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ALLAN HANCOCK PACIFIC EXPEDITIONS 



VOLUME 14 



NUMBER 3 



BRYOZOA OF THE PACIFIC COAST 
OF AMERICA 

Part 3, Cyclostomata, Ctenostomata, 
Entoprocta, and Addenda 

(Plates 65-82) 



BY 



RAYMOND C. OSBURN, Ph.D., D.Sc. 



h 




THE UNIVERSITY OF SOUTHERN CALIFORNIA PRESS 



REPORTS ON THE COLLECTIONS OBTAINED BY ALLAN HANCOCK PACIFIC EXPEDITIONS 
OF VELERO III OFF THE COAST OF MEXICO, CENTRAL AMERICA, SOUTH AMERICA, AND 
GALAPAGOS ISLANDS IN 1932, IN 1933, IN 1934, IN 1935, IN 1936, IN 1937, IN 1938 
IN 1939, IN 1940, AND IN 1941, AND VELERO IV IN 1949-1952 OFF THE ' COAST OF 
MEXICO AND SOUTHERN CALIFORNIA. 



BRYOZOA OF THE PACIFIC COAST 
OF AMERICA 

Part 3, Cyclostomata, Ctenostomata, 
Entoprocta, and Addenda 



BY 

RAYMOND C. OSBURN, Ph.D., D.Sc. 



The University of Southern California Publications 

Allan Hancock Pacific Expeditions 

Volume 14, Number 3 

Issued August 12, 1953 

Price $4.25 

The University of Southern California Press 

Los Angeles, California 



BRYOZOA OF THE PACIFIC COAST 
OF AMERICA 

Part 3, Cyclostomata, Ctenostomata, 
Entoprocta, and Addenda 

By Raymond C. Osburn, Ph.D., D.Sc. 
Plates 65 - 82 

A report based chiefly on the Bryozoa collected by the Allan Hancock 
Expeditions, 1932-1941, in the Velero III (see pages 1-2 of Part I) and 
in the Velero IV in 1949-1952. 

Additional material received from several sources has greatly enlarged 
the scope of this study. Especially should be mentioned contributions 
from the U. S. National Museum, the collections of the Alaska Crab 
Investigations from southern Alaska, and those from the Point Barrow, 
Alaska, Arctic Research Laboratory. Also practically every museum and 
marine laboratory on the Pacific coast of the United States and Canada 
has contributed some specimens of interest in this extensive survey. 

Order GyGLOSTOMATA Busk, 1852 

Busk in 1852 proposed the name Cyclostomata for this group of 
Bryozoa, since vi^hich time until rather recently it has generally been 
considered a suborder of the order Ectoprocta. In 1926 Borg pointed 
out striking anatomical differences between the Cyclostomata and the 
Cheilostomata-Ctenostomata and gave the former ordinal status under 
the name Stenolaemata. At the same time Borg (1926:490) included in 
the Stenolaemata the old fossil order Trepostomata of Ulrich, but did 
"not wish to give any decided opinion on this point." Later (1944:18-19) 
Borg definitely made the Trepostomata a suborder of the Stenolaemata, 
parallel with the Cyclostomata, and included in it the Horneras, Het- 
eropores, Lichenopores and their allies. 

While it is now generally recognized that the cyclostomes are suf- 
ficiently different from other bryozoans to warrant their separation in 
a distinct order, the merging of the Trepostomata with this order and 
the inclusion of the Horneras, Heteropores and Lichenopores in the 
Trepostomata has not been accepted. The Trepostomes are all Palaeozoic 
and do not occur above the level of the Permian. 

[613] 



614 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

If the Trepostomata are not to be included in the same order with 
the Cyclostomata, there appears to be no good reason for the use of the 
new name Stenolaemata and, at least until there is substantial confirma- 
tory evidence on this question, I prefer to continue the use of "Order 
Cyclostomata." Marcus (1941:12) suggested the name Stenostomata 
to replace Cyclostomata, which has been used also for a group of verte- 
brates, but since the rules of priority are not concerned with ordinal 
names, there seems to be no very good reason for substituting a new 
term for one which has been well-known and acceptable to zoologists 
for a century. 

Borg's separation of the Cyclostomata (1944:20) into five divisions, 
based on anatomical studies, follows closely that of older authors and is 
logical and well-founded, but he seemed to think it necessary to set up 
a whole new series of divisional names. Since it appears that Borg 
simply confirmed, by added histological evidence, the distinctions already 
made in the past, there seems to be no necessity for the discarding of 
well-known terms and the coinage of a new series of divisional names. 

The following table gives a brief digest of the essential characters 
of the five divisions of the Cyclostomata under the old established names, 
with those of Borg in parenthesis, to indicate the synonymy. 

1. Zoarium adnate, suberect or erect, never jointed, the first few tubules, 
at least, always adnate. Wall of the zoarium simple; the ovicell a 
gonozoid varying from simple to broadly expanded and often lobate, 
its polypide degenerating after first reaching maturity. Tubuliporidae, 
etc. (Acamptostega Borg, 1926). ,j^ 

Tubuliporina Hagenow, 1851.^ 

2. Zoarium slender, erect from the first zoid, always jointed, branched, 
rhizoids present. Wall of the zoarium simple; gonozoid simple (some- 
what expanded in Crisulipora) , its polypide degenerating before 
reaching maturity. Crisiidae. (Camptostega Borg, 1926). ' ^Lj 

Articulata Busk, 1859. 

3. Zoarium erect from the beginning, branched tree-like or wine-glass 
shaped, never jointed. Wall of zoarium double, increasing in thickness 
throughout life; gonozoid strongly dilated and usually situated more 

or less on the dorsal side. Horneridae, etc. (Pachystega Borg, 1926). .-, 

Cancellata Gregory, 1896. '<* 

4. Zoarium usually erect, sometimes adnate, often cylindrical and branch- 
ing, composed of autozoids and kenozoids with the apertures of both 
at the surface. Wall double; brood chamber zoarial formed by the 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 615 

absorption of kenozoids around a fertile zoid. Heteroporidae, etc. 
(Heteroporina Borg, 1944). 

Cerioporina Hagenow, 1851. 6^^ 
5. Zoarium adnate or short stipitate, discoid or semiglobular, sometimes 
complex by the formation of subcolonies ; zoids radiating in all direc- 
tions from the center and separated by alveoli (cancelli). Brood- 
j^ chamber zoarial by the fusion of alveoli around a fertile zoid. Lich- 
f^ enoporidae. ( Calyptrostega Borg, 1926). I . ' 

Rectangulata Waters, 1887. -'"^^ 

In this order the older workers based their descriptions and classifi- 
cation almost solely on zoarial characters, and even Hincks in the 1880s 
paid little attention to the ovicells. Waters insisted on the importance 
of the reproductive characters and Harmer, Calvet, Canu and Bassler, 
Marcus, Borg, Silen and others, including the writer, have accepted 
this point of view. 

The difficulty with zoarial characters is their variability, depending 
partly on their adaptation to the substratum and other features of the 
environment, and partly on the stage of growth. In the Crisiidae, the 
younger stages are so much alike that, in the absence of ovicells, the 
determination of the species is often impossible. 

In the encrusting species the nature of the substratum may determine 
the size and form of the zoaria, and the environment often modifies the 
appearance of erect species. Among the Tubuliporidae, encrusting species 
are usually flat and regular on flat surfaces, but when the same species 
develops on a small stem the zoarium may be variously contorted. In 
deeper, quiet water, erect species are usually more slender and more 
elongate, sometimes giving quite a different zoarial appearance from the 
same species in the surf area along shore. In protected areas the peri- 
stomes are usually much more elongate, and in crowded areas or on rough 
surfaces the zoaria may be much reduced in size. Numerous "species" 
have been described on such differences. 

By far the most constant characters in this order are found in the 
ovicells or brood-chambers, either zoidal or zoarial. In the Hetero- 
poridae and Lichenoporidae the brood-chambers are interzoidal or zoarial 
spaces surrounding a gonozoid and their position in the zoarium and the 
gonopores and their tubes (ooeciostomes) are fairly constant. In all the 
others the ovicell is an expanded gonozoid, sometimes only slightly en- 
larged, or again it may be greatly expanded over a considerable portion 
of the zoarial surface and may surround some of the autozoid tubules. 
There may be marked variation in the size and form of these ovicells, 



616 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

even on different parts of the same colony, but the position and form of 
the gonopores (ooeciopores) and their tubes (ooeciostomes) are again 
fairly constant. With most of the species of this order these reproductive 
organs are essential for exact determination, and even here a certain 
amount of caution is necessary. 

The order is ancient, dating back at least to the Ordovician, and the 
number of fossil species that have been described far outnumbers those 
that exist today. How many of these, both fossil and recent, may event- 
ually go into synonymy, no one can even guess, but undoubtedly a very 
large number of them, described from fragments, young colonies, or 
vi^ithout ovicells, may eventually be properly placed. 

Glossary 

A few terms which are not included in preceding glossaries, or which 
have a different use in the Cyclostomata. 

Alveoli. Pores of various sizes distributed between the zoids (see 
cancelli). 

Autozoid. The functional nutritive individuals of the colony. 

Basis rami. A small wedge-shaped base of a branch, characteristic of 
the crisias. 

Brood-chamber. A cavity, usually large, surrounding a gonozoid 
which opens into it (not an expansion of a gonozoid). 

Cancelli. See alveoli. 

Capitulum. An expanded "head" at the tip of an erect branch, usually 
with an ovicell or brood-chamber. 

Disc. The frontal area of a zoarium in the lichenopores ; in complex 
colonies there are often numerous discs. 

Fascicle. A series or bundle of connate tubules or peristomes. 

Gonozoid. A reproductive individual, often greatly expanded to har- 
bor the developing larvae. 

Kenozoid. Various types of greatly modified individuals without poly- 
pides, which serve other purposes in the colony, e. g., the joints of radicles 
in the crisias, and the alveoli (cancelli) in the heteropores and lichencn 
pores. 

Nannozoid. A much reduced individual similar in appearance to an 
autozoid. 

Ooeciopore. The aperture of an ovicell. 

Ooeciostome. The tube surrounding an ooeciopore, the morphological 
end of a gonozoid (ovicell). 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 617 

Pellicle. A thin calcareous layer. 

Proancestrula. A rounded knob formed by the metamorphosis of the 
larva, which buds ofiE the first tubule of a colony. 

Peristome. The projecting end of a tubule (autozoid). Not homol- 
ogous with the peristome of Cheilostomata. 

Radicle. Root-like or pedunculate structure for attachment. 

Radii. Radiating series of tubules, especially in lichenopores. 

Rhizoid. See radicle. 

Subcolony. Branches or areas similar to the primary zoarium pro- 
duced by budding of the zoarium, often very numerous in the licheno- 
pores. 

Tubule. The main part of a zooecium which contains the polypide, 
usually embedded and ending in a "peristome." 

Zoid. A functional nutritive individual. 

