in voyage of H. M. 8.
““Challenger,’’ Zoology X, pp. 4-216, Pls. 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. Sci.,
Philadelphia, p. 441.
‘Coppin, J.
1848. Description of a new genus of British Marine Zoophytes belonging
to the family Eucratiade. Ann. Nat. Hist., (2), Vol. IT,
pp. 273-274, Pl. X.
Ehrenberg, C. G.
1831. Symbole Physice.
Ellis, J.
1767. Versuch einer Natur-Geschichte der Corall-Arten. Nurnberg. Pp.
168, Pls. I-XLVI.
Ellis, J.
1767. An Account of the Actinia sociata, &. Phil. Trans., Vol. LVII,
pp. 428-436, Pl. I.
Ellis and Solander.
1786. A Natural History of the Zoophytes. Pp. 208, Pls. 63.
294 University of California Publications. [ ZOOLOGY
Farre, A.
1837. On the Minute Structure of some Polypi, &e. Phil. Trans., pp
387-426, Pls. 8.
Fewkes, J. W.
1889. New Invertebrata from the Coast of California. Bull. Essex
Institute, XXI, pp. 99-146, Pls. 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. Il, pp. 111-179, Pls. 19-21.
Goldstein, J. R. Y.
1879. New species of Polyzoa. Quart. Journ. Mier. Soe., Victoria. Vol. J.
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, Pls. VI-X.
Hincks, Th.
1880. A History of the British Marine Polyzoa. 2 vols. Vol. I, pp.
CXLI, 601, wood cuts 86; Vol. II, Pls. 83.
Hincks, Th.
1882. Polyzoa of Queen Charlotte Islands. Ann. Nat. Hist., (5) Vol. X,
pp. 248-256.
Hincks, Th.
1883. Contributions toward a General History of the Marine Polyzoa.
Ann. Nat. Hist. (5), Vol. XI, pp. 193-202, Pls. 2.
‘incks, Th.
1884. Polyzoa of Queen Charlotte Islands. Ann. Nat. Hist. (5), Vol.
13, pp. 203-215, Pl. I.
Hincks, Th.
1884. Contributions toward a General History of the Marine Polyzoa.
Ann. Nat. Hist., (5), Vol. XIII, pp. 356-369, Pls. 2.
Hincks, Th.
1885. Contributions toward a General History of the Marine Polyzoa.
Ann. Nat. Hist., (6), Vol. XV, pp. 244-257, Pls. VIT-IX.
Hincks, Th.
1888. The Polyzoa of the St. Lawrence: A Study of Arctic Forms. Ann.
Nat. Hist. (6) Vol. I, pp. 214-227, Pls. XIV and XV.
Jelly, E. C.
1889. A Synonymic Catalogue of the Recent Marine Bryozoa including
Fossil Synonyms. London. Pp. 322.
VoL. 2] Robertson.—Non-Incrusting Bryozoa. 295
Johnston, G.
1847. British Zoophytes. 2 vols. Vol. I, pp. 251-406; Vol. II, Pls.
XLVI-LXXIV.
Jullien, J.
1888. Mission Scientifique du Cap Horn. T. VI. Zoologie, Bryozoaires.
Pp. 924 Pls. 15:
Lamouroux, J. V.
1812. Extrait d’un memoire sur la classification des polypiers coralli-
génes. Bull. des sci. pour la Soc. Philomatique.
Lamouroux, J. V.
1816. Hist. Nat. des Polypiers coralligénes flexibles.
Linneus, L.
1758. Systema nature. 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. Mier. Sei., Victoria. Vol.
IE; jo: Ie
Nitsche, H.
1869. Beitrige zur Kenntniss d. Bryozoens. Zeitschr. wiss. Zool., Vol.
XX, pp. 1-36, Pls. 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, Pls. 3.
Oken, L.
1815. Lehrbuch der Naturgeschichte. Abt. 2.
Orbigny, A. d’
1850. Paléontologie Francaise-Terrains crétacés. Vol. V, Bryozoaires.
Text, pp. 1191; atlas, Pls. 800.
Pallas, P. 8S.
1766. Elenchus Zoophytorum. Pp. 451.
Pieper, F. W.
1881. Hine neue Bryozoe der Adria, Gemellaria (?) avicularis. Jahresber.
Westfalischen Prov.-Vereins, Vol. IX, pp. 43-48, Pl. I.
Robertson, A.
1899. Studies in the Pacific Coast Entoprocts. Proc. Cal. Acad. Sei.,
(3), Vol. Il, pp. 323-348, PI. I.
296 University of California Publications. [ ZOOLOGY
Robertson, A.
1900. Papers from the Harriman Alaska Expedition. The Bryozoa..
Proe. Wash. Acad. Sci., Vol. II, pp. 315-335, Pls. XIX-XXI.
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1811. Iconographie des Zoophytes de 1’Egypte, from the ‘Description de
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1867. Kritisk Férteckning 6fver Skandinaviens Hafs-Bryozoen. Ofv. K.
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1871-72. Floridan Bryozoa. Kgl. Svenska Vet.-Akad. Handlingar., Vol.
X, pp. 20, Pls. 5.
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1857. On some new Microscopic Organisms. Proc. Cal. Acad. Sci., Vol.
I-II, pp. 110-115, Pls. 2.
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Waters, A. W.
1879. Bryozoa of the Bay of Naples. Ann. Nat. Hist. (5) Vol. III,
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(2), Vol. 5, pp: 174-779; Pl XT:
Waters, A. W.
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— a. eee
Vou. 2] Robertson.—Non-Incrusting Bryozoa.
ABBREVIATIONS IN THE PLATES.
a.—area.
ab. ap.—aborted aperture.
ad.—adnate part of zocecium.
ap.—aperture.
b. sp.—basal spine.
b. f.—binding fibre.
bi.—bifureation.
bi. av.—avicularium at bifurcating
zocecium.
bi. sc.—hbifid seutum.
bi. sp.—hifid spine.
bk.—beak.
br.—branch.
c. av.—eurved avicularium.
cr. Sp.—erossed spine.
¢. Sp.—eurved spine.
car. val.
con. tu.—connecting tube.
cr. bor.—erenulate border.
cr. br.—creeping branch.
cardiac valve.
d.—denticle.
dis. ze.—distal zoccium.
e. br.—erect branch._
€. Sp.—erect spine.
fr. av.—frontal avicularium.
fr. ze.-—frontal zocecium.
fst. ze.—first zocecium.
giz.— gizzard.
gv.— groove.
i.—intestine.
in. sp.—inner spine.
int.—internode.
j.—Joint.
k.—knob.
lat. av.—lateral avicularium.
lat. z@.—lateral zocecium.
m.—mouth.
man.—mandible.
me. av.—median avicularium.
mem. ap._—membranous aperture.
mus.—muscle.
o.—ovum.
oe.—ocecium.
oe. or.—oeecial orifice.
@s.—cesophagus.
op.—operculum.
or.—orifice.
ou. Sp.—outer spine.
ov.—ovary.
p-——pore.
par. mus.—parieto-vaginal muscle.
pd.—polypide.
ped. av.—peduneulate avicularium.
ph.—pharynx.
pr. by.—primary branch.
py. val.—pyloric valve.
r.—yrootlet.
rb.—trib.
r. ch.—root chamber.
r. {b.— root fibre.
re. mus.——retractor muscle.
re. sh. mus.—vetractile sheath muscle.
re. pd.—regenerated polypide.
s. h.snake’s head.
se.—seutum.
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 zocecium.
tu.—tubular part of the zocecium.
u. r.—upward extending rootlet.
v.—vibraculum.
v. c.—vibracular cell.
v. ch.—vibracular chamber.
2e.—zocecium.
z2@. bi.—zoecium at bifureation.
ze. br.—zoecial branch.
zZ@. or.—zocecial orifice.
PLATE IV.
Fig. 1—Aetea anguina Linneus. Habit 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 zomcia. Adherent
(ad.) portion attached to the substratum like a stolon is part of the zowcium
which rises at its anterior end into the ringed tubular portion (tu.). The
polypide (pd.) can be seen inside several of the zowcia; when fully ex-
panded it protrudes through the operculum (op.) at the summit of the tube.
The zowcia are separated from each other by a septum (sep.) at the anterior
end of each. X 30.
Fig. 3—A. anguina. Part of a zowcium in which the polypide (pd.)
is in a state of retraction; upper part of tubular portion broken
away. Polypide retracted by contraction of retractor muscle fibres (mus.)
attached to the wall of the adherent portion (ad.) of the zowcium. In the
adherent portion the ovary (ov.) is visible. > 70.
Fig. 4.—A. anguina. A single zoecium much enlarged showing position
of owcium (0e.) just outside the aperture (ap.), below the operculum on
the ventral side of the zowcium. Within the oecium is an ovum in early
cleavage stage (emb.). X 70.
Fig. 5.—Aetea truncata Landsborough. Portion of a colony enlarged to
show variations in height of the erect, tubular portions (tu.), also length
and slenderness of the adherent portions (ad.) of the zowcia. X 30.
Fig. 6.—A. truncata. Habit sketch, about natural size.
2981
UNIV. CAL.PUBL.ZO0L.VoOL.II.
ROBERTSON DEL
FHOTO-LIE LGB SF
PLATE: V.
Fig. 7.—Eucratea chelata Linneus. Habit sketch, about natural size.
Fig. 8.—E. chelata. Enlarged to show zowcia and method of branching
(e. br.) just below the aperture (ap.), and the creeping branches (cr. br.)
given off from the sides of the zowcia. X 50.
Fig. 9.—£. chelata. Showing oecium (0e.) on a small aborted zowcium.
(After Hincks.) X 50.
Fig. 10.— Gemellaria loricata Linneus. Habit sketch, natural size.
Fig. 11.—G. lcricata. Portion of a branch enlarged to show arrange-
ment of zowcia and method of branching. X 30.
Fig. 12.—Menipea ternata Ellis and Solander. Habit sketch of speci-
men from Puget Sound.
Fig. 13.—M. ternata. Habit sketch of specimen from California.
Fig. 14.—M. ternata. Enlarged portion of colony from Puget Sound
showing zowcia in groups of three, except the omcial internodes (oe.). Lat
eral avicularia (av.) well developed. X 30.
M. ternata. Dorsal view of the zocwcium at the bifurcation of
a branch (bi. br.) showing the adjoining zoecia (lat. z@.) and root chamber
(r. ch.) with upward extending rootlet (r.). X 30.
Fig. 15.
Tig. 16.—M. ternata. A single zowcium to show the position of the two
kinds of root fibres, those anchoring the colony (r.), arising in a simple
root disk (7. d.) on the front wall of the zowcium; those extending upward
(wu. r.), arising from a rather large chamber (1. ch.) projecting from the
zowcial wall just above the lateral avicularium. X 30.
[300]
ih in ii i i i lt
| ROBERTSON
Uwnvv. CaL.PUBL.ZOOL.VOLII.
FHOTO-LUOH-L.G SF
ROBERTSON. DEL
PLATE VI.
Tig. 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.
Fig. 19—M. gracilis. Portion of a branch enlarged to same scale as
figs. 14 and 17, showing the greater length of the zoecia, the large flabellate
scutum (se.), the relatively small size of the avicularia (av.), and the shape
and position of the two sorts of root chambers (r. d. and r. ch.) similar to
those of M. ternata. X 30.
Fig. 20.—M. gracilis. Portion of an ocial internode showing develop-
ment of avicularia (av.) on the front of the zocwcium close to the owcia
(oe:).) 30:
Fig. 21.—M. gracilis. Dorsal view of the zoecium at the bifurcation of
a branch (bi. z@.), showing the lateral adjoining zowcia (lat. z@.), and
root chamber (r. ch.) with upward extending rootlet (wu. r.). X 30.
Fig. 22.—M. occidentalis Trask. Habit sketch of a single branch to
show method of branching. Somewhat diagrammatic.
Vig. 23.—M. occidentalis, A few internodes enlarged to show size of
avicularia (av.), 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
figs. 14, 17, and 19. X 30.
Fig. 24.—M. occidentalis. An oecial (0e.) internode consisting of five
zocecia. X 30.
Fig. 25.—M. occidentalis. Lateral view of a zoecium, showing a root
chamber (7. ch.) from which there extends a downward extending rootlet
(Gena, 2 clk
[ RoBeRTsoN| PLATE VI
Univ Cal. POBL.Zo0L. VoL. 1].
FHOTU-LITELG SF
ROBERTSON, DEL
PLATE VII.
Fig. 26.—M. occidentalis catalinensis, subsp. noy. Portion of a branch
showing finger-like processes on scutum (sc.), and bifid spines (¢. sp.). * 50.
Fig. 27.—M. occidentalis catalinensis. Lateral view of a zoecium show-
ing root chamber (r ch.) and downward extending root fibre (7.).
Fig. 28.—Menipea erecta Robertson. Habit sketch, natural size.
Fig. 29.—M. erecta. Enlarged portion of a colony from Kadiak show-
ing lateral avicularia (lat. av.), frontal avicularia (fr. av.), and spines
(sp.). X 30.
Fig. 30.—M. erecta. Enlarged portion of a colony from Sitka showing
absence of lateral avicularia (lat. av.); showing also rootlet (7.) springing
directly from zoccial wall. X 30.
Fig. 31.—WM. erecta. Enlarged portion of colony from San Juan showing
absence of spines. 30.
Fig. 32.—Menipea pribilofi, sp. nov. Habit sketch, natural size.
Fig. 33.—M. pribilofi. Occial internode (0e.). X 30.
[304]
Univ. CaL.PUBL.Zo0L.VoLII. | ROBERTSON | FLaTe VII
ROBERTSON, DEL.