Division I. TubuHpoHna Hagenow, 1851 
(Acamptostega Borg, 1926) 

This group appears to be the most primitive among recent Bryozoa, 
as noted by various authors. The zoarium, whether it remains adnate 
or becomes erect, is always adnate at first, the first tubule arising later- 
ally without a joint from the ancestral disc and attached for most of its 
length; and at least a few daughter tubules have this position. The 
resulting zoaria may take almost any form, uniserial or broadly flabellate, 
flat or contorted, semierect or erect, and sometimes profusely branched. 
The apertures ar e always o n the frontal sidei^wjth jhe exception of the 
Entalopkoridae,Iwh ere"lhey _ are dist ributed evenly around the erect 
stein. The ovicells vary from~simple pyriform expansions of the middle 
portion of the gonozoid to very broad, sometimes lobate, expansions 
which may cover considerable areas of the zoarium ; and frequently they 
surround and enclose the erect portions of neighboring tubules. In 
Entalophora and Fasciculipora the ovicells are narrow and greatly 
elongated. 

Key to the Families of Tubuliporina 

1. Zoaria adnate or more or less erect, the apertures opening 

only on the frontal side -u ' * 

Zoaria erect, the zooecial tubes forming cylindrical stems with 

1 • ^ r.r. oil cirlpQ . . Entalophoridae ;oo 
the apertures openmg on all sides ^ f 



618 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

2. Zoaria adnate, linear or flabellate, ovicells simple 3 

Zoaria adnate or erect, usually flabellate but sometimes with 

narrow, erect branches; ovicells expanded 4 

3. Tubules not seriated nor fasciculate; ovicells slender-pyri- . 

form or only slightly expanded Oncousoeciidae \ W 

Tubules single or in small erect fascicles ; ovicell short, 

between fascicles Frondiporidae 

4. Zoaria usually adnate and flabellate, but may be erect and 

branched ; tubules not fasciculate ; ovicells expanded laterally, ,- 

ooeciostome terminal or central Diastoporidae(: '' 

Zoaria adnate, with few exceptions, the tubules usually fas- 
ciculate or in series; ovicell usually ramifying among the ^ ^ 
tubules, sometimes more simple Tubuliporidae^, '' 

Family Oncousoeciidae Canu, 1918 

"The axis of the ovicell is parallel to that of the tubes. The ovicell 
is developed at the same time as the adjacent tubes which are not dis- 
arranged in their respective positions." (Canu and Bassler, 1920:687). 

The ovicell is a simple inflation of the gonozoid, with a terminal or 
sub-terminal ooeciostome, and is often as primitive as that in the genus 
Crisia. The development of the ovicell has some influence on that of 
the adjoining tubes, separating them to some extent, and, in the linear 
species, the tubules at the sides of the ovicell may be increased in number. 

Only three genera with recent representatives concern us here, viz.: 

1. Stojnatopora. Zoarium uniserial throughout, except around the ovi- 
cell which has tubes on both sides, no doubt for additional nourish- 
ment of the developing larv^ae. 

2. Proboscina. Zoarium uniserial for only a short distance at the proxi- 
mal end, then becoming biserial or with additional tubes over most 
of the length of the linear lobes. 

3. Oncousoecia. Zoarium rounded, flabellate or with flabellate lobes; 
the ovicells either narrow or sometimes slightly lobed. 

It is evident that this separation is largely based on zoarial characters 
but the groups present rather distinct facies and it is convenient to treat 
them separately. In Oncousoecia the ovicells are more embedded be- 
tween the adjacent tubules than in the other genera, where they are 
usually ventricose. In all cases the ooeciopore is terminal or nearly so. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA—CYCLOSTOMATA 619 

Genus STOMATOPORA Bronn, 1825 
Alecto Lamouroux, 1821, preoccupied. 

This genus was described and has generally been considered as uni- 
serial, adnate and branching. Some authors have introduced into it 
various linear adnate branching species with a biserial or multiserial 
zoarium, which preferably belong elsewhere. The only part of the 
Stomatopora which is biserial is the expansion about the ovicell. Geno- 
type, Alecto dichotoma Lamouroux, 1821. 

There has been no description of the ovicells until recently, but Borg, 
1926:358, has discovered them in S. eburnea (d'Orbigny) and S. granu- 
late (Milne-Edwards). They are very simple in nature, differing but 
little from those of Crisia except that they are more or less embedded. 

It is true that the zoarium of species of the Tubuliporidae (sens lat.) 
originates in a single zooecium and sometimes the uniserial condition is 
continued for several generations of zooecia before the biserial or multi- 
serial condition is developed. While the generic distinction is not too 
sharply defined, it seems better to retain Stomatopora for the strictly 
uniserial species. 

The three genera, Stomatopora, Proboscina and Oncousoecia have so 
much in common, especially in the nature of the ovicells, that sharp 
distinctions are difficult. For the purposes of the present treatise they 
will be considered as genera on the following zoarial basis: 

Stomatopora, uniserial, except immediately around the ovicell, which 
is simple, inflated, and may be slightly lobate, the ooeciostome terminal. 

Proboscina, biserial, or the linear branches may have several rows of 
tubules, the ovicell simple, inflated, sometimes slightly lobate, the ooeci- 
ostome terminal or nearly so. 

Oncousoecia. broadly multiserial, the zoarium rounded or with fan- 
shaped lobes, the ovicell simple or slightly lobate, more depressed be- 
tween the adjoining tubules, the ooeciostome terminal. 

Stomatopora granulata (Milne-Edwards), 1836 
Plate 65, figs. 1 and 2 

Alecto granulata Milne-Edwards, 1838:205. 
Alecto granulata. Busk, 1875:24. 
Stomatopora granulata, Hincks, 1880:425. 
Stomatopora granulata, O'Donoghue, 1923:11; 1926:17. 
Stomatopora granulata, Borg, 1926:359. 
Stomatopora granulata, Sakakura, 1935:37. 



620 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

The zoarium is adnate, uniserial except around the ovicell which has 
a series of tubules on each side of it, branching more or less at right 
angles. The branches are straight or curved, anastomosing when they 
come into contact. The peristomes curve up from the adnate tubules, 
becoming more or less erect, the diameter at the tip 0.15 to 0.18 mm, 
that of the aperture 0.12 to 0.14 mm; the base of the peristome some- 
what broader according to the amount of calcification. The distance 
from one peristome to the next varies greatly, usually 0.40 to 0.50 mm, 
but may be as much as 1 .0 mm. The tips of younger peristomes are finely 
reticulate; in older parts the peristomes and adnate tubules are irreg- 
ularly roughened. 

The ovicell, first described and figured by Borg (1926:359), is simple, 
the proximal end narrow, irregularly pyriform with small lobes extend- 
ing between the peristomes at the sides. The ooeciostome is an erected 
tube, shorter than a peristome, its tip free and the aperture circular. In 
young ovicells the pore is a semicircular slit at the base of the peristome. 

It appears to be a widely distributed species in the northern hemi- 
sphere, reported from the coasts of Europe from Norway to the 
Mediterranean Sea; Cape Verde Islands (Norman) ; British Columbia 
(O'Donoghue) ; Japan (Sakakura) ; and Waters listed it as a fossil 
from New Zealand. 

Hancock Station 1316-41, off Santa Catalina Island, 45 fms; and off 
San Pedro, southern California, on shells. 

Genus PROBOSCINA Audouin, 1826 

Peristomoecia Canu and Bassler, 1920:692. 

"The zoarium consists of multiserial elongate bands, which are simple 
or branched, and are always flat and adnate. The zooecia are cylindrical 
and narrow. The peristomes are flush with the surface of the zoarium, 
or slightly raised ; and they are usually distributed irregularly, but are 
occasionally quincuncial or in transverse linear series." (Canu and 
Bassler, 1920:658.) Genotype, Proboscina Boryi Audouin, 1826:236. 

This genus differs but little from Stomatopora except in biserial-multi- 
serial disposition of the zooecial tubules. It is true that the zoarium 
begins with a single series of tubules, but this soon becomes expanded 
into two or more series, while in Stomatopora the expansion is limited 
to the area immediately around the ovicell. The mode of branching is 
much the same except that in Proboscina the origin of the branch is 
usually at least biserial while in Stomatopora only a single tubule is 
involved. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 621 

Again there is but little difference in the nature of the ovicells. The 
fertile zooecium arises like any other tubule and is narrow at its proximal 
end, broadens to various degrees in the different species, and terminates 
in a slightly elevated and more or less terminal ooeciostome. The terminal 
expansion of the gonozoid misled Canu and Bassler into believing that 
the enlargement was a part of the peristome, and on this basis they 
created a genus Peristomoecia which has since been discarded. 

The genus also has a close relationship to Oncousoecia Canu and 
Bassler (q. v.), which similarly has simple ovicells of much the same 
nature, but has a flabellate zoarium. 

Whatever may be the ultimate disposition of these three groups of 
species, it seems better for the present to allow them generic status — 
for convenience in classification if for no better reason! 

Large numbers of "species" have been created on incomplete material, 
probably most often on immature specimens. It is practically useless to 
attempt to identify such specimens since only the mature ones with the 
characters of the ovicells show specific characters. 

Proboscina major (Johnston), 1847 
Plate 65, fig. 5 

Alecto major Johnston, 1847:281. 

Alecto major. Busk, 1875:24. 

Stomatopora major, Hincks, 1880:427; 1884:204. 

Stomatopora major, O'Donoghue, 1923:11. 

Diaperoecia major, O'Donoghue, 1926:23. 

Oncousoecia major, Canu and Bassler, 1930:46. 

The zoarium is adnate, strap-shaped, the branches narrow and widen- 
ing slightly, sometimes a little elevated at the tips on rough substrata. 
Zooecia in two to four series, the tubules distinct with well-marked 
grooves, and sometimes in more or less transverse rows; the peristomes 
moderately high, free, with round apertures. The diameter of the aper- 
ture varies greatly, from 0.14 to 0.20 mm and the peristome likewise 
from 0.20 to 0.26 mm on the same colony. 

The ovicells are located near the ends of the branches or just proximal 
to a bifurcation. The narrow proximal end is comparatively short; the 
middle portion expanded and rather bulbous; the ooeciostome sub- 
terminal, erect, rather short, smaller than the peristome, its aperture 
about 0.12 mm in diameter. Usually the inflated area is simple in form, 
but it may be slightly lobed between the surrounding zooecial peristomes, 
and occasionally a peristome may be surrounded. 



1 



622 ALLAN HAXCOCK PACIFIC EXPEDITIONS VOL. 14 

It is a widely distributed species in the northern hemisphere, in Eu- 
ropean waters from Norway to the Mediterranean Sea and the Cape 
Verde Islands, and in the Pacific from British Columbia to the Gala- 
pagos Islands. As a fossil it is known as far back as the Miocene of 
Italy. 