PLATE VIII.
Fig. 34.—M. pribilofi. Portion of a branch enlarged showing number
and length of jointed spines (sp.), size of lateral avicularia (lat. av.),
frontal avicularia (fr. av.), and root chambers in the two positions (7. ch.
and: 7 1ds) in e< 80:
Fig. 35.—Scrupocellaria californica Trask. Dorsal view of part of two
zoecia to show vibracular chamber (v. ch.), short vibraculum (v.), groove
of vibraculum (gr.), and lateral avicularium below (av.). x 50.
Fig. 36.—S. californica. Portion of a branch enlarged showing varia-
tion in size of lateral avicularia (lat. av.), the vibracular cell (v. ¢.), on
zowcium at bifurcation (zoe. bi.), visible from the front, while on other
zocecia only the short vibraculum (v.) is seen from the front, and only on
the lower zoccia which possess root fibres (r.). X 50.
Fig. 36a.—S. californica. Habit sketch.
Fig. 37.—S. californica. Three zoccia with owcia (0e.). X 50.
Fig. 38.—Scrupocellaria varians Hincks. 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 (v. ch.), and the long vibraculum (v.) present on
each zoecium. X 20.
Fig. 39.—S. varians. Dorsal view of a portion of two zoecia to show
position of vibracular chamber (v. ch.) with its long vibraculum (v.) and
its groove (gr.), with the root disk (7. d.) and root fibre (7. f.) on its
basal margin; also lateral avicularium with its much elongated curved beak
(c. av.). X 20.
Fig. 40.—Caberea ellisi Fleming. Habit sketch.
{306] -
{
oe ae |
ebre yy
Univ, CaL.PUBL.ZOO0L.VOL.II. | ROBERTSON] PLaTe VIII.
ROBERTSON. DEL
FAOTO-LITALG SP
a . 7 , 7 . ; en
PLATE IX.
Fig. 41.—S. diegensis sp. nov. Portion of a branch enlarged showing
form of zowcia (z@.) with large scutum (se.), spines on the upper margin
of the aperture (ap.) one or more of which are bifid (bi. sp.) ; also lateral
avicularium (lat. av.) and frontal avicularium on the zowcium at bifurcation
(bi. av.). On those zowcia not possessing owcia the vibracular chamber
(v. ch.) plainly visible with its long vibraculum (v.). 50.
Fig. 42.—S. diegensis. Dorsal view of a few zoewcia 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
zocecia. At the base of the vibracular chamber and to one side of the groove
is the pore (p.) from which the root fibre extends (7r.). X 50.
Fig. 43.—S. diegensis. A vibracular chamber enlarged to show detail.
Fig. 44.--S. diegensis. The avicularium on zoecium at bifurcation en-
larged to show its large muscular portion (mus.) and the obliquely directed
mandible (man.).
Fig. 45.—Caberea ellisi Fleming. A few zocecia enlarged. Vibracular
chamber visible from the front (v. ch.); the minute lateral avicularia (lat.
av.); serrated vibraculum (v.). X 50.
Fig. 46.—C. ellisi. Dorsal view of a few zocwcia to show vibracular
(v. ch.) chambers extending obliquely across the back of each zoccium,
the groove (gr.), and the long vibraculum (v.); also the rootlets (7.) 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 zowcia (z@.) with large aperture (ap.), margins of which termi-
rate in denticles (d.); also pedunculated owcium (0e.). X 30.
[308]
| ROBERTSON] FLaTe IX
Univ. Cau.Fush.Zo0L. Vo...
FHOTO-LITH LG SF
ROBERTSON. DEL.
PLATE X.
Fig. 48.—Bugula murrayana Johnston. Portion of a branch enlarged
showing zoecia (z@.) with margins of aperture armed with variable number
of spines (sp.); avicularium (av.) or spine (sp.) at base of aperture. Mar-
ginal avicularia lacking on this specimen. X 30.
Fig. 49.—B. californica sp. nov. Portion of a branch enlarged to show
spines terminating the upper margins of the aperture (sp.),long outer spine
(ou. sp.), and position of avicularia (av.). 50.
Fig. 50.—B. pacifica sp. nov. Portion of a branch enlarged to show
zoecia (z@.), spines (sp. and sp.’), and the shallow owcia (0e.); also the
embryo (emb.) in the upper part of the zoecium. X 50.
Fig. 51—Bugula flabellata Thompson. Habit sketch, natural size.
Fig. 52.—Bugula flabellata. Portion of a colony enlarged to show ar-
rangement and branching of zoccia, the spines (sp.), avicularia (av.), and
owcia (0e.). X 30.
Fig. 53.—Bugula pugeti sp. nov. Habit sketch.
Fig. 54.—B. pugeti. Zoecia with lorg aperture (ap.), marginal avicu-
laria (av.), the three spines on the marginal zowcia (out. sp.), and the
process on the distal margin (k.). X 30.
[310]
ls
BL.ZOUL.VoL.!
UNIV. CaL.Pu
FHOTO -LITEL SSF
ROBERTSON,DEL.
PLATE XI.
Fig. 55.—B. pugeti. A few zoecia of lower part of a branch enlarged
to show the embryo (emb.) in the upper part of the zoccium, and the
brown body (b. b.) in each zoecium containing an embryo; a regenerating
polypide (re. pd.) in one zoecium. X 30.
Fig. 56.—Bugula curvirostrata sp. nov. Habit sketch, about natural size.
Fig. 57.—B. curvirostrata. Portion of a branch enlarged to show zowcia
and the two sorts of avicularia (av. and av’.). X 30.
Pig. 58.—B. curvirostrata. Portion of a branch showing three zowcia
with omcia (0e.). xX 380.
Fig. 59.—Bugula longirostrata sp. nov. Habit sketch, about natural size.
Fig. 60.—B. longirostrata. Portion of a branch showing attenuation of
zowcia, and avicularia (av.), and the shallowness of the owcia (0e.) which
only partially cover the embryos (emb.). X 50.
[312]
[RopeRTsoN| PLATE XI
Univ. CaL.FUBL. ZO Be aN Ge tley
ty
4
A
E
es
e
8
ab
ROBERTSON DEL
PLATE XII.
Fig. 61.—Bugula laxa sp. nov. Habit sketch.
Tig. 62.—B,. laxa. Portion of a branch enlarged showing form of zoc-
cia, marginal spines (sp.) and oecia (0e.). x 50.
Fig. 638.—Beania mirabilis Johnston. Habit sketch.
Fig. 64.—B. mirabilis. A few zowcia enlarged to show mode of connec-
tion (con. t.), the erect and decumbent portions, the former with margins
armed with spines. 50.
Fig. 65.—Beania longispinosa sp. nov. Habit sketch.
Fig. 66.—B. longispinosa. Portion of a colony enlarged to show char-
acter of zomcia, spines, and the owcia (0e.). 30.
Fig. 67.—Stirparia ciliata sp. nov. Habit sketch.
Fig. 68.—S. ciliata. Portion of the stalk enlarged showing segmentation
(seg.), also the beginning of the zowcial branches (z@. br.). x 50.
Fig. 69.—S. ciliata. Lower segment of stalk. X 50.
[314]
UNIV. CAL.PUBL.ZOoL.VOLII. | ROBERTSON | FLaTe XII
ROBERTSONDEL PHUTO-LITA.L.G SF
PLATE XIII.
Fig. 70.—S. ciliata. Portion of zowcial branch enlarged to show shape
and arrangement of the zowcia (z@.), the curved marginal spines (c. sp.),
and the minute avicularia. X 50,
Pig. 71.—S. ciliata. A few zowcia enlarged to show pedunculate owcia
(oe.). X 50.
Pig. 72.—Stirparia occidentalis sp. nov. Habit sketch, natural size.
Pig. 73.—S. occidentalis. Upper portion of the stalk and a few,of the
lower zo@cia, showing a specialization of the segments (seg.) of the stalk,
their articulation (art.), and the sudden transition into zowcia where the
zoecial tufts begin (fst. z@.). XX 50.
Fig. 74.—S. occidentalis. Portion of a branch to show character of
zoecia (z@.), of avicularia (av.), spines (sp.), and omecia (0e.). XX 50.
Fig. 75.—Stirparia californica sp. nov. Habit sketch, natural size.
[316]
| ROBERTSON] FLaTe XIII
UNIV. CaL.PUBL.ZOOL.VOL.I.
ye et A
Lex OH
Vg gis bys
LD LLP AR
Lyf i,
g Lp “4 A
Le $
A f
FHOTOLITHL.S SF
ROBERTSON, DEL
= le
i: Beis Ey
ae a Ls b te |
PLATE XIV.
Fig. 76.—Stirparia occidentalis, sp. nov. One of the lower segments of
the stalk drawn to the same seale as fig. 73, Pl. XIII, to show difference
in length of segments in the upper and lower parts of the stalk. X 50.
Fig. 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. X 50.
Fig. 78.—S. californica. One of the longer segments of the stalk en-
larged to the same scale as fig. 76. X 50.
Fig. 79.—S. californica. One of the shorter segments of the stalk en-
larged to the same scale as the preceding figure. X 50.
Fig. 80.—S. californica. Portion of a branch enlarged showing Bugulan
character of the zocwcia, position of avicularia (av.), and the Bicellarian
like owcia (0e.). X 50.
Fig. 81.—Corynoporella spinosa, sp. nov. Habit sketch.
Fig. 82.—C. spinosa. A few zowcia enlarged showing method of branch-
ing, position of avicularia (av.), of spines (sp.), and rootlets (7.). X 50.
Fig. 83.—C. spinosa. <A single zowcium with owcium (0€.). X 50.
Fig. 84.—Synnotum aviculare Pieper. Habit sketch.
Fig. 85.—S. aviculare. Portion of branch enlarged showing character
of zoecia, root fibres (r.), of sessile (s. av.), and pedunculate avicularia
(ped. av.). X50.
Fig. 86.—Cellaria borealis Busk. A few zowcia enlarged to show their
arrangement, the avicularia (av.) just above the operculum (op.) sur-
rounded by the crenulate border (cr. bor.) similar to that separating the
lines of zowcia. X 50.
[318]
<4 ho —> Qt ieee
Univ. CaL.PuB.Zoo1 Vou.IL. | ROBERTSON] FLATE XIV
ve aH wits oad °
FHOTU-LITHL& SP
ROBERTSON, DEL
o
MTA:
=
; 7
—
oy
im
PLATE XV.
Fig. 87.—Cellaria mandibulata Hincks. Two zoecia, young stage show-
ing lateral crenulated border (er. bd.).
Fig. 88.—C. mandibulata. Several zowcia at an older stage, showing
operculum (op.) with its four denticles (d.), the immersed owcia 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 he-
low the zoccia, thus surrounding them. X 50.
Fig. 89.—C. mandibulata. Three zowcia at a still older stage to show
inerease of caleareous wall and final form of omecial orifice (0e. or.) re-
sembling a small avicularium. X 50.
Fig. 90.—Cellaria diffusa sp. nov. A few zowcia (¢@.) to show shape,
oecial orifice (oe. or.) with the lower margin developed into a mucro (m.),
and the avicularia (av.). X 50.
Fig. 91—Flustra lichenoides Robertson. A few zocwcia enlarged to show
erect spines (é. sp.), operculum (op.), and omcia (0e.). X 50.
Fig. 92.—F. lichenoides. The dorsal surface of a few zowcia showing
rcot fibres springing from the distal angle of each zowcium.
lig. 93.—Flustra membranaceo-truncata Smitt. Habit sketch.
Fig. 94.—F. membranaceo-truncata. A few zoecia showing arrange-
ment, immersed oewcia (0e.), and avicularium of simple structure in place of
zoecium (av.). X 50.
[320]
Univ. CAL.PuBL.ZO0L.VoLII. | ROBERTSON] PLATE XV
PHOTOLITALE SR
ROBBRTSON DEL.
Photographs to show habit of growth.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
o
PLATE XVI.
95.—Scerupocellaria varians Hincks. —
96.—Scrupocellaria diegensis Robertson. ane <n
97.—Bugula neritina (Linneus) Oken. a la :
98.—Bugula murrayana Johnston. Specimen from Orea, Al ska. my; ‘ |
99.—Bugula murrayana Johnston. Specimen from Puget Sound.
100.—Bugula californica Robertson. ;
101.—Bugula pacifica Robertson.
102.—Cellaria borealis (Rusk) Hincks.
103.—Cellaria mandibulata Hincks.
104.—Cellaria diffusa Robertson.
105.—Flustra lichenoides Robertson.
[322]
Unt. CaL.PuBL.ZooL.VOL.IL. [ ROBERTSON] Flare XVI
J.M_W. Photo.
UNIVERSITY OF CALIFORNIA PUBLICATIONS
; ZOOLOGY
Vol. 2, Nos. 6 and 7, pp. Bgss10. os" = Becdater 13, 1905
CONTRIBUTIONS FROM THE LABORATORY.
2 = OF THE... =~
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oe | vI tee
DIFFERENTIATION IN HYDROID COLONIES —
AND THE PROBLEM OF SENESCENCE
2 | i ot ee oc ae
- THE BEHAVIOR OF CORYMORPHA
BY =>
HARRY BEAL TORREY
_. BERKELEY.
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Vb
DIFFERENTIATION IN HYDROID COLONIES
AND THE PROBLEM OF SENESCENCE.
BY
HARRY BEAL TORREY.
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-
theeal pedicels, size, proportions and.ornamentation of hydro-
theeae 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'
(Torrey, :04, p. 16). The stems, usually unbranched, rise from
a ereeping hydrorhiza to the height of 20 or 30 mm. For a
few millimeters (2 to 5) each stem is free of polyps and is ecom-
pletely annulated. The annuli, short at the base, increase grad-
ually in length until they become, at the first hydranth pedicel,
1The Hydroids of the San Diego Region. Univ. Cal. Publ. Zoology, I,
No. 1, 1904.
324 University of California Publications. [ ZcoLoGy
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 to one hydrotheeal pedicel. The pedicels themselves vary in
length and annulation from the base of the stem toward the tip.
The first pedicel may exhibit 4 to 7 annul. The last hydro-
theca may possess no pedicel at all, or one of but a single an-
nulus. The number to a pedicel varies with much general uni-
formity from one of these extremes to the other.
From base to tip, then, each stem presents two main serial
changes. (1) Below the hydranth region, the cauline annuli
erow progressively longer. (2) Within the hydranth region, the
number of annuli to a pedicel progressively decreases. There
is a strong suggestion in these facts that they depend on changes
in the internal physiological conditions of growth. Since polyps
are readily regenerated where stems are sectioned, a method at
once presents itself for determining the differentiation at 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 at 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 cut 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 annulus, the number
characteristic of the distal region ?
The facts obtained from the experiments fall into three cate-
gories, according as the regeneration followed (1) a cut through
the distal portion of the stem, (2) a cut through the middle por-
tion, (3) heteromorphically from the cut basal end.
In all, 16 operations were performed. Of these, 7 were unsue-
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 26, No. 2. Stem with 24 hydranths and terminal
bud. Enumerating from base to tip,
\Vou.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 18 with 0 annuli
Pedicel 23 with 0 annuli
Pedicel 24 with 1 annulus
10am. Cut off terminal bud, 24, 23, and part of 22.
July 28,10 a.m. New hydranth No. 22 regenerated within old
eup, 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.
Il.—July 26, No. 3. Stem with 17 hydranths and terminal
bud.
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 0 annuli
Pedicel 13 with 0 annuli
Pedicel 15 with 0 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.—Aueust 5, No. 4. Stem with 15 hydranths and terminal
bud.
Pedicels 1-4 with 2 annuli
Pedicels 5-11 with 1 annulus
Pedicel 12 with 0 annuli
Pedicel 13 with 3 annuli
Pedicels 14-15 with 1 annulus
Pedicel 16 with 0 annuli
326 University of California Publications. [ ZooLocy
8:45 a.m. Removed terminal bud, 15, 14, and 13.
August 7, 9:45 a.m. Nos. 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. Stem with 17 hydranths.
Pedicel 1 with 8 annuli
Pedicel 2 with 6 annuli
Pedicel 3 with 4 annuli
Pedicel 4 with 3 annuli 3
Pedicel 5 with 2 annuli
Pedicel 6 with 4 annuli
Pedicel 7 with 3 annuli
Pedicels 8-9 with 2 annuli
Pedicels 10-11 with 3 annuli
Pedicels 12-15 with 2 annuli
Pedicel 16 with 1 annulus
Pedicel 17 with 0 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
bud.
Pedicel 1 with 6 annuli
Pedicel 2 with 4 annuli
Pedicel 3 with 5 annuli
Pedicel 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 9 with 3 annuli
Pedicel 10 with 2 annuli
Pedicels 11-14 with 1 annulus
8:55 a.m. Removed terminal bud, 13 and 12.
Vou. 2] Torrey.—Hydroid Differentiation and Senescence. pal
August 8. Nos. 12 and 13 regenerated, each with 1 annulus in
pedicel.
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. [ ZooLocy
clusively that, under the conditions of the experiments’ regen-
eration from cuts through the distal region of the stem produces
structures of the local, 2.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
bud.
Pedicel 1 with 5 annuli
Pedicels 2—5 with 4 annuli
Pedicel 6 with 6 annuli
Pedicel 7 with 3 annuli
Pedicel 8 with 5 annuli
Pedicel 9 with 2 annuli
Pedicel 10 with 4 annuli
Pedicel 11 with 1
Pedicel 12 with 2
Pedicel 13 with 1
Pedicel 14 with 3 annuli
Pedicel 15 with 1 annulus
9
1
1
annulus
annuli
annulus
Pedicel 16 with
Pedicel 17 with
Pedicel 18 with
annuli
annulus
annulus
10 am. Cut stem between 6 and 7.
August 8, 3:30 p.m. No. 7 has regenerated without annuli
immediately below the hydrotheca.
Result: regeneration according to the distal, not the local
type.
I].—Auegust 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 138-17 with 1 annulus
11 am. Cut stem between 4 and 5.
+The stems under observation were removed from the hydrorhiza and
rested on the bottom of flat glass dishes containing about 300 ¢.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.
Vou.2} Torrey.—Hydroid Differentiation and Senescence. 320
August 8, 9:45 am. Hydrotheea of No. 5 with 1 annulus
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
type.
IlI.—Aueust 5, No. 10. Stem with 20 hydranths and ter-
minal bud.
Pedicel 1 with 5 annuli
Pedicel 2 with 0 annuli
Pedicel 3 with 0 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 am. Cut stem between 6 and 7.
August 7, 10:30 am. No. 7 has regenerated in a manner
practically identical with that of the preceding experiment.
Result: regeneration according to the distal, not the local
type.
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.—Aueust 5, No. 4 (see above).
August 8, 10 a.m. Heteromorphic stem.
August 11. Heteromorphie 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
fie. 4.
IIl.— August 5, No. 8 (see above).
August 11. Heteromorphic stem similar in all respects to
preceding (1), with the exception that development had stopped
before a hydranth had appeared.
330 University of California Publications. [ ZooLocy
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. Heteromorphie stem with three annular
basal 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. Heteromorphie stem differing but
slightly from preceding (III), surmounted by a hydrotheea
(fig. 4).
Latter small, and diaphragm not apparent. Hydranth at-
tached to wall by a number of amoeboid strands which are com-
mon along the stem but are seldom met with in hydrotheca.
V.—August 5, No. 9 (see above).
August 8, 4:15 p.m. Heteromorphie stem with 3 basal an-
nuli, a long stem segment and a poorly defined annulus imme-
diately below the hydrotheca. Shoulder process on side of long
stem segment carrying growing stalk of normal type.
VI.—August 5, No. 5 (see above).
August 11. Heteromorphic shoot with terminal hydrotheca.
Three basal annuli and a very long non-annulated stem segment
which passes directly into the hydrotheea, 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 cases. 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 (1). The tendency to develop according to the
Vou. 2] Torrey —Hydroid Differentiation and Senescence. 331
distal type is therefore conspicuous so far as the pedicels are
concerned.
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 would 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. [ ZooLocy
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
consideration.
The resemblance of the phenomena of colonial differentiation
in OC. 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.
UNIVERSITY OF CALIFORNIA PUBLICATIONS
ZOOLOGY
Vol. 2, No. 7, pp. 333-340, Figs. 1-5 December 13, 1905
CONTRIBUTIONS FROM THE LABORATORY
OF THE
MARINE BIOLOGICAL ASSOCIATION OF SAN DIEGO.
Vit:
THE BEHAVIOR OF CORYMORPHA.
BY
HARRY BEAL TORREY.
In a former paper ( :04),1 it was shown that Corymorpha pos-
sesses an unusually wide range of activities for a hydroid. It
responds to mechanical and thermal stimuli, to chemieal 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, Clava 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 eases 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;
* Biological Studies on Corymorpha. I. C. palma and Environment.
Jour. Exp. Zoology. I, No. 3, 1904.
334 University of California Publications. [ ZooLoGy
after which, 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 it 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 deseribed are 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-
lation of fluids in the coenosareal 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
museular activity. The behavior not only of the stem as a whole
but 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 for bringing about a response. This mechanism
is expressed, I believe, in the large, highly vacuolated cells of
the endodermiec 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
diameter.
+ a ae Oe capt a cn ema
Vow. 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
existence.
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 Corymorpha, 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 wpward and came
to rest only when the stem had reached a position approximately
vertical, but wpside down.
Comparing this result with the movements of stems attached
by the proximal end, either resting on or hanging from the sub-
strate, it is obvious that no reversal of behavior in the axial cells
need be assumed. The variable factor 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 ease it never equals the ten-
sion on 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-
tropie orientation of the stem, has been strengthened to some
extent by a further experiment. In my original experiments,
euts were made at frequent intervals half way through the stem
. — —— “or
VoL. 2] Torrey—The Behavior of Corymorpha. 337
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
1 2 3
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
338 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 endoderm 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 restraint.
f 5
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 eut 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] Torrey—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 geotropie response, Corymorpha stands alone among the met-
azoa in possessing a tropic mechanism distinet 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 geotropie reaction. The axial endoderm cells
probably change their volume under a given tension which is
apphed 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
eradual, 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-
pretation.
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CONTRIBUTIONS FROM THE LABORATORY
OF THE
MARINE BIOLOGICAL ASSOCIATION OF SAN DIEGO.
Vili.
DINOFLAGELLATA OF THE SAN DIEGO
REGION.—I. ON HETERODINIUM, A
NEW GENUS OF THE PERIDINIDAE
BY
CHARLES ATWOOD KOFOID.
The investigations of the plankton of the Pacific at the San
Diego Station during the past three years has brought to heht
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 eall 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 (799) from the tropical Atlantic as members of
the genus Peridinum to wit: Peridinium hindmarchu, P. milneri,
P. blackmant, 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
pO
ee Byatt si gy tae
Pg ; AY ACT STi 1Sitypy
f f - %
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 deseribed for
the first time are in the collections of the University of California.
Heterodinium gen. nov.
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
margin or lst 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 if 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 interealary, six
premedians, seven postmedians, one furrow plate, and three ( ?)
antapieals, 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-lke 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 epitheea 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 theea 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 H. scrippst.
DETAILED DESCRIPTION.
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,
Von. 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 coneavity 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 seoop-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 eases as
in H. blackmani and H. hindmarchi.
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 distinetness 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
protuberant.
The hypotheca is usually subequal to the epitheca, being
longer in H. milnerit and H. sphaeroideum, and shorter in H.
blackmani. The posterior end may or may not show a bifurea-
tion into antapical horns. It may be broadly rounded, dome-
shaped and without any antapical differentiations as in H.
sphaeroideum and H. doma, with mere spinules with or without
lists as in H. milneri and H. murrayi, with slight median bifur-
cation as in H. whittingae or with typical elongated antapicals
344 University of California Publications. | ZOOLOGY
as in H. blackmani. In the form of hypotheca Heterodiniwm
thus exhibits a development parallel to that found in the allied
genus Peridinium. The ventral face of the hypotheca 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 H. milneri, or postmedian
in others as in H. scrippsi. The girdle is much more oblique in
the flattened than in the rotund forms. In H. 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 most
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 not a ribbed fin or lst 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 H. sphae-
roideum 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 theeal wall is made up of discrete plates, which, however,
are much less easily separated and much less clearly defined 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
cases 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. ~ 345
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 Euheterodinium 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 theea. In some eases as in H. scrippsi and H. blackman
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, 6, 7 (or 6 and 7) and apicals 1 and 3. This interealary 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 arrangement of the
premedians which attend its presence in species in which it has
been demonstrated. It is greatly reduced in size in flattened
species such as H. 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 epitheea of the flattened species is
formed by the two ventral-apical plates (2, 3) and the two
ventral-premedians (5, 10) but in the more rotund forms the
lateral-premedians (6,9) are partially exposed in a ventral view.
The left interealary in the flattened species is dorsal in position,
but in the rotund forms as H. murrayi and H. 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 band as in
Peridinium and appears to be variously subdivided by reticular
ridges some of which may be suture lines.
The plates of the hypotheea 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 the tip of the right antapical without subdivision
into anterior postmedian and posterior antapical moieties, ex-
cept possibly in H. sphaeroideum. The separation of the adja-
eent postmedian (17) from the ventral median antapical is
VoL. 2] Kofoid.—On Heterodiniwm. 347
often feebly expressed. The antapical 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-
ical horn.
The boundaries of the plates on the right ventral face of the
hypotheeca are subject to much obscurity and considerable varia-
tion in loeation. The relations of the midventral plate (19) to
(
are subject to considerable variation in connection with the vary-
the adjacent postmedian (17) and to the longitudinal furrow
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 hypotheea. In other forms as H. tri-
rostre and H. murrayi it reaches the antapical border. It is
without high membranous lists as a rule though one appears in
HH. murray.
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 H. 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. murrayt,
and H. hindmarchi.
These reticulations are often quite regular as in H. blackmani
and along the lateral margins of the epitheea of H. 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. blackman. 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 Californa Publications. [ ZooLoGy
theea. 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 H. doma are enclosing ridges so thickened by the filling in
of the angles as to leave a subcireular central area. Each reticu-
lation has 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
through the pores, for the polygons bear a 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 fill the interior of the
theea. The nucleus has the usual ellipsoidal form with beaded
chromatin reticulum and lies near the center of the protoplasmic
mass not far from the flagellar pore. It is small and is found
with difficulty. No instances of diffuse reddish coloration often
seen in Peridinitum 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
eases 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 far recorded appears to be H.
blackmani which has a length of 225y, and transverse and dorso-
ventral diameters of 1354 and 160» respectively; the smallest
appears to be H. sphaeroideum with a length of only 42u, and
transdiameter of 39x.
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:
VoL. 2] Kofoid—On Heterodinium. 349
Species Latitude Temperature C.
H. blackmani 7°-380° N. 25°-27°
H. doma 34°-39° N. 16.1°-18.9°
H. hindmarchi 19°—39° N. 15..6°=27 .2°
H. trirostre 262 IN: Dhan
H. leiorhynchum 19°—39° N. 15.5°-24.5°
H. murrayi 14°-28° N. 20°-25.5°
H. milneri 29°-31° N. 26.9°-27.2°
H. sphaeroideum
ier ea San Diego ee
Bly arcane B45 (1 ANTE 14.6°-22.5
H. inaequale
H. triacantha 55°=8il> 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 filtermg 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 inerease 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 Peridimum. 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 difterent
individuals do not indicate as yet any noticeable intergradation.