Hancock Stations: 155-34, 324-35 and 450, Albemarle Island, 45 
to 70 fms; 183-34, between Albany and James Islands, 50 to 70 fms, 
and Barrington Island, 52 fms, Galapagos; 328, Cocos Island, Costa 
Rica, 14 fms; 1150-40, 1187-40 and 1316-41, Santa Catalina Island; 
1064, Santa Barbara Island; and 1268-41, Anacapa Island, southern 
California. Also collected by Miss A. E. Blagg at Monterey Bay, Cali- 
fornia, and by Dr. John L. Mohr at the San Juan Islands, Puget Sound. 

Proboscina sigmata new species 
Plate 65, figs. 3 and 4 

A very delicate species. The zoarium is encrusting and consists of 
linear biserial to quadriserial branches with very symmetrical sigmoid 
lateral curves; the curvature is evidently due neither to the substratum 
nor to lateral branching, as there is evidence of only one such branch 
on the outside of a cune. The dorsal side, which is not extended laterally, 
measures about 0.25 mm in width, and is only slightly wider in the 
region of the ovicells. 

The zooecial tubules are narrow, slightly embedded, the separating 
grooves distinct. Their peristomes are thin, about 0.10 mm in diameter, 
very elongate, averaging about 0.65 mm in length but sometimes more 
than 1.0 mm, semi-erect, their walls thin and the aperture about 0.08 
mm in diameter. There is a tendency for the peristomes to arise in 
alternate pairs ; the bases of such a pair may be connate for a short 
distance but the tips are widely divergent. 

The ovicell is an ellipsoid swelling, pointed at the base where it 
disappears among the tubules and narrowed more roundly at the distal 
end where it ends in the terminal ooeciopore, near the base of a peri- 
stome. In the two ovicells in my material there is no evidence of an 
Goeciostome, but this may be due to incomplete development. Length 
of ovicell 0.65 mm, width 0.33 mm. 

Type, AHF no. 57. 

Type locality, ofif Rocky Point, southern California, at 45 fms, on 
the surface of a sunken buoy, Earl Fox, collector, two small colonies, 
both in reproduction. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 623 

Proboscina incrassata (Smitt), 1866 
Plate 66, figs. 1 and 2 

Proboscina incrassata Smitt, 1866:402 and 458. 
Tubulipora (Proboscina) incrassata Smitt, 1871:1119. 
Alecto retiformis Hincks, 1871:81. 
Stomatopora incrassata, Hincks, 1880:436. 
Stomatopora incrassata, O'Donoghue, 1923:11. 
Proboscina incrassata, O'Donoghue, 1926:17. 

The zoarium is white, adnate, much branched, the branches short, 
anastomosing to form often a fairly close network; on the basal part the 
branches are usually two tubules in width, but at the distal ends they 
may be 4 to 6 tubules wide. The peristomes are very irregular in ar- 
rangement, single or 2 or 3 in a transverse line; sometimes a few are 
clustered but they are never connate except occasionally at the base; 
sometimes also these clusters are elevated into short fascicles, especially 
at the ends of branches. The peristomes are usually quite erect, 0.50 to 
1.0 mm in height, 0.26 mm in diameter, and aperture about 0.20 mm. 

The ovicells, which seem not to have been noticed previously, are 
simple ventricose expansions near the ends of branches and surrounded 
by a row of tubules on each side (occasionally a peristome may be en- 
closed in the expansion) ; the ooeciostome is a small erect tube much 
shorter than the peristomes, terminal or nearly so, usually connate with 
a peristome at its base but the tip always free, the aperture round and 
about 0.13 mm in diameter. 

Described from Spitsbergen and recorded from Norway, Nova Zem- 
bla, Kara Sea, and the British Islands from Cornwall to Scotland and 
the Shetland Islands. On the Pacific coast O'Donoghue listed it for 
several localities in British Columbia. Point Barrow, Alaska, Arctic 
Research Laboratory, 328 feet, a common species, especially on stones, 
G. E. MacGinitie, collector. 

Proboscina lamellifera Canu and Bassler, 1930 
Plate 66, fig. 3 

Proboscina lamellifera Canu and Bassler, 1930:46. 

"The zoarium incrusts shells and is formed of sinuous branches joined 
together by a smooth calcareous lamella. The tubes are indistinct, short, 
seriated and terminated by a long peristome perpendicular to the zoarial 
plane. Measurements, — diameter of orifice, 0.12 mm; diameter of 
peristome, 0.16 mm; internal separation of tubes, 0.20 - 0.30 mm; width 
of branches 1.5 mm." (Canu and Bassler, 1930:46). 



624 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

This short description by the above authors and their photographs of 
the species (Plate 11, figs. 1 and 2) are sufficient for identification, 
though they did not have the ovicell. In our specimens the lamella, while 
it extends somewhat from the borders, does not come anywhere near 
connecting them. This may be due to the fact that our colonies are much 
smaller and probably younger. 

Four ovicells are present in our specimens. They are small, short and 
very bulbous, raised as high as the peristomes surrounding them, thick- 
walled and shining; the ooeciostome is a short, round tube, its diameter 
nearly that of the peristomes, and terminal, which, in this case, is nearly 
on the top of the semiglobular ovicell. One of the ovicells is slightly 
enlarged and its border encloses two peristomes. 

Described from Albatross Station D. 2813, Galapagos Islands. 

Hancock Stations: 143-34, off Wenman Island, 1°23'10"N, 91°48' 
45"W, at 100-150 fms; 155-34, off Tagus Cove, Albemarle Island, 
0°16'45''S, 91°22'52"W, at 50-60 fms; and 453, Gardner Island, 35 
fms, Galapagos Islands. 

Genus ONCOUSOEGIA Canu, 1918 

The zoarium is adnate, broadly multiserial, rounded or with flabellate 
lobes; the zooecial tubes are long, distinct on the surface, quincuncially 
arranged, the peristomes short and more or less erect. The ovicell is 
simple, often differing only slightly from the zooecial tubules, the proxi- 
mal end embedded between the neighboring tubules, distally expanding 
gradually between the adjacent tubules and sometimes extending laterally 
above them for a short distance. The ooeciostome is terminal, not asso- 
ciated with a peristome, usually short, erect, smaller than the peristomes, 
round, and not expanded at the tip. 

The genotype is Alecto dilatans Thompson (Johnston, 1847:281), 
and not Tubulipora lobulata Hincks, 1880, as indicated by Canu, 1918: 
325. Hincks confused two species in his Plate 61, and the figure 5, 
from which Canu evidently drew his description, is that of Alecto 
dilatans Thompson. (For details see Osburn, 1933:9-12). 

Oncousoecia diastoporides (Norman), 1868 

Plate 66, fig. 4 

Alecto diastoporides Norman, 1868:310. 
Stomatopora diastoporides, Hincks, 1880:434. 
Stomatopora diastoporides, Osburn, 1912:218. 
Oncousoecia diastoporides, Osburn, 1933:9. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 625 

Zoarium a flat fan-shaped or lobulate incrustation on shells and stones ; 
moderately thick, usually with two rows of incomplete zooecia bordering 
the outer functional ones. The zooecial tubules are elongate and hori- 
zontal for most of their length, about 0.30 mm wide, convex and the 
separating grooves distinct, finely punctured. The peristomes are short, 
suberect, quincuncial in arrangement, the aperture about 0.15 mm in 
diameter. 

The ovicells resemble the zooecial tubules, elongate, pointed at the 
proximal end, only a little swollen and more thickly punctate ; they were 
overlooked for many years, probably because of their resemblance to the 
tubules, but they are definite enough when one knows what to look for. 
The ooeciostome is terminal, short, erect, round and about 0.08 mm in 
diameter, not associated with a peristome. 

Described from the Shetland Islands by Norman and recorded also 
by Hincks from the British Islands. On the west coast of the North 
Atlantic it ranges from Cape Cod northward to Mount Desert Island, 
Maine, to the Gulf of St. Lawrence and Baffin's Bay. 

Point Barrow, Alaska, Arctic Research Laboratory, G. E. Mac- 
Ginitie, collector; also at Canoe Bay, southern Alaska, U. S. Alaska 
Crab Investigation, Sta. 25-40 at 25 fms. 

Oncousoecia canadensis Osburn, 1933 

Plate 65, figs. 10 and 11 

Oncousoecia canadensis Osburn, 1933:12. 
Stomatopora diastoporides, Whiteaves, 1901:110. 

The zoarium is flabellate or irregularly lobate, entirely adnate on 
shells and stones, the primary region, 2 or more tubules in width, is 
usually short ; thinner than in O. diastoporides. The tubules are com- 
paratively thin-walled, somewhat hyaline and vitreous, conspicuously 
perforated. They are more slender than those of diastoporides (width 
about 0.18 mm), and never does more than one row of incomplete ones 
appear at the margin. The peristomes are short, thin-walled, the aper- 
ture about 0.10 mm in diameter, and never connate nor seriated. 

The ovicells are usually like small thin-walled blisters; the fertile 
zooecium arises in the same manner as the infertile ones but soon ex- 
pands both frontally and laterally and the adjacent tubules appear as if 
separated by the growth of the expansion. Sometimes the expansion 
extends in very short lobes on either side of the ooeciostome, but occa- 
sionally it may be as simple as in diastoporides. The ooeciostome is 



626 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

situated terminally, between but not in contact with the adjacent peri- 
stomes; it is rounded or slightly elliptical transversely, short, erect or 
slightly bent proximally; the aperture 0.06 mm in diameter. 

Described from Mount Desert Island, Maine, and recorded by Osburn 
from the Bay of Fundy and the Gulf of St. Lawrence (for details see 
Osburn, 1933:13). 

Point Barrow, Alaska, Arctic Research Laboratory, to a depth of 
50 fms, G. E. MacGinitie, collector, common on shells. 

Oncousoecia ovoidea new species 
Plate 65, figs. 8 and 9 

The zoarium is broadly flabellate; the ancestrula produces a single 
short tubule; from this two tubules arise, and then four tubules begin 
the wide expansion. They are arranged in quincunx, the peristomes all 
well separated. The embedded tubules are about 0.13 mm in diameter, 
the peristomes 0.11 mm and the apertures 0.08 mm. The tubules are 
very definitely cross striated, the peristomes smooth and semierect, the 
longest ones 0.13 mm. 

There are two ovicells on one colony, the expanded portion ovate, 
ventricose, 0.15 to 0.18 mm in width, the distal end rounded, the ooecio- 
stome terminal, short, erect, the aperture rounded and 0.05 mm in 
diameter. The proximal part of the gonozoid is like the other tubules 
for about half or two-thirds of the length and the expansion appears 
suddenly. 

Type, AHF no. 58. 

Type locality, Hancock Station 276, San Esteban Island, Gulf of 
California, 28°38'30"N, 112°36'W, at 32 fms. Two colonies encrust 
smooth shell fragments, while a third is on the rough pebbled surface 
of an echinoid spine; the largest colony is less than 2 mm in width. 