350 University of Californa Publications. [ ZooLocy
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-
nium, unless indeed the slight midventral expansion on the
suture line be taken to represent a degenerate 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 a narrow transverse furrow which is complete distally.
Its plates (see Schutt (’96)) are not known and it may prove
to have a closer relationship to Heterodinitum when these are
definitely determined.
The midventral pit on the epitheea of Heterodinium resembles
the so-called ‘‘pore’’ in Poroceratium gravidum (Gourret) but
bears a different relation to the thecal plates and is probably not
a homologous structure. In Poroceratium the ‘‘pore’’ hes near
the middorso-ventral line in the middle of the dorsal and ventral
apical plates, whereas in Heterodinium it lies in the suture be-
tween the two ventral apicals.
The generic distinctness of Heterodinium is thus beyond ques-
tion and it belongs with Ceratium, Peridinium and Protoceratium
in the sub-family Ceratiinae.
SYNOPTIC KEY TO THE SPECIES OF HETERODINIUM.
Sphaerodinium subgen. nov.
Body spheroidal, antapical horns not present or only slightly
developed as spines. Epitheca rotund without stout lateral
sutures.
ft; With nosamntapreall horns: Or (Sp Une St sete cae eee sgvtzeteacs 2
ie SWithantapicall’ spines yas oss es eee ees eee See ieee oe 3
2. Outline smooth, sutures faint, reticulations minute..H. sphaeroideum
2. Outline subangular, sutures prominent, reticulations coarse..H. doma
3. No apical horn, thecal markings faint S22 e ee H. milneri
3. Short apical horn, thecal reticulation prominent .............. H. murrayi
Bp) slong apical) horns sects ene eee ee eee H. triacantha
VOL. 2] Kofoid.—On Heterodinium. 351
Euheterodinium subgen. nev.
Epitheea 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
oblique.
1. Epitheca with strongly convex sides, apex broadly rounded _....
ee Ae Sat Neeson eae ore sea ane asec eats eect ten ES ANOEG UAE
1. Epitheca with straight or concave sides, apex not broadly rounded... 2
2. Epitheca low, its altitude about one-half the transdiameter -............
Be Pattie = Maen gn 8 Mes Rn dasa nee ides sect s eomaswtae ean aet kesh Ate edad H. rigdenae
2. Epitheca high, tapering, nearly equal to the transdiameter _............ 3
3h TUGHE Gale gonkel lomut wince Neel eae ae ae ce eee H. trirostre
See Wehteambant cele no tbc cate Cee terpenes ee ea ee eet ee ee ee 4
4. Reticulations very coarse, scantily developed H. leiorhynchum
4. Reticulations subregular, very delicate ..............-..--.-- H. blackmani
4. Reticulations medium sized, very heavy _.....................- H. hindmarchi
Platydinium subgen. nov.
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. Antapieal
horns present.
i, A\ratigyjorioal Innes): CWNAETAO ENING 5 eee cee eee eee H. scrippsi
He A\inireyanicewll INOS; COMET ECG oe see occas ee cere H. whittingae
Heterodinium sphaeroideum sp. nov.
Pl. 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 epitheea, 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 theeal wall. Longitudinal furrow short,
its length less than 0.5 distance to the postmargin, broad and
shallow, its distal two-thirds enlarged.
Theceal 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 postmedian 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 dense, heavily vacuolated, chromatophores irregular,
peripherally located, greenish yellow; nucleus near flagellar pore,
ellipsoidal.
Dimensions :—leneth, 42; transdiameter, 391; width of fur-
row, 4-5; diameter of polygons, 2-3».
Taken once in vertical haul from 165 fathoms off San Diego
in June.
Although this organism does not have the deficient 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 (799), p. 327, Pl. 30, fig. 3.
Plainly belongs to Heterodiniwm 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 Atlantie between 34°-
Bo Ne
VoL. 2 | Kofoid.—On Heterodinium. 353
Heterodinium milneri (Murr. et Whitt.).
Peridinium Milneri Murray and Whitting (’99), p. 327, Pl. 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. murrayt.
Reported from tropical Atlantic in 29°-31° N.
Heterodinium murrayi nom. nov.
Peridinium tripos Murray and Whitting (’99), p. 327, Pl. 30, figs. 4a, b.
non P. tripos (Muller), Ehrenberg (’33), p. 272 = (Ceratiwm tripos).
The specific name tripos must be rejected as it was previously
introduced into the genus Peridinium by Ehrenberg’s (733)
transfer of Cercaria tripos of O. F. Miiller (1786) to the genus
Peridinum. 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 interealary
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
oirdle 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 Atlantie in 14°-31° N.
354 University of California Publications. [ ZooLocy
Heterodinium triacantha (Jirg).
Gonyaulax (?) triacantha Jorgensen (799), p. 35.
Ceratium hyperboreum Cleve (’00), pp. 14-15, Pl. 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 midventral suture of the epitheca
deflected to the left, the widened distal end of the transverse
furrow, and the longer right antapical spine. There is also
some indication that 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 theeal plates in full. In so far as they are indicated in Paul-
sen’s (’04) figures, they conform to those of the genus Hetero-
dinium. There are difficulties in reconciling Cleve’s (’00) figures
with each other, and with those of Paulsen (’04) as well as with
Jérgensen’s description, probably due to the fact, that, as Paul-
sen suggests; Cleve’s figure is reversed, 7.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 Sphaerodimum,
though it does not possess a spheroidal body. It is characterized
by the absence of antapical horns and post indentation, concave
sides of the epitheca, the developed apical horn, and the three
antapical spines.
Dimensions :—length, 72-84; transdiameter, about 50,;
dorsal-ventral, about 45p.
Reported from coasts of Norway and Iceland.
Heterodinium inaequale sp. nov.
[EAL ibs mates Sh Alo)
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 coneave. 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. 359
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-
eavated. The broad midventral suture runs from the longi-
tudinal furrow to the apical pore swerving towards the left at
the ventral pit.
The hypotheea 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 imcurved 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-
dexiotropie 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 theeal 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; transdiameter, 100u; dorso-
ventral, 754; 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.
Pl. 18, figs. 6-8.
A small pentagonal Peridinium-like species with a coarse
polygonal reticulum and slight obliquity of the girdle, resem-
bling P. 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 sloping in a straight line from the apical pore to the girdle,
compressed dorso-ventrally and somewhat excavated in the mid-
ventral region, lateral sutures prominent and doubled. Hy-
potheea also compressed, the 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
eurved 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
Vor. 2] Kofoid.—On Heterodinium. ay
flagellar pore little more than 0.5 of the distance to the post-
margin.
Theeal 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:—leneth, 120-1254; transdiameter, 90-92p;
dorso-ventral, 155; girdle width, 10»; polygons, 2-10p.
Taken in vertical hauls from 90-100 fathoms to surface in
June off San Diego.
This species resembles H. h indmarcht (Murr. et Whitt.) im
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, Pl. 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 Atlantie.
358 University of California Publications. | ZooLoGy
Heterodinium leiorhynchum (Murr. et Whitt.).
Peridinium leiorhynchum Murray and Whitting (799), pp. 326-327, Pl. 29,
figs. 2a, b.
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 epitheca in the sight shoulder on the left
epithecal margin, and in the incomplete suture lines of the latera!
view.
The species is characterized by the somewhat differentiated
apical horn, the unequal, pointed, divergent antapicals termi-
nating in spines, the rotundity at the girdle, prominent suture
ridges (incomplete in figure), smooth or coarsely reticulate theeal
wall, and fenestrated lists of the girdle.
Reported from the warm temperate Atlantie from 20°—40° N.
Heterodinium blackmani (Murr. et. Whitt.).
Peridinium Blackmani Murray and Whitting (’99), pp. 327-328, Pl. 29,
figs. 6a, b, ¢.
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 Heterodiniwm 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 Heterodiniwm.
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 Hindmarchii Murray and Whitting (799), p. 326, Pl. 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 epitheea, 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 epitheea, and coarse reticulations.
Reported from tropical Atlantic from Panama to 34° N.
Heterodinium scrippsi sp. nov.
Pl. 17, figs. 1-5.
A large species with short antapicals, scoop-shaped epitheeca
and coarse irregular reticulations. The body is subheptangular
in face view, swollen at the girdle, and with shallow posterior
bifureation. 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 epitheea 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. | ZooLocy
The postmargin is not set off from the inner margins of the an-
tapicals with which it forms a fairly regular are. The distance
between 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 its width. The longitudinal fur-
row is about 0.6 distance to postmargin in length and is wide
and shallow.
The plates of the theea are typical, the left interecalary 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 to the left and contains the
reniform ventral pit. The suture lines are well developed and
are laticed in places. Hyaline lists are found on the lateral and
postmargins of the hypotheca. The thecal wall including the
eirdle is coarsely and irregularly reticulate with well developed
polygons of 3-5 sides, each with a single central pore. Near
the lateral margins of the epitheca these polygons are somewhat
recular 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 subspherical greenish chromatophores and
an ellipsoidal nucleus near the flagellar foramen.
Dimensions :—length, 140-1554; transdiameter, 100-105, ;
dorso-ventral diameter, 602; width of transverse furrow, 10-12,;
polygons 4-8p, rarely 12p.
Taken in vertical hauls from 95 fathoms off San Diego in
June.
I regard H. scrippsi as the type species of the genus.
_——>--
Vou. 2] Kofoid—On Heterodinium. 361
Heterodinium whittingae sp. nov.
TeAb ley aaweys alaleals}.
A large species with very oblique girdle, elliptical outline,
and shallow rounded bifureation. 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 hght suture
ridges except in the case of the lateral sutures between the apicals
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 hght
and delicate. The polygons are relatively numerous, 97 having
been recorded on the dorsal apical plate.
362 University of California Publications. | ZooLocy
The plasma is exceedingly hyaline and coarsely vacuolated
and its total amount is relatively very small. The nucleus is
minute (12), 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 in transdiameter and 60u
in greatest dorso-ventral extension. Polygons 5-12, in diameter.
Taken in vertical catch from 85 fathoms to the surface off
San Diego in July.
VoL. 2] Kofoid—On Heterodinium. 363
LITERATURE CITED.
Ola7e, 1s 10
: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
Richtung des kleinsten Raumes. Abh. Akad. Wiss. Berlin,
1833, pp. 145-336, Taf. 1-11.
Joergensen, E.
799. Protophyten und Protozoen im Plankton aus der norwegischen
Westkiste. Berg. Mus. Aarb. 1899. No. VI, pp. 1-112,
I-LXXXIII. 5 plates.
MiullerssO rk:
1786. Animaleula Infusoria fluviatilia et marina. Pp. LVI, 365, 50 Taf.
Murray, G., and Whitting, F. G.
799. New Peridiniaceae from the Atlantic. Trans. Linn. Soc., London.
Botany (2), Vol. V, pp. 321-342, Tables I-IX, Pls. 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.
796. Peridiniales. Engler und Prantl’s Nat. Pflanzenfamilien. I
Teil, Abth. b, pp. 1-80, 43 figs.
EXPLANATION OF PLATES.
PLATE 17.
1.—Heterodinium scrippsi ventral view. X 725. ap. p., apical pore;
fl. p., flagellar pore; l. f., longitudinal furrow; v. a., ventral
area; v. p., ventral pit.
2.—Dorsal view of same. X 420.
3.—Oblique view of left side of same. X 420.
4.—Diagrammatie apical view of same. X 420.
5.—Reticulations adjacent to posterior list of transverse furrow.
xX 2725.
[364]
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)
’
Linty. Can ust. Zo ou. Vat.11. [ Kororo | Pate XVIL.
| on Ae oe 2
«
PLATE 18.
Fig. 6.—Heterodinium rigdenae, ventral view. X 410. ap. p-, apical pore;
in fl. p., flagellar pore; l. f., longitudinal furrow; v. a., ventral
F area; v. p., ventral pit.
Fig. 7.—Dorsal view of the same. X 410.
Fig. 8.—View of left side of same. X 410.
Fig. 9.—Ventral view H. inaequale. X 420. Abbreviations as in fig. 6.
Fig. 10.—Oblique view of right side of same. X 420.
[366]
een EE ————————— —_= =
— —_————
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Univ. CAL-Fus.Zoou. Vout.
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mid oe) ’ van :
Pod oa 2
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wan 7 ca
ee -
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‘4 ; : 2 : oat _ v tg
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.
PLATE 19.
Fig. 11.—Ventral view of Heterodinium whittingae. X 420. ap? p., apical
pore; fl. p., flagellar pore; J. f., longitudinal furrow; v. a.,
ventral area; v. p., ventral pit.
Fig. 12.—Dorsal view of same. X 420.
Fig. 13.—View of right side of same. X 420.
Fig. 14.—Optical section at girdle of same. X 420.
Fig. 15.—Ventral view of H. sphaeroideum. 1500.
‘,
{368 |
Untv. Can.Posw. Zon Vo. IL. | Koro | PLATE XIX.
INDEX TO VOLUME II.
Synonyms in italics, titles in boldface.
PAGE
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
LECCE SVGIG! QSL EUSTIG 2's Raat oe bad ot ere? 2 ea PRE nt oe ae ee a 108
AN ONGLDUNEH ICN 4%, cect epesectr cease ee = ea cae a Oe SRE ON OR Se 21
PAOLO LDS LG NUL CIC © re meee eee R ne antes cad eS 8 yee a 266
UCU 2 SS a8 2S Re ne Ey ee OO Dm 2(4
PNGHER. san eet hep cn EER ee ee, Oe ae EOD OGRE ROC Rae 243
aT CET EAE Oe ee Se Na tol ck RON oe 2 ae 241, 243, 244
UWRETUDONGLEN EES eee 0 i Aa BR Oe Ce eee PEE 241, 243, 246
PANES TF1(Cl C: UG me tment et ae re Pass occ t ene aM eee ey 144
ARTS) CAF UT See ee Oe See Fes eee eee eee sos See eee ee ca 145
PRGWONOUES: | op 8 kao a es ee ne Ea oe er en PEPER LE xd eit Sree EE 243
ENGI GUINEA Se eae ee ee ene ee 143
PaO eM ey GCM ONISIS mee) eH Chee sees. sa ceecesteie Sens Aes ees 2, 0; 38
RACE ONS TOC peee soe eee hee ce ne eRe ee es ee oes ee ie I ce 28), 3
Fo UL a pee cement SA eS Se eect ae ee Pe REE Sg 2, 5, 34
SHEL RUST MI OVC TCG (GY S| eS Sali Se MC oot Bec uane me Oe eee An 2, by aD
AUGHHIY HOUR TE eg eRe oe a Pe ne Ae a aN te An eR eee EN ed G5)
LUD UC nee Aen ege ce SEAS Ss OP ch ee, <n ee PO ES Pate 170
LOT OST CDE sc Race sd oe BEE ae ae es eee ey = Ee ee AOE ee EI 243
ENON OMAGH ICCD, ee RPE a I ee atta pee ORE Bee ee 244
AHA OUPUOR omen PR ee ee ea a a ee re ac ae Pm UCD)
WUGOLUCCTNGIUC Oman S288 Lain oe a AOD eer ee 2 eee ee an ae ae 216
PATO UMS UTC VIC Cl smilNh Cigrecen cast catre tetany Nts Unni tu. fie Pe ie we Se Aes ii
@liiona aie cere Oa Clit OT Speen eee ee Re ST AM RES 2 Py ah EIR, pet iv
Physical tcoudiiions, Of 0.2 es cee ee vill
GS SOTA O ULL aNO dL: Reve onee Aeeecete ys ee eee wee eee ee eee ee oe vi
PATEL ES GNU StS 5S US eee neces eee Reet BRC ee ete ee 189
ANTPUICGTIU EYE NCGTL 2. 5 te SPR a ele oe A ee OD ee eno Oe tee ee Re orede 240
PAIS CT CIIAC Ce ie eeneen enter eae kee dc en ee ee ee ee ees Lae 97
PAT EDP GUISE) meses see eter ee SS a eae ee ee ae ON ES ee 187
[evi (CPUS EEUU sees oe eee ee Ee ee LA 188
LEON COMIN cetera Oe PP ee OOD
ULLLALLUS 12.07 CT ene eT Er aD ae ee ce so ee OG
TH OUPOMICHPUh. ooo oe acne occa Se at ee eee ee eens %e ee)
ELT TAUG NPG: 9 fT DCU GR EY, cs ee Bean 270
VB SPDUMIEY ee acter pnt ear ee a ea SB PP a 264, 276
TUTTO WIGS eeeeee Se eI Nye OP ee tse Pe Seal ce at Recker 237, 238, 276
TaN BETES OW ANOS em a ne Sr Sern of Ic 201
370 University of California Publications. [ ZOOLOGY
PAGE
Behavior of Corymorpha, 2-2-2 ee ee ee 333
Beroé forskealli, 25.05. s2ce fe eo oso e cco eee aw acc epee eae one nee nee ce 47
ero (ee 5s cscs sk Pa a snot Be Secs eee eee seen ea wee nec a oe 47
1 <9 0) (0: - eee RR Re ne A ee eee 47
Bibliography of
(HOROPHOROG, (65 550k nes ee ceed reer oe aco tee oe 49
ebero Gimtumn eee eos sa ee tee co ra ee eee 363
Ds Ub go boo) Co f= Weer eer ener et te Reena me Pei See Sr Sb eS rn a eee ee 40
Won-Inerusting /CRUGSEOMICS | a nace oe ee eee 293
Pela pies Cop epi Wek sa aca ma a oe 227
Pelsoie Wimmicatan, oes. se sec cca ctnehseracean teen nceneppene ae ae ener ee eee 102
BA CEUAN A oss 8 en al oe, ee ep eee eee en ees 278
PRUE UATE Ee ca. ee cenc ce a nea ape aise Sa a ar ne ee eee 264
BRCCULORRCOR ons ors cateware epee aaeem arene seaern ee ae aie ano e gear epe escent en ee 264
PB Cel errr Fa ea a ae a Pe ee ap ree eye ee aa 242, 264
A GANG 0c) eit: Nig eee pre te Senn oe ee Meee eB Bn ee Nin SES eS Eee en? 2, 3, 6
re TL: | acd iene ie Ree eee nie Diohe Premeeeet a anbee We ete tn een a 2, 3, 6
1 CF A 221 Sh: ee Rn Snr nena ent oom ten Mee SETS peers eee xiv
BVDRONG oat wk a alps ov cocsfanhe es mere neo da eS Sonn Ge a 53
Bird's) ead | Corane 20 nn 2. co entecennnse Sth are chips ban tenadts pened oer eee 270
SES UMS, UCL OP HOLE yt acc oo nso ce Sen eran emmen ape ence eer ose 47
GOP TSMETLON ALIS, oo asec nn sew enannnns Seen tmrer se ceiserecee ne eente ent agoncsnn neste 47
2) a eeeneee ener I Sears TNR see es Tee a eR SET nr PA aa ES hee 8 ce 47
BUEN, onan cin see cee n nn enmeen tne cnsne cv nnnniwen nese emonentnapng aes atennnae ae sorseaeae Say 6
_ STAG 1 Re le 2 eee a ee peers sa Sonstiges Ooo occ Seca teenie concede aes ee een nee OE Fat
Vaso Vege utah eG Foy ees eee mere een Pere Sse dos cece Seat caus cepa eee cacao earned 2, 3, 6
Bowarbankia, inbeata S:5...2- S50 ee 239
BREW OZOR, asec cient eens pee ere cea ne ere ae cee 237
MUNG a a cee ae ant en nea wn eens aes oe nner 264, 265
UNCLE a ran a ae ee a ae ee deere eee 270
COLEMITT a0 tn cl pg ae PE re a Se oA 265, 267
GUAT VUE OB USL Shi cape ceeae a ae e 265, 272
FU CUUGE an cscs scene enema cen Sec Sogo Nee 271
Ale el Sate, acess esa ers ce wna a er eee 265, 270
TOUS UT MAES ee eee ann need nee neers eee eee er er ee 270
UR ee og a ee ee ae 265, 275
longirostrata -........-.-----------csc-ereeceseceeeeeneeeeneeneeeeennenennenenennensnesnanenanaaaaans 265, 274
FAULT Ba: eR ee ce PaO eee 265, 266
MOLY GUT acces eee oa ee ee ae ee oes eLearn ec 265, 266
PACHCA —.......n ene eeeneeneeneen ec eneeeeeeeeceeeneneceeeeneeecerennenennnnensnemnenanscenenesntneneann 265, 268
PUGEEI -_...n. a --neeneennneenenencneeneeeeceeeeeceneeensneenenemerenecesasmneencecsansceseseemeranaans 265, 271
PUTPUTOUINCEA ~..--------------2---0nnenneenenesnenceeennneeeeneeeneneneecsenenesnanenunenetenuntmennetaasns 268
CGT oY ve st: me te ee eee ere eee ee Sect Preece tee Renee erp 250, 265
ellasi (ee PE LED Ee. ieee rere en ere ee eS Sas etre eee 263
Galberesidse <2 e 249
Caries Se ae ee ee ee 130
MUG CELECT TUNE. cso a a ee era 133
Index to Volume 2. 371
PAGE
(Cand etaa cl ae Ng eae se er che ree be en he nee Soe oleh ds 122
(© aleapriain sey esos eed eee ke nee See eee oe PML es eee. oe ees eee 122
CHEM A ORL OYIC K Beak PS ae a ee = a ee ee ee 123
(OLDIE EST pe ea ae PL Cet os SR SR a eee 130, 135, 139, 142, 150
UGE Cie eee eee Bn Se SAE SEO Se ME So ee 153
NON OUOS. tse MERE R ee CE eo pee Sue ae ec 133
[NOP OES. SS a a a ee fla a Se ey he Pe eR OPS ORE. Wee 140
6 SN ENA UT Semen eres eens oss te Ae ce ene roe hee em eens ees ee a 123
LOLS ES 2S 5 yee EN eA ae 129
UU AWTAVE RCC) NOY XS] acs se eh i Ee Be ee RR ee ee ee 25
FRE ePAY OUI) 5 a 2 aks eh RR Or eee ee eg oe PE 128
TILAIHITO () Tosa ere = Se et ne eer See es SSDs eR Te ec 126
GTS UC OCG wee Ae Ne a a CI Nee EL ety A ec eS eet Ste See 125
TCDS, ODES UO eee Se Bee A eR, ee eee Cee 129
(UCEHE UICC TANS) sae i Sg ee eee ere a ae 127
OH ODS es RE a SSS eg ESS Pa rec 126
(CREE ere pst a ee Ee, eae ee pan AU
(Can ype] MENS dete ae ere See Ra Mee ec PE en a re 2 Oy Io)
(amiga airvesse Ciweik Sig essen tei eee we cen ee eee eee en Be Se cas ea reeneer 2545 at
CRS Ee Tee eg ee eee 2, 4, 12
VAM SF) OY £9 kre ae ee ee ee 2, 4, 12
FIT algae Ae pee REA 2 Res Feat eo Sea ce Sea ae ES 2, 4, 13
RV Opole ee = tee ea es ce Ree ete 5 uk UA Lebeau beteccn oc eeeree-conase 2, 4, 13
Gear earn an a a ee ct ae se eee eee vee Zeros del
(CGH (sR a ee ee 258, 265
CONDUC HOH Tae eae oe ae ee Ee SE Ue EE ee fener eee 192
(ANY DUDUON EDL, = ee tee na ae Say cad a Bs ESTE IEP RSIT NB ele Oro ree eee ree 195
(GOUGING ee cE Re oe 196
DET HOY OU OE CBee ee Mea ea reo ere ee er 196
(ULGLICHPUOY OES, Ap ee ae Ee ee ea EEE eee 196
FIG GOL CCL eee ae 193
arc Cla eee eek a Re ee ee Eee ae eee es 192
aE Fa pT ca ne a ee ee 195
| OYONIEDU EN Eh Ae an met mee ene ee aeeee le ee e eeer eee epee See ee ote een aE 195
CRIT GEL a oe ease 196
jQVSLCLR UEC AE) Ca aoe eect ete ef coe eee eae nO ae eer ar ee eee 193
(Wear eae epee er coo rae 192
(CUNO oh oer eee eae Re eae a a RR Ee SEE SEE 291
OCCT Ce a ae 247
CONOUER ADD ec asses ee se So eae 248
(OLE GCGY ah (2) eee ee Oe ea 171
(CRUG ac ree eee eee a ae PPE 244, 247, 248, 249, 251, 258, 266
CFIC SAUCE = ab Rs i Sle a ec en eo eB eee Ee EE Pee ee ee eer 244
I ETPUGOLOGR asso ee ee Re SE 266
CHEAP TOTUREY, “cos as ee SE or CE SE ee 251
(ell irre Om eels ete eee eee cee Se ere eh ea eRe eee ee 287
TMEHION OWING), eget eran eee ec ce ca nee abso meee pee eer SESE EEE CE eae ERS ene Reo Ser 287, 288
(STEWS, araeekecetecentenosseskece secede come be cstetecceher 6 oe Bet oe ape teen gar oeaee Rees Nese BeOS ee 287,
372 University of California Publications. [ ZooLoGy
PAGE
COU TTC ssc ann ee hk aie Sa SS et See eM EC ae cae eee 286
Wellarind se eee ee pane se Sasa aan cs eee eee eee 242, 286
COPLEY! 11: eee nen eS cea ec ciasene Pe enM ey Otte «ape ) See SORE eto te bake 250
QTOCMAS® abc sac sake Geax HRS aaneaen Re cbe an ote oo ee ees a eee 251
OcliilarTig se ae 244, 247, 248, 249, 250, 251, 258, 263, 265, 266
anguina .......: Ne eR te 9 REE Ie ay el oly 244
CUD ROUNGT A ec cE a a oC ae 270
PO OVE ET, 9 ccs oe vn amnesia Se 263
OP TUT 00 5 £9 ate Oe RR br eR RRS Pe Ne tees SC oP a Ba NER aS ES TR pee eae 249
COC THIOVE 1 1) Saha eR ee Seeley eer sone SRE p A USCSEACA EOE RULE ne Heh ies oe Aer e ie 242, 249, 250
(NUE O PMO sae ee Ce race iyi
ITEM CLI cee cc el ee 172
WUOLACEUB: Fiero eet ec ie eee aca as ee Cee a 172
GF gt ioe |: a ane i aeeee ce SSS Osunp Mt Noarwee van SI A 170
CSG NT OURS esc sens eden nee rim ence erp Fone el eee 171
C032} 1 ie eee ely nen CR SUSE ace Set Grameen re Be ae in eS ee SRS oe See 350
COT OGINUB in cee acco aoe cewek ona Pe NE pao es cae oe 123
PUG OGTR OUS sR aoe a wcrc poco tt Sans oe ee Re ner ae 125
VOTKOMMEITIIR Oo aan co atelectasis oo cee tsa ee ran parla ea se 128
MINOT ooo. Se ee ee RE eRe Se PP A Mes toe 126
Ce) vat Fofsy ay 6142): ae ene eer ee ee net he, a OA =r MRR CORE nn A rkle Se Ree ee 45 oh 243
OTST @ CUTIES tesserae ceeseues ean saan ee ee ae OT ENO we neo 141
CGlapidacalsimis SAG mC ORNS co coe snes ence een cp cee ape ne ee ee et. eck 142
G@liytemmestra 2. 6oe. ion. aaac wwesacsacesteneserasenaee gd eect ta ate dec aoe Bote ook ee 213
RES LTS EM a ache ec See ce term nnin a tose a am eg es 214
DOVER are oe tea wo Este enn a eo ee ep ea er 214
gE 0) «AR AERO Vee aetna a Ses eaiee SLR ieee mene ee veep SUM oF 2, 4, 16
TUNE E) Oc. «ARs ae 2 oop ee ane ee OS CMaN be ten rhea ae, eee SL rr ee: a Wf
hendersoni ...............-.- ELEN ap A a Per eee gee eh ee ees EAS 2, 4, 18
COC OK TAT COT UA og ree ce ene nent Sener ee ei ee 249
Copepoda, collections made ...... Birt tk erp iene oe are ere con ete bo ee eect rer 113
EQ TMALTL OL Oya Ne ce sg ase ee le ee ee 114
COV DUUIG (OTL BUF OV ATONE noe ance aware nee a oe 244
LO 163 3 7c: ee Rea ee Se BENS Eni oe SRE Rare Sp bee ree otras nn ome ence Sere eae te ope eS 218
CT CEU cee a 224
EISCUS) hore ooo eens cated cecere eee eee ee ae ees a Bete ee 225
COMINGS absence Nee be cosa cups eee ope soe ace we ee ons dese Sc ae ee 226
Wareywao rp yeh aa eee aca son cee eee ween 2) 3), 0
Glory Ort ay” a a ep ea rc ee ec Pata
MR LR OE sc wae ce ao we ooo
GRO EROS a cn ac cee pee 334
Uo cXoy 01-00) 70) & MR eee ee et ee nL ean SMR eeee nk SE SSD AE meee er Pars SAPO eee 2 334
prehensile mechanisny <5 <<. c52 oc ee eens ian eee 333
reaction to amechsmical stitrmully 2 seee a ee 333
TEE CRON RGVON 6 sce Sc 323
Goryeo pore, See ea eae a ee 264, 284
SS POLIO eh 9 so sa cece See ene caesar 284
Index to Volume 2. Biles
PAGE
Gtenophores of the San DielO) PVC LON eee ene 45
(Oar C110) ONG Frege eee ae eect pede Serio ee oe nae eNO eRe ane ee ee ee ace 207
Meee el ea ee ras Oe ee one Bo
Sue ripe mcs Ronee ee se Pe ee es aN ee A ec ne Are ce des ca Piste ee eee 54
\CAOTH OF O48 Ve Se a ee ee Eee ae ee 45
IDONOPO ISO) as a Se Se eS a RE ee 53
IDRC ET NOI RES pc EE Ne a Se ee 276
STOVE TD Eg OTTO TLS Wa ee a ce ee eee le ie:
CL UOTIUU TU CTU Cree sae a os Ss a See ee cee tg ahs cease Ee Eee Eee heats? 174
Differentiation in Hydroid Colonies and the Problem of Senescence... 323
FTE) LS ens NI ed SC Ace a Fake er eects ie cal payee, racy sbae eso 2038
Dinoflagellata of the San Diego Region.