Oncousoecia abrupta new species 

Plate 65, figs. 6 and 7 

The zoarium is small, delicate and entirely adnate ; the proximal por- 
tion very slender (0.45 mm at the widest part) for a distance of 5 mm, 
then abruptly becoming broad and round, about 2.5 mm in either direc- 
tion. The proximal part, just above the pro-ancestrula, which is wanting, 
is 2 tubules in width, alternating, and widens to 4 tubules near the 
expansion; the peristomes moderately short and free. On the expanded 
area the peristomes are much higher (to 0.50 mm), rather regularly 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 627 

distributed and not seriated, free with a few exceptions where two are 
connate at the base, slender with an outside measurement of 0.09 mm. 
The apertures are round and about 0.07 mm in diameter. The tubules 
are so much embedded that their measurements cannot be determined 
the surface with minute pores and light transverse striae. 

There are 3 ovicells side by side, occupying practically the full width 
of the terminal border, each broadly pyriform and with short lobes be- 
tween the surrounding peristomes; a little ventricose and cross-striated 
and the surface with minute pores. The ooeciostome is erect and mode- 
rately high, situated beside a peristome with which it is connate at the 
base and the upper portion free, its base cylindrical and about 0.05 mm 
in diameter, the tip expanded transversely and its aperture measuring 
about 0.09 by 0.03 mm. 

The basal portion alone would readily be mistaken for a species of 
Proboscina, but the expanded part is similar to that of Oncousoecia in 
the nature of the ovicells and ooeciostomes. 

Type, AHF no. 88. 

Type locality, off Rocky Point, southern California, about 33°49'N, 
encrusting a sunken buoy recovered from a depth of 45 fms, one colony 
by Earl Fox. Also one colony collected at Santa Barbara Island by 
Dr. H. R. Hill. 

Family Diastoporidae Gregory, 1899 

Diastoporn Lamouroux, 1821; Berenicea Lamouroux, 1821; Mesen- 
teripora Blainville, 1830; Bidiastopora d'Orbigny, 1849; Actinopora 
d'Orbigny, 1853; Microecia Canu, 1918; Plagioecia Canu, 1918; Di- 
aperoecia Canu, 1918; Diplosolen Canu, 1918. 

Diastopora, genotype D. foliacea Lamouroux, 1821 :42, though it was 
described from a fossil without ovicells, has rather definite zoarial char- 
acters and has been much used for recent as well as fossil species. In 
the absence of an ovicell, however, it is impossible to place this genus 
except as a member of the present group. It is useful to the paleontolo- 
gists when ovicells are wanting, but should not be used when the ooecial 
characters are present. 

Berenicea, genotype B. prominens Lamouroux, 1821 :80, is so indefi- 
nite as to be meaningless. Norman, 1903:569 and 1909:299, and Borg, 
1944:61, have maintained that Berenicea is not even a cyclostome but a 
cheilostome form. Diastopora and Berenicea have been used rather 
indiscriminately for the same species. If they are synonymous, Dt- 



628 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

aperoecia takes precedence by its earlier appearance in Lamouroux's 
work, but owing to the uncertainty as to its nature Berenicea had better 
be discarded. 

Mesenteripora, genotype M. micheUni Blainville, 1830:397, was pro- 
posed for erect contorted forms otherwise similar to Diastopora. Here 
again the genotype is a fossil without ovicells. 

Bidiastopora, genotype Diastopora ccrvicornis Michelin, 1846:241, 
was founded to include erect diastoporas with bilaminate folds, and the 
genotype is a fossil without ovicells. 

Actinopora, genotype A. regularis d'Orbigny, 1853:763, was based 
on the arrangement of the zoids in regular radiating series and like the 
preceding included only fossil species without ovicells. The ovicelled 
species which have later been placed in this genus have the ovicells and 
ooeciostomes definitely of the Plagioecia pattern. 

The above generic names were all properly founded but only on 
zoarial characters, so their complete nature is uncertain. They may still 
be useful when it is necessary to catalog specimens which are incomplete 
in reproductive characters. 

In 1918 Canu attacked the problem of the old Diastopora complex 
with the ovicells as the basis, and proposed a number of additional 
genera, Alicroccia, Plagioecia, Diapcroecia and Diplosolen. 

Microecia Canu, 1918:326, was mistakenly founded on Diastopora 
Sarniensis Norman, 1864:89, and is synonymous with Plagioecia. 

The other three genera agree in the mode of early development in 
the tubuliporoid manner, in the closure of numerous older peristomes 
by a calcified porous membrane, and by at least the occasional enclosure 
of peristomes by the ovicell. 

Canu went so far as to propose several new families, Mecynoeciidae, 
Plagioeciidae and Diaperoeciidae among the diastoporid forms. The first 
of these, which included Microecia, has already been reduced to syn- 
onymy by Bassler, 1935:10. While Canu's analysis of the ovicell is of 
the greatest importance in the separation of species of this group, he 
apparently was not sufficiently familiar with the intraspecific and inter- 
specific variation in the ooecia and ooeciostomes. 

For the purpose of the present work I propose to accept only the old 
family Diastoporidae with the following genera: 

1, Diastopora Lamouroux, 1821, reserved for species in which the 
reproductive characters are unknown. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 629 

2. Plagioecia Canu, 1918, ovicell usually much broader than long, 
not proliferated beyond the level of the ooeciopore ; ooeciostome terminal, 
at or near the middle of the distal border ; one or more peristomes often 
surrounded by the lateral prolongation of the ovicell. 

3. Diaperoecia Canu, 1918, the ovicell completes its development by 
proliferating distally in advance of the ooeciopore and surrounds few 
or many distal peristomes in the process; the ooeciostome, which repre- 
sents the morphological distal end of the gonozoid, is usually located 
somewhere near the middle or occasionally even near the proximal end 
of the ovicell. 

4. Diplosolen Canu, 1918, miniature zooecia (zooeciules) present, 
scattered among the normal tubules of the zoarium; ooeciostome sub- 
terminal at or near the middle; peristomes occasionally surrounded. 
Diplosolen differs from Plagioecia in appearance only by the dimorphic 
nature of the zooecial tubules. 

It is sometimes difficult to assign a species to one of the above genera, 
owing to variation in the size and form of the ovicell and especially in 
the occasional occurrence of ovicells as simple as those of Oncousoecia. 
When these occur on the same zoarium with more highly developed 
ovicells, the latter has been accepted as the proper generic association. 
The simple ovicells usually occur on the older parts of the colony and 
if only this form of ovicell is present the species would necessarily he 
assigned not only to another genus but to a different family. A rather 
exaggerated case of this is found in Plagioecia ambigua new species 
(q. v.), but examples may be found in other species of this genus and 
also in Tubulipora. The enclosure of peristomes by the ovicell, on which 
Canu (1918) based the genus Diaperoecia, is subject to much variation, 
and this condition is also found to a greater or less extent in Plagioecia 
and several genera of the Tubuliporidae. 

In spite of these variations, where a fully developed ovicell is present, 
the position of the ooeciostome is usually diagnostic, median and terminal 
or subterminal in Plagioecia, more centrally located in Diaperoecia, m 
which genus the ovicell continues to develop distally beyond the ooecio- 
stome. The ooeciostome must be considered the morphological distal end 
of the gonozoid. 

Genus PLAGIOECIA Canu, 1918 

"The ovicell is a long transverse sack obliterating a certain number of 
zooecial tubes and developed in the vicinity of the zoarial margins. The 
ooeciostome is small, equal to or less than the zooecial diameter. The 



630 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

tubes are isolated from each other. No adventitious tubes." (Canu and 
Bassler, 1922:26). Genotype, Tubulipora patina L,aimrck, 1816. 

The above description, drawn from P. patina and correct for that 
species, requires some modification as the ovicells are not always much 
expanded laterally and sometimes in other species may even be longer 
than broad. The ovicells are symmetrically developed and the ooeciostome 
is terminal in the midline at or near the distal border and the ooecial 
expansion is not continued beyond it. The inflation often surrounds one 
or more, sometimes several, peristomes; even in the same species the 
shape of the ovicell and the number of the included peristomes may vary 
considerably due to the amount of lateral expansion. There is much 
closure of the older peristomes by a calcified membrane which is either 
perforated by a number of small pores, or provided with a small erect 
central tubular pore. 

The zoarium is usually adnate, but may be erect or semierect and 
more or less contorted and either unilaminar or bilaminate. 

Key to the Species of Plagioecia 

1. Zoarium more or less free, erect or semierect 2 

Zoarium entirely adnate or sometimes free at the edges ... 4 

2. Zoarium with narrow, bilaminate, contorted branches which 

form a free reticulum meandrina 

Zoarium with broad bilaminate lobes, contorted 3 

3. Zoarium erect from a rather narrow base, sometimes stipitate, 

contorted, the folds thicker than in other species . . grimaldii 
Zoarium beginning with a broad adnate base from which erect, 
contorted folds arise, much thinner than in P. grimaldii . . . 

tortuosa 

4. Except near the center of the zoarium the peristomes are in 

uniserial, radiating rows, connate only at the base . anacapensis 
Peristomes not in radiating series 5 

5. Zoarium broad and thin, irregular, with occasional smooth 

areas containing aborted tubules without apertures; peri- 
stomes very short; ovicell low and flat, surrounding a few 

peristomes, irregularly rounded tubiabortiva 

Zoarium rounded or with flabellate lobes, ovicell conspicuous 

and usually transverse 6 

6. Ovicell usually several times as broad as long and slightly 

arcuate to conform to the zoarial border, the lateral ends 

often enclosing peristomes patina 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 631 

7. Zoarium irregularly rounded, peristomes thin, 0.09 mm at 

their tips; ovicell only moderately broad sarniensis 

Zoarium lobate, peristomes 0.13 mm at their tips; in the one 
specimen there are two simple, narrow ovicells and one ex- 
panded one, the ooeciostome slightly subterminal . . ambigua 

Plagioecia patina (Lamarck), 1816 
Plate 73, fig. 4 

Tubulipora patina Lamarck, 1816:163. 
Diastopora patina, Hincks, 1880:458; 1884:206. 
Diastopora patina, O'Donoghue, 1923:14. 
Plagioecia patina, O'Donoghue, 1926:21. 

Zoarium variable in form, rounded or lobate; entirely encrusting, or 
partially free. The zoids are embedded for most of their length, the 
free part of the tubules, "peristomes," being semierect and usually short. 
The apertures are somewhat elliptical or nearly round, measuring about 
0.10 by 0.12 mm. The embedded portions of the zoids are slightly convex 
and are perforated by minute pores. The peristomes show no tendency 
to be arranged in series, and are not connate even at the base. In older 
parts of the colony, especially, the apertures become closed by a peculiar 
calcified membrane which is perforated either by a number of pores or 
by a single larger tubular pore. The basal lamina often forms a distinct 
border beyond the functional zoids. 

The ovicell is a prominent swelling, moderately large and distinct, the 
edges usually sharply outlined. Normally it is transversely very elongate, 
several times as wide as long, but varying considerably in size and form. 
Usually a number of peristomes are surrounded, to 7 in my specimens. 
The ooeciostome is terminal and free between the zooecial series, ordi- 
narily occupying a small notch in the middle of the distal side; it is 
rather short and either erect or flexed slightly toward the proximal part 
of the zoarium, the aperture rounded and 0.06 to 0.08 mm in diameter. 

This well-known Atlantic species resembles P. sarniensis in its general 
appearance, but the zooecia are distinctly larger with shorter peristomes, 
and the ovicell is much wider transversely. On the Pacific coast it was 
first noted by Hincks at Cumshewa, and later by O'Donoghue at Bull 
Passage, British Columbia. 

Hancock Stations: 143-34, Wenman Island, 100 fms, 147-34, Albe- 
marle Island, 30 fms, and 352-35, Chatham Island, 35 fms, Galapagos; 
299, San Jose del Cabo at the southern tip of Lower California, 82 
fms; 72, Guadalupe Island off Lower California, 17 fms; Santa Barbara 



632 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

and San Miguel Islands and various other places off shore along southern 
California, 15 to 76 fms. Also collected in Puget Sound, Washington, 
by Dr. J. L. Mohr, and two colonies from Cleveland Passage, Frederick 
Sound, southern Alaska. 

Plagioecia sarniensis (Norman), 1864 
Plate 73, fig. 3 

Diastopora Sarniensis Norman, 1864:89. 
Diastopora sarniensis, Hincks, 1880:463; 1884:206. 
Berenicea sarniensis, Harmer, 1915:114. 
Microecia sarniensis, O'Donoghue, 1926:21. 
Plagioecia sarniensis, O'Donoghue, 1926:22. 

The zoarium is usually encrusting but sometimes the borders are free 
and slightly contorted ; the basal lamina forms a distinct border. There 
is much resemblance to P. patina in the zoarial form, but the smaller 
size of the zooecia and the form of the ovicell easily distinguish them. 

The zooecia are embedded for most of their length. The semierect 
"peristomes" become suddenly smaller, their diameters only 0.09 mm 
and their apertures 0.07 mm in diameter. The peristomes are usually 
longer than those of patina, never connate and not in series. 

The ovicell varies in form from irregularly rounded to short trans- 
verse, occasionally somewhat bilobate, and often one or two peristomes 
are surrounded. The ooeciostome is terminal or sub-terminal at the 
distal border, isolated, erect or curved proximally, the ooeciopore round 
and 0.05 mm in diameter. 

Norman and Hincks both figured a small oval or rounded ovicell, 
though the latter states (1880:463) "Ooecia transversely elongate, 
subelliptical inflations of the zoarium, of a considerable size." Doubt- 
less it was the small size of the ovicell figured that led Canu ( 1918 :326) 
to select this species as the genotype of his new genus Microecia, which 
he placed in his new family Mecynoeciidae, now discarded. If there is 
such a fossil group of species of generic value, the selection of sarniensis 
as the genotype was most unfortunate and the generic name Microecia 
is invalidated, for sarniensis is certainly congeneric with patina. While 
the ooecial characters are of the greatest importance in the study of the 
cyclostomatous species, it is necessary to recognize the fact that these 
characters, like all others in nature, are subject to variation, and this is 
especially true of the size and form of the ooecial expansion. I have seen 
several cases of simple ooecia on the same zoaria with those of larger 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 633 

size in sarniensis, and in patina there is much variation in width. 

P. sarniensis has now been found in so many parts of the world that 
its distribution must be considered cosmopolitan. Hincks first listed it 
for Pacific waters at Cumshewa, British Columbia, and O'Donoghue 
recorded it from Banks Island and Lowe Inlet, British Columbia, and 
the San Juan Islands, Puget Sound. 

Hancock Stations: 484, Barrington Island, Galapagos, 0°49'S, 90° 
06'40nV, 52 fms; 423-35, off Port Utria, Colombia, 5°59'20"N, 77° 
2r50"W, at 20 fms; 276, San Esteban Island, Gulf of California, 
28°38'50"N, 112°36'W, at 32 fms; 72, Guadalupe Island, oflF Lower 
California, 29°N, at 17 fms; Santa Barbara, Anacapa and San Clemente 
Islands, of? southern California; and San Juan Islands, Puget Sound, 
Washington. 

Plagioecia tortuosa new species 
Plate 67, figs. 8 and 9 

Mesenteripora meandrina, Robertson, 1910:251 (not Wood, 1844:14). 

Dr. Alice Robertson has given an excellent description of the zoarium : 
"Zoarium bilaminate, forming a contorted, convoluted mass . . . begin- 
ning as a simple, primitive disk from which there grow tubular zooecia 
curving in opposite directions, and forming a fan-shaped expansion 
similar to any young tubuliporidian colony. The two layered condition 
results from the ridges which occur at irregular intervals over the 
unilaminar sheet, . . . and which growing upward form the erect, 
bilaminar layers, the laminae becoming highly convoluted," The en- 
crusting base sometimes covers a considerable area before the bilaminate 
folds are formed. 

The zooecia are alternate, in quincunx, embedded but with the frontal 
surface convex, with numerous pores and sometimes transversely ribbed. 
The erect tubules or "peristomes" are usually short but may be as much 
as 0.50 mm in length, narrowing only slightly, perforated only near the 
base, about 0.13 mm in outside diameter; the aperture short oval or 
round and about 0.10 mm in diameter. 

The ovicell, partially described by Robertson, is a distinct inflation 
which is usually considerably broader than long, surrounding 6 to 12 
peristomes. The ooeciostome, which Robertson was unable to find, is 
sub-terminal, median, somewhat removed from the distal border, short, 
erect and slightly expanded at the tip, the pore round, 0.08 mm m 
diameter and the tip expanded to 0.13 mm. 



634 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

In all characters, zoarial and reproductive, except the bilaminate adult 
zoarium, this species agrees closely with P. patina and must be considered 
congeneric with it. 

Robertson recorded it from three localities in southern California, 
down to a depth of 32 fms. 

Type AHF no. 107. 

Type locality, Hancock Station 1662-48, off Santa Cruz Island, 
southern California, 33°55'45''N, 119°31'05nV, at 23 fms. Also taken 
at 1130-40, off Laguna Beach, southern California at 25 fms; at Cortez 
Bank, 32°24'N, 119°22'30"W, at 131 fms; and at 1190, Puerto Escon- 
dido, Gulf of California, 25°48'04''N, lll°18'53nV, in shallow water. 
Another fine specimen from Station 275, Raza Island, Gulf of Cali- 
fornia, 28°48'N, 113°W, at 40 fms has 4 ovicells more or less centrally 
located and 3 others partially developed near the margins of the zoarium. 

Plagioecia grimaldii (Jullien), 1903 
Plate 66, fig. 5 

Mesenteripora Grimaldii Jullien, 1903:118. 

Plagioecia grimaldii, Osburn, 1936:540. 

? Mesenteripora meandrina, Smitt, 1866:432. 

The zoarium consists of erect contorted folds arising from an encrust- 
ing base to a height of 1 or 2 cm. The folds are bilamellar, the growing 
edge showing the basal lamina with the tubules arising on both sides. 
The colony may be stipitate, as in Jullien's figure, plate 15, fig. 4, but 
is often broad and irregular. The embedded tubules are convex and quite 
distinct on the surface of the zoarium, 0.25 to 0.30 mm in width, with 
moderately deep separating grooves and perforated by numerous small 
pores. The peristomes are usually very short, often rising scarcely above 
the zoarial surface, but in protected areas they may rise, semierect, to 
a length of 0.40 mm. The apertures vary considerably, from 0.14 to 
0.18 mm and are often closed by the characteristic diaphragm with a 
small tubule at the center. 

The ovicells, here described from Baffin Bay specimens, are variable 
in size and form, large enough to surround 5 or 6 peristomes, prominent 
and sharply outlined ; the ooeciostome smaller than a peristome and 
scarcely elevated above the surface, median and terminal in position. 
There is a tendency for the ovicells to be slightly broader than long but 
in one case the^ length is 50% greater than the width; another much 
smaller ovicell, which encloses only one peristome, is round. Since so 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 635 

much attention has been given to the shape of the ovicells by Canu and 
Bassler it is important to note the amount of variation in form. 

Jullien described the species from the Grand Bank of Newfoundland, 
at 155 meters. The only other positive reference is that in Osburn's 
report on dredgings by Captain R. A. Bartlett in Baffin Bay, three 
colonies w^ith ovicells, (previously unknovi^n), at 140 to 210 feet. It is 
probable that Smitt's Mesenteripora meandrina in the Torell collection 
from Greenland (1866:432) should now be referred to gnmaldit rather 
than to the fossil Diastopora meandrina of Wood. 

Point Barrow, Alaska, Alaska Research Laboratory, at 217 feet, G. E. 
MacGinitie, collector, several fragments agreeing with Baffin Bay speci- 
mens in all other details but without ovicells. The range of distribution 
is evidently high northern and probably circumpolar. 

Plagioecia meandrina (Canu and Bassler), 1930 
Plate 66, figs. 6 and 7 

Diaperoecia meandrina Canu and Bassler, 1930:51. 

The zoarium has a very striking appearance, consisting of a broad 
encrusting base from which arise at intervals narrower bilaminate 
branches or fronds which often anastomose to form large quadrangular, 
pentagonal or hexagonal fenestrae. The branches are usually at right 
angles to the plane of the zoarium. On the encrusting base the zooecia 
are arranged in quincunx, but on the erect branches they tend to run 
in rather regular series more or less transverse to the branch; they are 
not connate but well separated. The peristomes of the base are short 
but on the branches, especially near the growing edge, they are mod- 
erately elongate and nearly erect. The basal lamina of the base extends 
rather broadly beyond the functional zooecia and on the branches there 
is a similar but much narrower lamina projecting from between the two 
zooecial layers on one edge of the branch. The zooecial tubules are very 
little inflated and their outlines are often obscure. The peristomial 
apertures are round and about 0.10 mm in diameter. 

The ovicell is a distinct inflation, irregularly elliptical, transverse and 
parallel to the edge of the branch and surrounding a number of the 
peristomes, most of which are closed, like those of P. patina, with a 
calcified membrane in the middle of which is a minute tubule. The 
ooeciostome is small, short, nearly erect, situated near the middle and 
terminal, free between the peristomes, and measures 0.08 mm. 



636 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Canu and Bassler placed the species in their genus Dtaperoecia be- 
cause of the peristomes surrounded by the ovicell, but indicated that 
"It is not yet a true Diaperoecia." As a matter of fact, the ovicell is 
almost exactly like that of Plagioccia patina in form and location as well 
as the nature of the ooeciostome. The perforation of the ovicell by the 
peristomes sometimes occurs in its relatives, patina and sarniensis, and 
in a specimen of P. (Microecia) tubiahortiva (Canu and Bassler, 1930) 
I have observed as many as 8 such enclosed peristomes. The closure of 
the peristomial apertures also is exactly like that in patina, a porous 
calcified membrane with a minute short tubule at the middle. 