I. On Heterodinium, a new Genus of the Dinoflagellata _......... 341
TD pov Ove) TET EES) cece Ae Cer R En ace See ee tere eae ee 2 Ooo
plllirnisstinee? mene eee ee es AU eet | ON ee 7 Sone oa ee 2, 5, 36
TOY EOIN 2a see cee ne toa er ee oS ae eS ere eee 52, 82
VED Qo LCOS ag 5 a cee PSN a ea A ee eee ee 83
LEE Gr Cry, Cl @ ree tem ane cee sree enc Be Oe See Ge cata SE nee Se ee ere 84
nomenclature -......... Pa ees Cee Lad okt 0 in ct eee ee ee eA 84
TAU OS eee ea ee ee BRR TE ra SEs Mee Pele hao a Ses eeeeee 83, 84
DVO ULO UTM CUCU UUILOLE ULI) meee ee eee 85, 91
LD Tat neol yn a 1D seetey ol ayes cots My Oke R ee Me Amey Oe fee Roe ncn Ce nee ee et 91
EXON MOVADO Se a eee re eae ree ae 91
COVA OYONG | eee ec eet cet NA BFE AO Ne ee Oe ee ee oer ee ee 91
PUNO TSO ZC OU ee eee ee ee eas oe ee eee 94
(SISLOY OLAV OAT LOY CG) SS ees ee Set res oe er Sa ene er eS 94
TPDUEL Ieper 9 ek ae a Oe eee eee wT tae ane eee ene 95
CEO OVA DOG bate ere eo oer Pee Se Se ce See emcee ce ee 95
LOW AYO NG | cesta ee eae ee reer Bee ene eee Lea ea! 96
FLUO UH UPA OYE UG OO ak cer Se ee ee ae oe a Ore oii
SECON ENO ZOO MCMG sis ea ee ae cre nee ne 97
PUT LLU TUT Ue Wi ae eee Re ae Pa Re se ee eee tea 95
[Dolo einie tyres Om Sees ee Ss ee nue ye Le ehh, 2 ee eee “Oe ee 85
REGU TEC Zi A ean a eee 85
CZ OL pee aa aE 1 i ere i caved eae eee 86
jOU24E NOVA STO) Lil wees a Cee snes SON eee eee ee a eee eeeer eee 88
(Ple{O) S1aNCVAS| OFS EN gh oe See Sine aes ON ccm ee ee ea ea Rone oa aan cr Se 89
TD VEY OR LON (OY TOI ce Pees che eS Pee ee nee eee Ca 142
DUB TUE TNL GTOL: goes ees haere ee Ae OM ery ue a fee ee EE OS saat epaneta cet oe eee ae an nate 53
DDS a2 0 (2) 042 gute see Sat Oar oe eee nce eRe pe pane eam See Eee cee 2, 4, 30
COE TR C1eT 2A s ewee 2,5, 23, 30
DEE GLO1UO S O11) Cee ee a a an ae Sede ee aes soe ee 210
DEY 0) 0 01 Oe: fu eee ee eee Se Sen CEE ee Ce oe 242
TES TOT EID eae ee eR a ae ee eee EE 250
1 SHAY C0 0010 08: eu nee eee Pe emo 114
LD 00S) (002 eee 240
FERS CPC Ci ee a en ee nea 291
374 University of California Publications. | ZOOLOGY
c PAGE
VUUSIOUD~ Bosc sees an sac = cae coe aca a ce 266
Batorly, (C. Os eccost tee Se an en ee 113
Wea Bin 88s se 2 Sache cs a a ce eC 130
TN Cueva ae cw sa es So sa, Cee eee 1230
AROMA EUS 2 2u2: Be ceees as ec cc ee ee 133
CUR SSIS Ecc eae a ee ee 134
BlGmps ines os cine he oe oe en ee ee Se 131
UGE a a ee ee a es ne ne ee ee re 135
PE EC OUT UU: oe oe se aca se ee ee 142
BYU Hep Ga Pye s 6 eee ad acs es hee ee ee Se eR eR ee 156
PY cLN 2 PS aon EPMO DiC eR ERSTE Ae Wee Tae AP Ao oee Bair sc foe aE ae a ad a 157
rats {0 bb: At eee ee a 1 mata eee pee Oe Cede SPS s MMS Yo oats SAU ARGS AE Ens Mace et 160
BYINOSA 253s oh ae eh Se eee ee ees 159
OTE (0021 Nees aan eee RSE Oe ee Sa Ree ere ae, TP ee Ree Syma ee ee Ein 150, 147
WO05OR (ke ced Se kg Se Pe ee ee eo eee 152
ELL! Nit ae eels CR = a Ba ape Pee cae Seed mor eee Sa Bee 8 2) Ee 0e) Ss a as 154
WBTUR CESS LATLES | esos ees eee spre oc os ane en ee ee ee Be ret Sa Are oe es ee eee 156
BMYCH ARIS, OTR UE OMNI os -oeaSac sence cee ope cee ee eee ek ere cn enacees 47
HOt oleic =) lk: ile nine Se eee See SeenON ae Be En oe RARE, TiN (Se tthe SESS oA Eo A. 5h tS 150
F111 0216: ie ee oe OR as Ree Rt scene PRET chan eee ANSE Oe PAL RES as! 155
galesta, oe sera to Se Se ee Oe PR alan a ee 155
TE) GSU TEOTASISS fae no wo ee cede wc gcc oe Seen epost 151
PUEDE eee gh ad SS wa ap ok ah eos PE eRe RC aS ee 153
0{: (01: eae ee ed een aE ace ee PONS Be Bde canes RH Resi i palen is 152
POUCTUUS: phe aa een ach tee asada ee a COR SR 156
ia er ates. ©). 2e 5 eke. cena te ee ee ee ete Ee PA ee rs a Se 247
UVC) SG a ss ae waka aes sags ee co cee nn Oc a a eo 248
LD efor ym hig 2s 5 Semen ee enna a MIN sn Serie Siwiphineae eae h ynein saa en Bede Leh 242, 247
TPA er Gl nn wrk Ch ges 8 a cee Pe ee cee ac tect cee ar ee yy ve
LDAv Col cyVo hghoh ev Wy g2 a 40-] yb 1 Ree een hae Satna cent np Mateenyn) inom Pa a teeth nit SST 2, 3, 8
TEVTVLO SUM Y e oos a eee gcc o es aae 2, 3, 8
SDS Wcbbehse cee reine Ooh en Sage anes ee Gn ERO STL TN ee Pa te 2, 9
Mp EHO OANA ITN sees occas occ se neat cee eh een ae ee 351
Huplokamiig “CaliEOrnensisy aes e nes: era ceee ce encasement ee 46
Buberpe: AewtaL Ons: ec ees ee cece a a ce 212
MCW Nene esse aaa ce a oo 212
dV Gin 2: eas ee ST RE SOM Petes es otal PON Ce el ik Beeline Set ee A npn meet 291
OUR CULTS, co 20 a ee Oe ee te ee ee ee ee 270
ML CE @VLOUGLE SS cw sak sce a a ge a rae ee 291
MEM DTANACHO CLUTCH GAN ocak ee Te op ee aaa eee ee 292
SEU COC: Co cies oa a ee a ee ne Sik 263
POUUS EPO oR ae aaa a nee 290
TBST ec 2a ec eae ree ae 242, 290
CAF ce: ee ele RPL we PS es Ms Rohan yea er pe cen pe Pee aS aaa eB 114
GALS) escort ne 145
_—.— ea Sw ows
Index to Volume 2. PAS
PAGE
(Creme ara elgg eee ok ax oS hae Bo OEE oo es at oe Pee. ee tee 248
UCU COUSIN SE chore Rea ee EEN. Ss Ten ee eee A Ld eee 286
IESVCONIE) 2 2a Setar ne See ie enon ET eect Aas ee oe eS EP 247, 249
UU TURE 22 Oo EO ae anes RE SRR Er. Sa ge Mees BR Poe IETS 2% 249
CICLO CUIGHI TOHORIG , SG a ie oe en | Oa PY, See eee y Ona ere nee pe le 285
(Giyanin Olle ma asters eat cere kh te Wee Sete eee ee eee
Generation, solitary of
Cyclosalpa
CULTLTIN Sige oe eae ee 2 ae aN i Le Rea ee Ps ee, Rive Sede ae OL see 60
ail Cr nent a tere | esa eee eee eek he sh om Be Ree p ne Meee ee 54
Salpa
confoderata-scutigera =. 22 Seed eect Sd een ce ee 81
ayia cline © ge wee erence Pee cx Bee he ee eee 79
CLEMO GT evbI GARMIN CN O IN eyit cl pam ee tse eee e Seea ae es ere Stee 73
[ial KENSHUTSCCCO(ST SEITE yee So See SS ne ee ee errr ae 70
Generation, aggregate of
Cyclosalpa
A ATO Siig sen ee ee deere eee m5 Ps ere Raiden eee Oe Sse ALC LEA a a eh oe a 61
[OGHIRENT, “Sos se Sete ie Ne epee Se ee Ce ee eee eee ee 55
Salpa
Comba demaitiass Cutie ere eee eases cee see nea cree eee we eae 81
GRANT ONG Her A ase ec ce eee ee eR Seen eee ee oe ee ees Ae 80
Clemo Grab Gam CT OTA ee ae ee ee es ar oe ee oe eee 75
fet Te OUTIL S aT CID A Gein meet ate ONES A resect a een oe eins Se ee eee 65
ATEN TT CYNE SEY pec epee eee ee ee een ees eee Me Re oat See 71
PZ IMENTAL A= COT OLA OIG LIN GS pepe i ee meee ee See a 77
COMI OTIS CLAUS geen ae wate ns cae a nce ee et ee = ORE re ne woe RE Pe Ame ee ke 213
(POPUOV INTE GOP CCH LOE eer eae eR ER ye Ee er EE Re 341, 354
Gaya OL asthe ety ee Sees eae tes ae Te CE es ae el AM ree tes 2-25, Oy 0
(Gry TTT Oo epee sense ee ee ace nee MU ee a ae ee So ee ile}, Mlee way
TFTA Ci Gs ens Saeco, ee oer a Pe eRe ie ee OS 2 eee ee Be Bis IO)
ede ee r peA MTNA ALTA eee ese eee ee 2, 3, 10
OO Cine ee eRe as ies recreates meee eneee fon ve vn pute Soe a rene ey tre UN Beat. Vall
SVU SS HED CFs AG pape reae eee Seen sen eee Seen ene ce ae eee Zo Bh, dul
PETIGID GLG UA CUS acre ae eae me, Se eS tence 9 Oe eas nee es Pe ee 210) 212
EM GTTUUCCLUCLTULUG Rare see Pee Ose ee, ea eee ON ene Ne ae ates AR WAL, aisy¢/
TEE UP RON TRG cy Bend or GO eer Pa ee ae on ES 181
NEG CIO CHUCEG Or ter eee cee cee eee Se EL te «Shen ee tee ocean ee 182
ClOUSi) See Ce 8 cece REP eS Sea nh oe ee a AR Po Ce ee 185
WOTUGUC ON NG eee tates eet wen ks es 2c eae Neer wee SSG aa eee 186
FSH TE © CO MIND TIT Nip pesos eee eee eu ie ee ere ee 341, 366
TI Sa earn sa eno a ence pn ee crs Boece Cee ORL cast eae ne ecateses 358
LOWE Aoki ee wet EE oe ee ee ee Namen eae ee ne 352
Farrar hirsute ere eee nace ne Ses Set 8 Rae ote a ac Aen (GOL
TOS EY NEES as eres ee arc ere re Eo PP 304
Meteors banyan G Darna eee nee eee eee eee eee Di SA ea Ae Pe ER 358
ENON) Tae es es et A ern a ae a ee Re SER we ee Pact eu eats 353
376 University of California Publications. [ ZOOLOGY
PAGE
1441.0 Vd) 1 eee ene oP een ee RSE” See eemeacie fame ee ee eee 393
Da 0 Coy co eee or eee Rear tec rR eee so ene acc oceans coed See 356
(63 0) 072) ee See er ee ES oe ee ee eh acdsee 359
)OYSF: e151 00 (21) 112 eae eee ee Ree RU aMeeen Shoe Mee ater encores Meee eee OSS ETS 351
ETT CRIA a See vane w cca se aa ec ce ee 354
ETN TEOBE TC a eee ee eer ct ogee Se 357
3 FL 15 BT 2g: > a A RP eee PSone SP Ae irs Ree a ro 361
Biblio eraphty 08 0 ce... - nt sebe ngs sacnads asa ancce ode ansstnane anes ee ngse eens teens erence 363