Our best developed specimen measures about 35 mm across the en- 
crusting base and the fenestrate erect portion is about 60 mm high and 
wide, with 7 complete fenestrae. In most cases the growing edges of 
the branches are oriented in the same direction. 

Described from the Galapagos Islands, Albatross Station D. 2815. 

Hancock Stations: 143-34 Wenman Island; 170-34, Chatham Island; 
201-34, Hood Island; 450, Albemarle Island; 452, Charles Island, and 
453, Gardner Island, all from the Galapagos. Also at 1662-48, Santa 
Cruz Island, southern California; and collected by Dr. Carl L. Hubbs 
at Guadalupe Island, ofl[ Lower California. The geographic range is 
wide, from Santa Cruz Island, southern California (33°35'45''N) to 
Hood Island, ( 1°21'55''S), and the bathymetric range from 23 to more 
than 100 fms. 

Plagioecia tubiahortiva (Canu and Bassler), 1930 
Plate 73, fig. 2 

Microecia tubiahortiva Canu and Bassler, 1930:48. 

The zoarium is broad and flat, with a very irregular outline; the 
surface even, with smooth areas free from apertures and consisting of 
aborted tubules. The zooecial tubes are completely immersed, except for 
the very short, semierect peristomes which usually project only slightly 
above the crust. The diameter of the peristomes is 0.12 or 0.13 mm, that 
of the apertures 0.10 or 0.11 mm. The aperture is rounded to slightly 
elliptical. The peristomes are irregularly spaced, never connate and not 
in series. The basal lamina usually forms a distinct border. 

The ovicell is a low inflation, rounded, expanded laterally or irregular 
in outline, usually surrounding a few peristomes (8 in one case). The 
ooeciostome rises barely above the surface, median and terminal in posi- 
tion, its aperture measuring about 0.06 mm. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 637 

This species is evidently congeneric with patina and sarniensis, judging 
by the nature of the tubules and especially by the characters of the 
ovicell. 

Described from the Galapagos Islands, Albatross Station D. 2813. 

Hancock Stations 143-34, oflF Wenman Island, Galapagos, 1°23'10"N, 
91°48'45"W, at 100 to 150 fms, several colonies on shells. 

Plagioecia anacapensis new^ species 
Plate 66, figs. 9 and 10 

? Dtastopora catillus J. Y. Johnson, 1897:61. 

The zoarium is round and flat but the margin is more or less turned 
up to produce small saucer-shaped colonies, which are attached by a 
comparatively small peduncle. The basal lamina is of moderate width. 
The zooecia about the center, with short peristomes, are arranged quin- 
cuncially, but beyond this area they form uniserial radiating rows which 
extend to the margin, similar to the fossil Unitubigera of d'Orbigny, 
1853. Additional shorter series are interpolated toward the margin. The 
tubules are embedded, convex on the frontal surface, the walls perforated 
and later often transversely ribbed. The peristomes are semierect and 
beyond the central area become longer (0.25 mm or more). The central 
peristomes are always free and isolated, those in the radiating series 
sometimes connate at the base but the tips always free; diameter at the 
tip 0.13 mm, the aperture round and 0.10 mm in diameter. 

The ovicell is inflated, its outlines distinct, transversely elongate 
(usually more than twice as wide as long), surrounding one or more 
peristomes. The ooeciostome is terminal at the middle of the distal 
border, short, erect, its rim flared like the bell of a cornet, the ooeciopore 
round and 0.08 mm in diameter, the rim circular and about 0.13 mm 
across. 

When this material was first examined I placed it at once under 
Unitubigem d'Orbigny, 1853, but the large non-seriated central area 
is different and the nature of the ovicells (unknown in Unitubigera) is 
distinctly like that of Plagioecia patina. 

There is a possibility that this species may be the Diastopora catillus 
of J. Y. Johnson (1897:61) from Madeira; his description is fairly 
similar, but he did not give a figure and did not mention the ovicell. 

Type, AHFno. 113. 

Type locality, Hancock Station 874-38, off Anacapa Island, southern 
California, 34°0r30''N, 119°21'W, at 45 fms, one colony on a shell. 
Also two colonies recovered from a sunken buoy off Rocky Point, south- 



638 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

ern California, 45 fms. One of the latter is almost exactly like the 
type specimen except that the ovicell encloses three peristomes. The 
other colony diflfers only in having the 3 or 4 marginal rows of peristomes 
suddenly much elevated. 

Plagioecia ambigua new species 
Plate 66, fig. 8 

The zoarium is flat and thin, entirely adnate, encrusting the smooth 
surface of a shell. The proximal portion is narrowly flabellate, with a 
very simple ovicell ; beyond this the zoarium becomes broadly flabellate 
with a similar simple ovicell at one side and a very broad ovicell occupy- 
ing much of the width of the lobe. The zooecial tubes are elongate, 
moderately distinct on the surface, slightly cross-striated and perforated 
with small pores, 0.20 mm in width. The peristomes are only suberect 
and directed strongly forward, the diameter 0.15 and the round aperture 
0.13 mm; arranged in quincunx. There is only a single row of incom- 
plete tubules at the margin. 

The simple proximal ovicell is about 0.40 mm in width by 0.75 mm 
long; the simple lateral ovicell 0.30 mm wide by 0.70 mm long and the 
large ovicell is about 1.60 mm broad by 0.80 mm long. The ooeciostome 
is terminal in the small ovicells, erect at the proximal side of a peristome; 
in the large ovicell the ooeciostome is similarly situated, but is somewhat 
subterminal as the ooecial cavity has extended slightly beyond it ; diameter 
of aperture 0.08 mm. 

Type, U. S. Nat. Mus. no. 11049. 

Type locality. Point Barrow, Alaska, 130 feet, Arctic Research Lab- 
oratory, G. E. MacGinitie, collector, one colony. 

This is a very unusual specimen, with characters of several genera. 
There are two simple ovicells like slightly expanded zooecial tubules 
and with terminal ooeciostomes, much like Oncousoecia diastoporides, 
except that the ooeciostome is associated with a peristome. There is also 
a much expanded ovicell, transverse, surrounding several peristomes, 
with a subterminal ooeciostome and resembling Plagioecia, except for 
the position of the ooeciostome proximal to a peristome. The latter 
character is more like that of Tubulipora. The ovicell surrounds a num- 
ber of peristomes, which would place the species under Diaperoecia. 

With such a combination of characters, one is naturally in doubt as 
to the generic relationship, but I am accepting as the most important 
character the fullest development of the ovicell, expanded laterally and 
with a subterminal, median ooeciostome. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 639 

I have observed a number of cases of similar ambiguity among the 
Diastoporidae and Tubuliporidae, but never quite to this extent. Pre- 
sumably all such cases may be interpreted as examples of the repetition 
of ancestral characters and therefore useful in tracing the evolution of 
the group. At the same time they present a problem in identification, 
for if only the simple ovicell is present the species must necessarily be 
assigned to a different genus and even a different family than if the 
expanded ovicell is developed, according to Canu's analysis. 

? Plagioecia lactea (Calvet), 1903 

Diastopora lactea Calvet, 1903:163; 1907:466. 
Plagioecia lactea, Canu and Bassler, 1930:48. 

The zoarium is flat and discoidal, with a narrow basal lamina, and 
is attached by a peduncle. The zooecial tubules are immersed for most 
of their length, their surfaces rather coarsely cross-striated and punc- 
tured. The peristomes are moderately short, semi-erect and well sepa- 
rated. Occasional apertures are closed with a lamella with a central 
minute tubule. The orifices are round or slightly elliptical and measure 
about 0.08 mm; the peristomes 0.10 to 0.12 mm in diameter, depending 
on the amount of calcification. 

While no ovicells have been noted in the Hancock specimens, the 
descriptions and figures given by Calvet and Canu and Bassler, and 
measurements by the latter are all in agreement. 

Recorded by Calvet from the Gulf of Gascony at 300 meters and 
from Cape Spadel, Morocco, at 717 meters, and by Canu and Bassler 
from the Galapagos Islands, Albatross Sta. D.2813, at 40 fms. 

Hancock Station 143-34, off Wenman Island, Galapagos, 1°23'10"N, 
91°48'45nV, at 100 to 150 fms. 

Genus DIPLOSOLEN Canu, 1918 

Diplopora Jullien, 1903:115 (preoccupied by Gumbel, 1866). 

Interspersed among the autozoids are nannozoids or reduced indi- 
viduals, irregularly distributed, their minute peristomes shorter than 
those of the normal tubules and often inconspicuous. The ovicell is a 
prominent swelling, usually surrounding a number of peristomes; the 
ooeciostome smaller than the peristomes, short, erect and isolated. Geno- 
type, Tubulipora obelia Johnston, 1838. 

Older authors placed the species under Tubulipora, Berenicea and 
Diastopora, but the constant presence of nannozoids, the function of 
which is unknown, appears sufficient for generic standing. 



640 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Diplosolen obelium, (Johnston), 1838 

Plate 73, fig. 1 

Tubulipora obelia Johnston, 1838:269. 
Diastopora obelia, Hincks, 1880:462. 
Berenicea obelia, Okada, 1917:352. 
Diastopora obelia, O'Donoghue, 1923:14. 
Diplosolen obelium, O'Donoghue, 1926:24. 

The zoarium is thin and flat, rounded or irregularly lobate. The 
zooecia are embedded for most of their length, though the semierect 
peristomes project well above the surface ; the aperture is round or 
slightly elliptical, 0.08 to 0.10 mm in diameter. In Alaska specimens 
the aperture is noticeably larger, 0.10 to 0.12 mm in diameter (var. 
arctica Waters, 1904a :1 71) than in southern specimens, but there seem 
to be no other differences of importance. The nannozoids are similar in 
form to the autozoids, but are minute in size ; their peristomes are much 
shorter and are only about 0.03 mm in diameter. 

The ovicell is considerably inflated, varying in size, oval or arcuate, 
transverse, and encloses a number of peristomes of both autozoids and 
nannozoids (as few as 2 and as many as 20 have been counted). The 
ooeciostome is isolated, short, its aperture rounded and intermediate in 
size between those of the autozoids and nannozoids, usually more or 
less central in position. 

It is a well-known North Atlantic species, extending into the Arctic, 
and reported from Japan. On the Pacific coast it has been recorded by 
O'Donoghue from several places in British Columbia and from the San 
Juan Islands in Puget Sound. 

Hancock Stations: 1194-40, 43 fms, and 1294-41, 34 fms, at Santa 
Cruz Island, southern California. Also among the collections are speci- 
mens from Puget Sound; from Alitak Bay, Alaska (U. S. Fisheries 
Alaska Crab Investigation) ; from Nash Harbor, Nunivak Island, 
Alaska; from the Bering Sea, and from Point Barrow, Alaska (Arctic 
Research Laboratory, G. E. MacGinitie, collector). 

The species is common in Alaska waters, less frequent farther south, 
and Santa Cruz Island, southern California (34°N. Lat.) is the most 
southern record. 

Genus DIAPEROEGIA Canu, 1918 

The ovicell continues to develop after the calcification of the tubes 
distal to it and often surrounds a considerable number of peristomes. 
The ooeciopore is usually not terminal and is often proximal or near 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 641 

the middle of the ooecial swelling. There are several types of ooecio- 
stomes, which may eventually result in the separation of the genus as 
suggested by Canu and Bassler (1920:740). (1) In some species the 
ooeciostome is not associated with a tubule but is quite independent among 
them; (2) in others it is a high tube at the side of a peristome and 
more or less connate with it; and (3) in still others it is a transverse 
or arcuate pore at the base of a peristome and without an ooeciostome. 

Pustulopora intricaria Busk, 1875:22, which is the genotype, is an 
erect, branching species with the ooeciostome isolated and situated a 
little proximal to the middle of the long ovicell, which surrounds a large 
number of peristomes. 

In my opinion Canu and Bassler have depended too much on a single 
character, that of the ovicells surrounding peristomes, for this character 
appears not infrequently to a lesser extent among other genera, even 
in species of genera that do not ordinarily show it, such as Tubulipora, 
Plagioecia, Fasciculipora, Frondipora, etc. Even in Plagioecia patina, 
the genotype of that genus, a peristome may occasionally be surrounded. 
It is also true that in Diplosolen and Crisulipora, which usually have a 
number of included peristomes, ovicells occasionally occur which have 
failed to surround any peristomes. 

The erect species of our eastern Pacific members of this genus agree 
in having an elongate ovicell which extends much beyond the isolated 
and more or less centrally placed ooeciostome. Others, such as those 
described from the Galapagos Islands by Canu and Bassler, D. striatula, 
D. subpapyracea and D. meandrina, with transversely broad ovicells and 
terminal, median ooeciostomes, more properly belong under Plagioecia 
notwithstanding the inclusion of some peristomes. 

It would appear to be true of any species that when the ovicell con- 
tinues to grow forward around a distal peristome, the walls may come 
together and coalesce to enclose it. However this may be, there is cer- 
tainly a group, Diaperoecia, with a well-defined facies which shows an 
extended ovicell enclosing numerous peristomes and with a non-terminal 
ooeciostome. 

The species of the present list show two distinctly different types of 
ooeciostome; in D. intermedia, johnstoni and clavifonnis the ooeciostome 
is a narrow tube at the side of a peristome and is proximal in position, 
while in californica and ftoridana the tube is wider, broadly flared at 
the tip, entirely free from the peristomes and situated more medially. 



642 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Key to the Species of Diaperoecia 

1. Slender, erect or semierect, branching species, rising from a 

small encrusting base, ooeciostome free 2 

Adnate species, with expanded lobes, the lobes sometimes short- 
erect, ooeciostome at the side of a peristome 3 

2. Branches very narrow, seldom as much as 1.0 mm in width, 

apertures of peristomes about 0.13 mm in width . . floridana 
Branches wider, 2 mm or more, apertures of peristomes 0.20 
mm or more, the lateral peristomes usually in short connate 
series californica 

3. Zoarium adnate and branched laterally, with short, erect 

fertile branches which expand into small capitula con- 
taining the ovicell intermedia 

Fertile lobes adnate 4 

4. Fertile lobes flabellate, usually more or less triangular, 

peristomes projecting high above the ovicell .... johnstoni 
Fertile lobe rounded, peristomes projecting only slightly 

above the ovicell claviformis 

Diaperoecia californica (d'Orbigny), 1852 
Plate 67, figs. 1 and 2 

Idmonea Californica d'Orbigny, 1853:732. 
Idmonea Californica, Conrad, 1855:441. 
Idmonea californica, Gabb and Horn, 1862:168. 
Tubulipora dawsoni, Hinclcs, 1884:205. 
Idmonea californica, Robertson, 1910:253. 
Idmonea californica, Canu and Bassler, 1923:199. 
Idmonea californica, O'Donoghue, 1923:12; 1926:27. 
Idmonea palmata, O'Donoghue, 1923:12. 
Diaperoecia intricata, Canu and Bassler, 1928:41. 

The zoarium is composed of erect or spreading branches which fre- 
quently reach a height of 25 mm and occasionally as much as 50 mm. 
The branches may anastomose and often form reticulated masses. 
Usually, in deeper water, the branches are narrow in proportion to their 
length, 2 to 3 mm in breadth, but in exposed places along shore the 
zoarium is more consolidated and the branches shorter and wider and 
less erect {Idmonea palmata O'Donoghue) ; sometimes procumbent and 
attached to the substratum by the radicles {Diaperoecia intricata, Canu 
and Bassler, 1927:41). Radicles or supporting processes are frequently 



NO. 3 OSBURN: EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 643 

present on the dorsal sides of the branches and these may fuse with the 
substratum or with another branch. The dorsal side is more or less 
striated transversely. 

The zooecia are large, their outlines distinct on the frontal surface; 
the peristomes curved into an erect position, arranged in fascicles of 
usually 4 or 5 zoids on either side of the midline, sometimes connate to 
their tips but often only at their bases. Or they may be entirely free 
from each other, and there are often isolated peristomes in the midline. 
The apertures are large, round, averaging about 0.22 mm in diameter. 

The ovicell is a large inflation, usually spread across the whole width 
of the branch below a bifurcation and frequently continuing on one or 
both branches; it surrounds often a large number of isolated peristomes. 
The ooeciostome is isolated, sub-terminal, moderately short, usually bent 
distally but it may be tipped in any direction, large, its base wider than 
a peristome, its tip flared and compressed, as much as 0.60 mm in 
diameter in the long direction, the pore long elliptical. There is much 
variation in the form of the pore, which is sometimes round, and the tip 
of the ooeciostome may be trumpet-shaped without compression. 

This is an extremely common species all along the California coast 
and I have examined hundreds of specimens, ranging all the way from 
the short, palmate form to tall, slender branches from sheltered localities 
and deeper water. It is common in various Pleistocene formations, where 
it was noted by Conrad, Gabb and Horn, and by Canu and Bassler. I 
can find no difference between the Pleistocene and recent specimens and 
I have even found the ooeciostome, which was overlooked by the paleon- 
tologists, except by Canu and Bassler, 1923:199, who show it in Plate 
43, fig. 6. 

The Diaperoecia intricata of Canu and Bassler from the Hawaiian 
Islands is undoubtedly calif ornica, as the differential characters by which 
they distinguish it are exactly those of californica, "par ses colonies reti- 
culees, par son grand ooeciostome et par son ovicelle perforce par des 
tubes ecartes les uns des autres." 

Hincks' description of Tubulipora dawsoni from British Columbia 
fits the zoarial characters of californica perfectly. He made no mention 
of the ovicell. 

Hancock Stations: dredged at more than 100 stations and found at 
shore stations in abundance. The northern limit of its range, as far as 
known, is British Columbia, including the records of Hincks (Tubuli- 
pora dawsoni) and O'Donoghue (both Idmonea californica and /. pal- 
mata.) It is common in the Gulf of California and along the west coast 



644 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

of Mexico; the most southern record is Hancock Station 460-35, at 
Playa Blanca, Costa Rica. The bathymetric range is from low tide to 
about 100 fms. 

Diaperoecia floridana Osburn, 1940 
Plate 67, fig. 3 

Idmonea Milneana, Smitt, 1872:8 {non d'Orbigny). 

Diaperoecia radicata, Canu and Bassler, 1928:160 (no;z Kirkpatrick) . 

Diaperoecia floridana Osburn, 1940:331; 1947:5. 

"? Diaperoecia rugosa Osburn, 1940:332. 

The zoarium is erect or sprawling, idmoneiform, irregularly branched, 
the branches slender, 0.60 to 1.0 mm in width, sometimes anastomosing; 
both dorsal and ventral sides more or less wrinkled ; strong unjointed 
radicles developed on the dorsal side. The tubules are elongate ; in 
younger branches the outlines are definite but the lines disappear with 
age; 4 or 5 to 6 or 7 tubules make up the width of a branch; the 
peristomes are curved, sometimes more than 1.0 mm long but usually 
about 0.40 mm, varying in diameter from 0.16 to 0.20 mm, the aperture 
varying from 0.13 to 0.17 mm; in older specimens transversely wrinkled 
nearly to the tips, perforated at the base. 

The ovicell is elongate, usually located near the end of a branch and 
may extend up both branches at a bifurcation, usually surrounding one 
or more peristomes ; but smaller ones may fail to enclose any. The 
ooeciostome is independent of the peristomes, usually situated near the 
middle of the ovicell; but when this is branched it is located near the 
base of the fork. It has the same width as the peristomes, usually bent 
sharply toward the base but in the forked ovicells it is more or less 
erect. The tip of the ooeciostome in any case, when fully developed, is 
broadly flared, irregularly elliptical, and measures from 0.20 to 0.35 
mm wide by about 0.16 mm in the shorter dimension. 

Pacific specimens have been compared with those from the Atlantic 
and seem to show no essential differences. Also I am inclined to place 
D. rugosa in synonymy, since in our abundant material there is much 
variation in the size of the peristomes, the amount of striation, and the 
form and position of the ooeciostome. 

Described from off Beaufort, North Carolina, and recorded also by 
Osburn from the southern shore of Porto Rico and from several localities 
on the southern shore of the Caribbean Sea; by Smitt (Idmonea milne- 
ana) from Florida, and by Canu and Bassler (D. radicata) from the 
Gulf of Mexico and the Straits of Florida. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 645 

Hancock Stations: 275, Raza Island, Gulf of California, 40 fms; 
305-34, Clarion Island, west of Mexico, 15 fms; 1978-50, Ranger Bank, 
Lower California, 71 fms; and 1143-40, off Portuguese Point, near San 
Pedro, 34 fms, 1413-41, San Miguel Island, 34 fms, 1064, Santa Bar- 
bara Island, 38 fms, and 1240, off San Diego, all from southern Cali- 
fornia. 



The O'Donoghues have recorded under this genus a number of other 
species from British Columbia which have not appeared in our material. 
D. (Entalophora) capitata (Robertson, 1900), 1926:22. 
D. (Entalophora) clavatn (Busk, 1859), 1926:23. 
D. (Stomatopora) expansa (d'Orbigny, 1851), 1926:23. 
D. (Stomatopora) depressa (O'Donoghue, 1923), 1926:23. 
D. (Tubulipora) labiata (O'Donoghue, 1923), 1926:23. 
D. (Tubulipora) striata (O'Donoghue, 1923), 1926:24. 
D. (Entalophora) vancouverensis (O'Donoghue, 1923), 1926:23. 

Diaperoecia johnstoni (Heller), 1867 
Plate 67, fig. 4 

Criserpia Johnstoni Heller, 1867:126. 
Stomatopora Johnstoni, Hincks, 1880:430. 
Stomatopora johnstoni, O'Donoghue, 1923:11. 
Diaperoecia johnstoni, O'Donoghue, 1926:23. 