Dietimitiom: i Ores |e sani cle oe sce eee ie eee 341
Detailed. discussion of structure. -....2<:3 ee esse cet tence 342, 350
BEGG Bis REMOTE OE en Sa te 350-351
Deseripiion of apecies “OF <2 ie ese eae ee ee 351-3862
Wastrarbutiom 5k cscs cen ac esata ace ee 349
DLO Gator OP. a2 2 scsescescasc canoe ener newer nena cee Diooedeios Settne ee 349
Uc] Estoy cto) ito) ae Saale eRe mer SER Rascletas EME Sareea ten eeRN OREe <a e aba Se RE 350
TEV eho T Sh Ma Ch ta 7 oc Ses ccs ak ce sg ee er 181
GGG ON NAD CUS occu, 5 -csee ease cae ee ee ee rae 182
=) OL HAD Gi 0) 1): AMORA. SUNMEMEESOL CP CRA ON Be nee Ub ere Sop Sere Srey Sees sy RR sy sorocc ca 183
CANIS tees lee Coenen eM) Himes a ote rth eu rene re Ee eee coe a AS ih 185
VOMAPEROOR YIU ccs wea a ces strgenrs ch Sap dete ga See ae ee 186
CNet 22) ene cen ee einer SameneeenEr Siesenpeneescra ty we ACT Fe oper st ee SE SNS 184
POLESIOS) ais pan at nec ieee Bee Bibs erent MMe Broo ht nana e. Saabs: Mer 186
History of Survey—
AS TOD 2 ocec soca ocak Nan lca hanes eae cA Ae a Ree ee ce era xii
CO CIS ca is sae nS eee aaa So Since an inc gcd Cem see nee rece ces ee ct cee xiili—xiv
UU AS 6 Se oo a serch pans el a og Sc oS ae A ek ee xiii
DRO CRON 9 own cance pi eee xii
ope lege OAT eT 2 0TM asp ete de ele nite ee eee ee Ra re Reena xill
Sth aTatd ets cfu epee eee se SMR Ale oe Oe etc ee ens rae rE ee co xii
SER OUOEIUUUT ALE | <5 haces a re a ace aa 53
BON OT UUINE, seek het nen once een sree ale ee cee deanna eter eet Seen 76
Eby dractimin..calitorninee jxo 2s Sas eaten es nan be ee ee 25.8509
Ds ge hegY4 F Vs EY - ee nape eM Semesee Mar SSaweerseeceemeomennees 8 1 0 ke Anne Pep By)
Hydroids of the ‘San Diego Re wion qo 2 sen acces ee 1
TPG OP TOT OC scx 5 os a eo wa eens re yf
WE (Xap re oe ae ee Bea ete mre eo AUER reenter eee ome crea eee net ape anes 192
TESS sc roa esa ce I 8 EE ce ac ee 53
Keys to
Eby drords Same Dero) Repu ons 2.2 se ese eee cece eee eee 3
families ‘ot Chilostomiste: 22... ee 242
genera of Bicellariide .......... Ns ete os eee ee 264
Ceasar iss ee ee a ce ae eee ee 250
Gey UUAN CSR eae er er 118
hos (0) 0] |<: re eee USO NO Nee Sve EOE ME ee ot 205
BPOCLER OL ACC 52S cee ce a ea 243
Whine gy * hc ec ohn a 265
(Claeys ka ce ee cc is ee ce ae 5 ee 124
ean’ T-
Index to Volume 2. 377
PAGE
LOEB GG ETE ieee Eo Sie eee en em _ apa ee vn ee cero 193
We are eae era ok Re wR See eo Ses hs | 287
DD AUKCET IT a(S) os ee OE ee Ne Serpe 131
TIONG EIEN Sextet ee ore eee aE ee pee ee RRR CE pis te 157
seh eee ee eee se tersen nad. cones tees Se ee Meee eon ee ee See 151
TDAUVGI SHUG Ee SS ke oe etna ee ee ei eae ne ee eee dS 247
EVO RETO CUI Tes see ace a ase eet ce eS es eee eee 350
Hee GET OTN) (LU Se eee eee ee Sse cer ee ec Se eens eee 182
INIT emf al eae ered a me ce cent IY tee ee ee eS Se eee ee 251
COLE Gib Nase tee ee SIM ie ok Nee Ad RET eats CW oer ak 163
SS GETTY ONTO Rare Naa Be Oe Ne Je eR a ne ce Pe 258
SUSE ORV Alp. “Seeeten ace ees aha ee eRe the ON Coen eee eel metye EAE 278
HCoordhe (Charl Gi Atiyy,0.0 Cae eee ane argos eae rent ee Sew 241
DEH Ono Koxc ener tea hie Cee eee ete eee Re Ue fe ED eRe OM ES ad 199
TEAS OUTDO Sen gape ere ceed Nelo ret We Ure Sr let see te eS tee 200
TW cast OG earaie GLDILLT @) elie een nee Oe soe Pee oe ee ee cen 2, 4, 20
VOSA OER VS wr coh Sac ORE See eRe En ee eee EE aE 2, 3, 20
TG. COME, "sts eh ere SN ne ee ee Aen ee ee ee ee eee eee ee ee vi
JE OXOOR CH DTS FM f ODP LES — esas cdc Seg ee aos ee eee ee 1ST
LEGON DTU RELL reek Nee ORR nk eee lente Sap NE UO Bere tt Ser A ok gO Ee PO SS 179
TETRAMER eee se Bea ean Lee ak eee Benen Peer 179
List of species
sETi@ er. elim er epee see ee oe ee ee ee a ee 349
IFAS GLU ON CLS et re eee nee nee Were A Oc Ss 2 no ne 2
Nomen ers Gury ora Thy OZ ale eee ese saes eee eee ce 241
Re) ove MC OP Sp Od al Lea a eae Sc ee ae ace bee en ER 116
IPelleverike, MM WRACIEY oe aceee co See ne eet Naas ee Pee ea) Ona Pa zen alas aaa eee 52
VOOR esses ese eee ee ee See a aR Re ape eA 9 Ree ORI ee tet 46
VES) OI CAA OLE ULR © SLE EI Nase ei) eek ee Pee De ee sete 163
Tp ervoqentiinbn ey Be ks Eas ee Se ee eae ee a 17/9)
RUNG CUI Ta Se Soe See Rt eee eae ee RC POR NE 8 SEA oe 179
MammemSiolo cicali ASSO Cla OM pees eee ee es eee ii, xi, xill
Mecynocera clausi .............. SE eae Se Pe Re ee eee en an 137
Mienupeal 22 a a ee a ee arene Ne, eas oe ee Nee eR ad Ate 240
COTA CCUG parece re es as areata. Oe Cae ee ee oe Aloe ee eee 254
TLC CUE pce ree ne eR Pe en Pn Nace eae el te 251, 256
(OTSA See em ee a me a ce A ras eect Soe 251, 253
(OX CKEWGKES AT FEBS Bes cape ese see ce ae Pee eae nee ee eR re 251, 254
Ocerdenttalli sie cartel CTSis ieee. ee eee eee nee eee eee ee ee 251, 255
clo tag eee: eee cee me ON Pee, ae SES Sh eS ee ae 251, 257
GQTERNEN Gh Mee See Set aes a ns eee ee ee ee estan Nace re ee een 251
Meteorological conditions of La Jolla,
SEDUCE OW eNO Bo sere ee ee ae a ee ee iv, Vv, vi
IQUECEN Bren (0 WIEN 0 HBr HO RO ag Re ee le Oe, Se ie iene nee ee Oe ee 177
VOX EXC cnt th BU TA eS a ee 178
YU OLS GDN ea ee a a Cy RR EO UEC
TRV Grea) ae Se aa re ee ee ee ee eee ae alee
378 University of California Publications. [ ZooLocy
PAGE
Microsetella 222. ud oe ee Se i ee 210
OBC © 2a eccte re Sone ss eae oe 211
Molluseoida. .sic2ek. Saeko edt ee 242
TE OVO CUM AUS tsa osha ea A eee ee pe eee ee 123
Neoscolemithraa: «iss sacs ee eh ee ee 163
Non-Incrusting Chilostomatous Bryozoa of the West Coast of North
JAMO TIGA 55 co ha ets Sia en ee Sy Nt er 235
Fife © Sel Sane a eee eee a eee sete ee Rs er eres RMmOnoe eT) et BUA: Be 248, 285
GQUICULOIIIRS. Solves Sek Se oct ee rp Ire a 286
COT ROUL ATG 9 a3 cee dao 249
Noten dip te of. oso heen con a ose Ree A a oe pa 242, 285
Obelia A corona Wes cee oe eee ae oe ee ee eee eee 2, 4, 14
CU CHOCOMD, » eocse asi. SP ato as en ee 2, 4, 15
SOM CUA Hebe scctee teste erect bee atte eee ee eee, Dee ee BE ee 2, 4, 15
Oitwons, NANA) Sede soe Aes te et ee oe 209
plumufera fienc seek ce Ste en ee ee ee ee 207
SPUN OSTING - sa Reece en cateencncd ep ees pa ee oe ee 207
[Ol iYc:-\s | es. ae Cont ener oe Re we! eee Nee A 215
Omereay. Hs2cees ees Ske See oer See en Mee Pas eae OE a oe Rad ig PE bt, ONT
conifera ........ Pe en En e neers en een SEY! EE Pe eee eee ha coo 216
TUN ba. (ie ece sees dca cn espana aoc esdnee eaeee e 217
COPE Cla) a8 8 re ce ee ee Gee 240
OPNATROROTO” wxss.5 kate tcc ice nce 265
RUC OU ED. ca caine etree eo iicnc eee Re ee ine yo Oe
Onrntmo poring): 2208 8 Be hres eh es een
OUUOULOTVG Socal ee a ee eee ee eee 270
Paracel sme: t--5 eee ee 5 Scere a Pat ttae oes Een eee 138
IPOVACRIANUS se ee ete poe oc eee oe Sek ec oe eS 139
| a8 eee a ee See ct nes SUB RaOe Pere ane Rear act havens he AREER ME learn at A . 140
PGT GCOLUNUS: TAU ETMNCUG a oo ee acco ct ie et ee oe ee aif
POV GVONTOWANG «x3. oh Se oe ee Se Te oat, een eee eu
Parapontelliniga: 2.2. :.8e. 5c o Scie s occtk ee en eee ees Le eee ee eer ee
PC GF Bh He ech pat fa ca Rea arg a he he 1 ane gle os)
CONF GOT OTE 5S Re ree Pe A ae 81
Pelagic Copepoda of the San Diego Region, The —........... 113
Pelagic Tunicata of the San Diego Region, excepting the Carvacca= 51
alternation /0© \generatlon: Wn 2 sees see ee ere ee ee 52
Pe rciriytrinn 97 oes aie re oe ee ee ee ee one ee 341, 342, 344, 349
Peridinium: iblackomany <.55-.5 0s ee ese 341, 358
GOTT ee oe ec ea Se Sn eh 341, 352
PANAMGTCRY i SE A RR ee re ee 341, 359
LEtOTRYNGHUM. 22.5.4 =. = Se coer oe ee 341, 358
TOLIUON RS (2S ruse A Be ee, a are ee See 341, 353
CEU OS Rae ea aC Se 341, 353
tHrostve. foe ee ee ee ee 341, 357
Perm ar vise: ssc 8 oe a aoe eee ee 2535.9
By opus e- 5S ss cae oe oe eee » HEY
bidentatis® 2s. ee ee 191
Index to Volume 2. 379
PAGE
Thy SiColc ON CIGTON Sm OE AT Caine ee ces ica! eee eee cc eld oo ee eee ee viii
AT eshte ys CUNT VAD gas ates eee a 6 ek ve A Ee ee ence i een eg ee ee 351
AEs airs corre In eapereteree one eres ves Aa eee ne ee ee oo coe 45, 46
IO ce ammenities dae Ae RO ees om as are eee 46
JES WER) one N el WUTC S) Aas seen eee ee RA oh Ue ee eee ee ee ere ese . 45
PU CAAT OTN Cl Meee de oo eee 2 eI ed 7 SE ee Re eee ee ee 174
TeAlmitaatyyil keygeaieeh Soy ON TR ee 5 ay eR gt OS eee ee eee 2, 5, 37
SA BX EYEE Lay OLS 0) ONE a eters a ere Se oe SE eee ee 2, 5, 38
era a EDT OVC CS ye aoe oe eee ee a ee a See 2, 5, 38
SC UC Cele rn ete Af Ree ee ae Poe cara 2 oad er ewe Oo
UE lpaataace id a exis 0 Ee AO as ee a Nr eee eee 2, 3, oo
Pleuromamma abdominalis —... Bios Mea eT Area I RS POY 2 174
LO LOTTUCT UCL C iss some ee ee aa ae ARE weet ome ee ey Soe eed Hep eae ee 174
Fee OIT NS ee ge ce ee a PRM ee ERE aE 176
Diy OU AUIESY Sa Be Ss re see eee se ee ee EO ee eee 176
TO CLO Gamma acme cre See sc Mies a oa ee eee 113, 205
VEO E20 ie Se ce Ox es Ie Ae EE er me eer Peeiee ane 237
EXO ARMA HIG Esc Lele it a Pe See eee a reser, ee Te et See ee ce a IE PIE oa ee 199
TE Xowat eS UhicG eek = ean eke ce ne Ee eae lee roe, ee ER ee Rane eee Pre 198
TONIC CLUCTUG Cece eer es et a OE A fel SO se ND ee I ee ee IBS 199
FER oat ee IN en ses wpe ee ie SR Sn ee eee ee A eda ee 198
OT. 6 COTA GIUTTI puree ee ee ae ea AN 2 ee Sees ee eee 350
ET GO CST UE TUT N gee eee eee ee te a ls SoA dg ee ce 350
J PEAT RONIAD RG meee a ee ee Se gaa 53
Ia O'S UNL EA eee nee a oe ea es Ce oe Nor D soces Sh eras nee ones 53, 98
nom GUT we ee ee Se es ee rs n Sac dA ee ae 98
A ya gC OYSK OW SUING EE 2 cee ae Se a ree ER SE 53, 97
FERPUUTUG GE CHGS arrears OR Pe Seta ee IY ot Bn Se 5 SL eee eae ee 135, 189
J BUTE ESTAR NN i D6 gE se a ee Peer i, 51
TRA UAE TREE ces ae ee eR eS cs NOIR OS See ete MO ES ete Bn Oreos 339
TRR(G)| OSH SRO ole ENT KON ape eA RPE Es ene EEE 235
SUC OTIUCLTIUC) meee ae 0 Sand a ee ee Rae Se oe Ben ee TA og aloe eee ae 287
AS CVG OIIUCH AUC Ca tee erate eS om een ce ca ee ee pe eee 286
[SUCH SY he BS ot pee ese Nt Dele IER ORR PRN PE OE Ee 53
Sal pa SF ad cee Rt tan RN RA SS Un SOR lee hs 28 See rt 53
SURES oes eee a ae anes ER Ll eB PR et eee 59
(CGT OV ON Ate Cte ee eee ec ER Oly EY Alpe ee ne eA 73
BON MORON ROMA De sya Be hen Pot er Oo Re a tae ee a 80
COMLOSMOTAGA-SCUbMME EAN Me cece se eiar ees Ae Se AN ct Be eae eee 80
CORGUOTINTS eee ena SBA bien WINE Pete A Ona ih E ee RS ats Se aan ree 76
COGN OMIT US 9), OUY GT LUC Clare eae ee wee ee ee 76
LO TRONG OCT SHEA OOM IUD. 2px Sc i a eae aR Ne eee 76
COSTE ea eet A rr na RR RA sO cca Ponce on ater ce cthne ae 70
COS CQEC= LLCS 11 ae ee ee enn aN Wile te on ee es 2 ae 8 eee 70
LET OCT OLE UCC gure ee eI ok oe Se, ae RE wre nce he gee not rsd Sd seee ssa 73
(LEVIN Cras Cea ITU CT COT Ese ee a ee nee ee 73
FRCTETOUGAUTUG Cet ce eae he Na Baas Pe ce “econ eee 81
380 University of California Publications. [ ZooLocy
PAGE
PUGET OP INOS oe reat on tase ke Ss pit RR SV eee eee ee 64
PUSiL ORM SSM CLA Ga ee sec ceenco ee oa cae oe cee ee 64
LOVIN QC R TT ean cs ac See a 67
MUU CT ONGC 58 AR ea csats och eae area ie een cer 13
OCLOP ONE ae Oe on roe de 80, 81
PO OUY CR ECO area ae ae See ee ene 76
MTCC Ch nce a ep ae a ochre re 64
PUMCULAUA-f UST OVINAS,.ascciscoctcn 8 oo ceen ce nee cee ee 64
BOW ORO sere ears rece a a Re ee ee 80, 81
SCULIO ETG-CON,OCGCTOLG. \-s55-cncn. Re ee ee ee 81
LL F 7) eee eMart PEM as Se PS eaten FSS a ETE eS Oe eo ee eS noc 70
a] GUT COS GAH sess Sao an ee Ete Sas eee pa 70
SSO TN esc a Se a ee 76
ZONATPASCONOREOTIOIS aes sex cnkcn son cee cece ae ee et ee 76
SS BULLER sf A oe ccee tee ie ee a ee ee eS a eS 53
LUD UGE) SET OSS UA Feo cag sa roonecesece a5. seteneccaee neice Bene St ee AE 246
San Diego Region
ACLONODNOTES = OL. chee Sst acct ent se ie Seale Se oe Se ee en 45
Wimotla eellatal tO c.A< b ns hea aeace ee peep ha Ree pees Lape eee 341
EV VOM OLS HOT, 62253, sere ees scan enc ua Coe mes concen eae ete NOR Pee ee 1
Nonzinerustiine: (Bryozoa .0%° -.... 2 2 eee Sos aes ne DO
Pelagic Copepoda rx, ta eee eee een en eatin Se aot arte 113
Peleionc: Tnmicate OL. secs eon seecacss ene Oe wee eee pee 31
RSS ELUNE Seen: ccnp Sea cege sae teen deeceoce ces Bigs sae year id Sete eect natn NE 218
Era DUS] 2 ae ee re por mens eos Metab coe ak eae. See teaser nt ne atte ork ste Ss 221
CULAR ch ere aI Se ea ene a an a sree oe 221
LONG ase i eg eS See ES oe oe ae ee oe ee ee 221
CORN C/L LT Caer ee Rd a een Eas MERE act SORE os SORE ares ent gape cs He oet Jace 0% 219
OS a ce er Oa ae ee 219
MOTI GG apc R cade case desea avedacis sear eeae beens Slee ooo g eae ee ee ae ena
1G A 17 (eam ar A? ee Se ee in ek eens en Sue EY ie RNa ae 219
Fo1CC ol Fi: pe na ge tee REPOS SOME RRP mcg Rip rh Sn EE MR eae eenere ian gO ARES Sins 222
Sieve Fevov urls (Cr b 61; - eee Raeien Me ee Semenaetie We Cee Sern ner Aieyge oe nee 5 ea ebntas hae Eon 162
SCOLG CUED Ig Soph acah sone 8k cre Oe cca epee eh atria aA NaS cs tea eee cee ee ee 163
DGUGLY AY ext eet en re ee 165
C11: eae en ee ee PENCE Cael en ST eet PEED EPR im Penn Se 164
ESO GSMTAS | cee ae eer ee 166
SUS a, isaac ca a oa Se 170
Bub Gem Gae <i casc kkcata ee n e n ee ee S 167
Sertbellig SOviO a. ce ore Fe a ee eee 207
SSOP UTA cance apis seach ore a eae er ro 247
CLOG, oi cach wwe ec coc cece eee ee 248
Sormpocellanne oan eee ee eee Prenaereem Wee Somer ane 250, 258
DT OUUS CLES, aso ah od a ac S cette ee Se ne ea 259
californiGa, 2.57 2 a ee ee ee 258, 259
RG CTU SES ase a ag ere 258, 261
E6017), 31 a ae A Te et 256
WABUANG 5 ce he ac ee eae cee ee 258,
a
we
SO Pe ae
Index to Volume 2. 381
PAGE
peer eee OT OU nee eno ccs Claes en eS Bo pe ee 2, 4, 26
ell CHT cle ne Sent afte aS oe ree ee ee en Ee ee 2, 4, 26
[OIG TURN SalSt: 2, on «ea ee eee. RenenneMmennrces a nBerm se. wnat 2, 4, 27
Lennie cieementeeeeese Serre fe eS ce) ee Se ee en aE ae 2, 4, 28
EROS) UCT TS i oo Se ae eee mE eae ei eR CR aa RN NLA Rs 2, 5, 28
iV0; 7 C) eer eerie ee coe ns banc ee oe Re eo A ee EN eet eee ZnO 29)
PSCC UET UTE RTE. aN G( a2 Tg eae ge eee Se tne ee a I eg ea pee 32, 33
(CABSITIV Tg UNS) se Sek a eNO ee AR SL ete tee Oe eae eae PR re PS REE 2, 5, 30
CULT CU Coane ee ie ie sate eae ce ence tee ys et We Aa ne ee yO Be
GUI Cel Uae eee ee = SE OES | eee Bee eee hse Se A nea ee aoa
Sertularia
HUG UU Cee Det LE We ot ee oe ee anal wet terete VE ae oe We 244
CUCU OT Care oe ea es ede Ree eee eee es Ee le he es ee nea 248
NO RUC CHU Cerro neat eek tA ee ee A A ee ee eee 249
EP UIA De a RS Sed HE ar eC BC ee RE See 266
Serr tu ATG] ae eee cee se eer eee ores emcee a EES eh ee eed 2235) 2
SAU DEGhUN OM COTGIUVTUG Ime ae teed eee aes ere ene Ae Me a 266
SS] OUMEZY2N RGN TAD se a ae ee Re eee 350
Slariies sallarie clam Ora Orsi tage eee ee ee ae ea eae eee x, Sau
PS icuteUIS mer OIE Tt ystems eames mete oe Ae aie eee 2 SOOO aoe erie tence a ene ae ee xi
ST AVG OCH 2 WINE NTI Ne Sete We GS ced en aa us See aay CPP aN Ane PRONE AW even RRP eRe Bee xi
LUMO T SU Ver Steg] leat ete kere ee Ee eee SANS a eee ae ee xi
USS CUS ys casas eee oe cee Oe cece te oe eee aan Sows Rea oe eee Sete ce oe eee eae ees xi
OLEN CONG peered tes ree me ek cee AE SR a OD bate mee eee ee xi
SE eg ee ene Ae ee De Ee ye ECL Sa ee eee eee xi}
SHEN Pei os oe Se nS a OP Se eee ee ee ree 264, 278
Gallic apenas es eee SS ee a ee oP Ce Ae 278, 281
CUT tan gee ee eee sa ae cee ae ee ee ee ee oni Ce ace 278, 279
OG CUCIS Mitel lS pare seers ead ee eat ce rn eRe RCN SoU Jue neyo 278, 280
TSA ATMA QDI See Re es ea te Ce enh oS ley toon aE NSN er 285
EvSval GUI eA © Vane seen te eae sees se re eye ver Se a ee Se 286
SE May aan OVOLOOT CA) nse. Saree Seer nt kg Le FE ee ii
CENT OVS) CO) Bec eee eee rp Ee We Een Snr SER Re Oe ed iil
CURSE ws ae eR a ON ed A SO re ees et Se, BEY ae 1
HIS i UGS ONE a a Ea ewe eee ne Senha ons eS He Seas xii
ISHN HE CO) RTI ae See ey re Oe ers 2, 22, 26
MSTA CETTE et ee Oe re neat ee 173
HIS Gay rece oe see ws ee es Me ee Ae Ser ea a ea cee ana aes Seecee 53
LS TCL cee eet re oe eg ee Te Se ae Se eae Nena See Ses Se 3
PT cap GG ea ae www ace ES a Se a esa a Oven AES e-em 58
CUTIES a fe te ee ope See UE ud os SR se ye Oe 2, 4, 21, 32
ANAS YOOVOUE TG HITE A SY ee decd. Sse se se sae SS ee eee a ee ere ee ee eee 24, 25, 26
Women Eleeriyy Ss Cel tee mee os cee nae oe ore tcesecee ee ee 45, 323, 333
Ayal} ill aye ay AGT: O CO ea) ey caer te oss cease weee presen ake ee eeu ae en oeoem as teseseeccs eo 2, 3, 10
(Win eu Hee ea eS waa ee ae Sec SE oces a Jogde case enact) Se See 147
TOME (O)LG pe eee ate ne ae AE ee oe Po ee 148
SMAI Gepost Sees Se Rees has eterna saa tp Re Nees Bevel as Do eatoee 149
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