Our specimens agree very closely with the Stomatopora johnstoni of 
Hincks from the British Isles, though Hincks did not have the ooecio- 
stome. 

The zoarium is encrusting, branching usually dichotomously ; the 
branches short, narrow at the base with 1 or 2 rows of tubules for a 
short distance, beyond which they suddenly become fan-shaped or tri- 
angular with 4 to 6 or 8 tubules in cross-section. The tubules are about 
0.26 mm in diameter, convex and conspicuously perforated; the peri- 
stomes are moderately high, more or less erect, not connate and not 
seriate, 0.18 to 0.22 mm in diameter, the aperture about 0.17 mm. 

The ovicells are as Hincks described them, "dilated and very ventri- 
cose, wedge-shaped," though there is considerable variation in the form ; 
thickly perforated with conspicuous pores; surrounding from 1 to 5 
peristomes. The ooeciostome, located near the middle of the ovicell, is 
as high as the peristomes and about half as large in diameter, the orifice 
0.09 mm, situated at the side of a peristome and connate with it for a 
short distance at the base. 



646 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Heller described the species from the Adriatic Sea, Hincks redescribed 
it from Great Britain, and O'Donoghue listed it from several localities 
in British Columbia and Puget Sound. It is possible that the species 
should be placed in the genus Tubulipora, but our specimens are incom- 
plete in certain respects which prevent a final judgment. It has much 
resemblance to D. intermedia O'Donoghue but the measurements are 
larger and the fertile branches are adnate. 

Point Barrow, Alaska, 21 fms, Arctic Research Laboratory, G. E. 
MacGinitie, collector. Also two specimens from Nash Harbor, Nunivak 
Island, Bering Sea, 8-10 fms, on a shell. 

Diaperoecia intermedia (O'Donoghue), 1923 
Plate 70, fig. 5 

Tubulipora intermedia O'Donoghue, 1923:10. 
Diaperoecia intermedia, O'Donoghue, 1926:23. 

The zoarium is encrusting and branching, with short erect or semierect 
branches which form small capitula. The zooecial tubes are all on the 
ventral side. The stalks of the free branches are about 0.60 to 0.70 mm 
wide and the capitula may reach a maximum width of 3 mm. The peri- 
stomes are all free and moderately long to a maximum of 0.90 mm, 
width 0.16 mm, the aperture 0.13 mm. 

The ovicell has its origin on the ventral side and expands upon the 
top of the capitulum where it surrounds several peristomes ; it is con- 
siderably inflated and thickly perforated. The ooeciostome is more or 
less connected with a peristome at its base, nearly as tall as a peristome, 
and noticeably smaller, its aperture 0.10 mm in diameter, varying in its 
position but usually somewhere near the middle of the expansion. 

O'Donoghue very properly questioned the generic position of this 
species, as the adnate portion of the zoarium is similar to that of 
Proboscina and the ovicell bears some resemblance to that of Tubulipora. 
The nature of the ovicell, enclosing a number of tubules, and especially 
the position of the ooeciostome near the middle of the expansion (occa- 
sionally quite proximal to it) suggest Diaperoecia where O'Donoghue 
finally placed it. It may possibly be one of the various northern species 
which have been described without the ovicell but there is at present no 
proof of synonymy. 

The species was described from Departure Bay, British Columbia. 

Our specimens are from Point Barrow, Alaska, 125 to 522 feet, G. E. 
MacGinitie, collector, common on shells and rocks. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 647 

Diaperoecia claviformis new species ^ 

Plate 66, fig. 11 

Zoarium encrusting on a shell, consisting of a ligulate branch, 0.65 
mm wide, which terminates in an asymmetrical rounded expansion 2 
mm in width. The tubules are short, their outlines inconspicuous, 2 rows 
on the basal portion; the peristomes moderately high and unusually 
close together, not connate and not seriated, 0.16 to 0.18 mm in diameter, 
the apertures 0.13 mm; the younger tubules have the walls perforated 
with small pores but these become closed with age. 

The ovicell is a conspicuous, ventricose area covering most of the 
expanded part of the lobe and enclosing 12 peristomes which are quin- 
cuncial in arrangement; among these the narrow lobes of the ovicell 
are evenly distributed around the peristomes. The ooeciostome is near 
the proximal end, a cylindrical erect tube, connate with a peristome at 
its base only, the orifice round and 0.10 mm in diameter. 

The species has some resemblance to D. johnstoni, especially in the 
narrow ligulate branch and suddenly expanded lobe, but the measure- 
ments are smaller, the peristomes much more closely associated, and the 
meandering branches of the ovicell very narrow. 

Type, AHF no. 92. 

Type locality, Hancock Station 1624-48, off Santa Catalina Island, 
southern California, 33°23'48"N, 118°21'05"W, at 36 fms, one colony 
on a shell. 

Family TubuHpoHdae Johnston, 1838 

"Zoarium entirely adherent, or more or less free and erect, multiform, 
often linear, or flabellate, or lobate, sometimes cylindrical. Zooecia 
tubular, disposed in contiguous series, or in single lines. Ooecium an 
inflation of the surface at certain points, or a modified cell." (Hincks, 
1880:424). 

"Cyclostomata in which the zooecia are restricted to one surface of 
the colony and are commonly arranged in connate alternating series. 
Cancelli are absent in the majority of the species. The ovicell is a modi- 
fied zooecium which is usually much dilated in the region where the 
embryos undergo their development." (Harmer, 1915:119). 

This is a large and difficult family and its analysis is complicated by 
the great number of fossil forms, often without ovicells, that have been 
described. As a rule they are adherent to the substratum, more or less 
lobate, with the zooecial tubes arranged in fascicles, and the ooecium 



648 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

lobate among the fascicles. The early development is characteristic as 
they all begin with a few adnate tubules radiating in a flabellate form 
from one side of the pro-ancestrula ; later the zoaria may assume various 
forms and even become erect and branched. The tubules are usually in 
connate series or groups but may be single over a large part of the 
zoarium or its entire surface, and may be biserially arranged or scattered. 
While the ovicells are usually broad and lobate between the fascicles, 
examples may be found in which they are almost as simple as in Crisia 
or Oncousoecia, and even in species with lobate ooecia, simple ovicells 
may appear on the same zoarium with lobate ones. Also the ovicells may 
occasionally surround tubules or fascicles as in Diaperoecia. The ooecio- 
stomes are very important in the determination of the species, but in the 
various genera they may be terminal, subterminal or more centrally 
located. 

Key to the Genera of Tubuliporidae 

1. Zoarium adnate with slender lobes; tubules in connate single 

series, on each side of the midline ; ooecium spreading the 

full width of the lobe between the fascicles, the ooeciostome ,V 

proximal to the first tubule of a fascicle Platonea* 

Ooecium not so arranged 2 

2. Ooecium arcuate and much depressed between the fascicles, the 

ooeciostome terminal at the middle of the arcuate ooecium ; 

zooecial tubes thick- walled Bathysoecia^S^ 

Ooecium not arcuate, the surface inflated .3 

3. Zoarium composed of extremely high, folded fascicles ; ovicell 

very elongate, simple, like a somewhat enlarged 

tubule Fasciculipora 

Zoarium usually flat and adnate, rarely erect and branched; •'J ' 
tubules in clusters, radiating series or single; ooecium 
usually broadly lobed between the tubules or fascicles, some- 
times smaller and simple Tubulipora 

Genus TUBULIPORA Lamarck, 1816 

Zoarium variable, encrusting and lobulate, repent and branching, or 
erect and branching. Zooecia all on the frontal surface, arranged more 
or less in transverse series or in groups, usually single near the ancestrula 
and occasionally over the whole zoarium. The ovicell is an inflated 
gonozoid between the tubules on the frontal surface, simple and pyriform 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 649 

to broadly lobate with the lobes extending between clusters of tubules; 
the ooeciostome is usually located at the side of a zooecial tube, some- 
times free from it or more or less connate, varying in size, height and 
form among the different species. Genotype, T. transversa Lamarck, 
1816 ( = T. liliacea Pallas, 1766). 

Key to Species of TubuUpora 

1. Zoaria comparatively large and coarse, often irregular in form, 

the ovicell usually much ramified, ooeciostome tall .... 2 
Zoaria smaller, often simply lobate, ovicell little ramified, the 

ooeciostome shorter 5 

2. Ooeciostome high, much compressed, the aperture slit-like . . 3 
Ooeciostome high, little compressed, aperture ovate 4 

3. Peristomes connate in series or bundles, ooeciostome tall and 

conspicuous, connate with a tubule at its base, enlarging 
upward and slightly flared, the aperture compressed (in- 
cluding var. fasciculifera) tuba 

Peristomes sometimes in series, ooeciostome smaller and less 
conspicuous, its aperture more narrowly slit-like and not 
flared flabellaris 

4. Peristomes connate, forming high fascicles, the tips free; 

ooeciostome scarcely compressed, tall, about as wide as a 
peristome (0.25 mm), slightly expanded at the tip . admiranda 
Peristomes not at all connate; ooeciostome slightly smaller 
than a peristome, not compressed, its tip slightly expanded, 
not connate with a peristome egregia 

5. Zoarium with erect slender branches, idmoneiform; ovicell 

small, ooeciostome slightly distal to the first member of a 

fascicle, short, flared ftexuosa 

Zoarium adnate, small neat-appearing species 6 

6. The dorsal side of the zoarium has numerous short attachment 

processes and near the base these give the edge a serrated 

appearance pulchra 

No attachment processes ' 

7. Zoarium widely flabellate; ooeciostome comparatively short, 

connate at base and widely diverging, flared at the tip to a 

width of 0.18 mm pacifica 

Zoarium lobate, ooeciostome short erect, connate at base or 

free, flared at the tip to a width of 0.12 mm .... concinna 



650 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Tubulipora tuba (Gabb and Horn), 1862 
Plate 68, fig. 9 

Semitubigera tuba Gabb and Horn, 1862:169. 
Tubulipora occidentalis Robertson, 1910:249. 
Tubulipora occidentalis, O'Donoghue, 1923:8. 
Tubulipora tuba, Canu and Bassler, 1923:198. 
Tubulipora tuba, O'Donoghue, 1926:24. 

The zoarium of this abundant species is always adnate, flat and rather 
regularly rounded or sometimes lobate on flat surfaces, variously con- 
torted on stems; rather coarse, white, gray or purplish in color. The 
"peristomes" are nearly erect, 0.12 mm in diameter, varying greatly in 
length, as much as 2 mm in sheltered locations but usually much less; 
single near the primary zoid and sometimes over a considerable area, 
then connate in small fascicles of 2 or more, the marginal fascicles 
increasing in the number of peristomes to 6, 12 or even as many as 30. 
The fascicles are usually uniserial or biserial and radiating, but occa- 
sionally occur in rounded or irregular clumps ; sometimes they are more 
or less biradial in arrangement, but this is rare. 

The ovicell is usually a large lobate inflation extending between several 
fascicles, but not infrequently it is smaller, and even simple Crisia- 
like ooecia may occur on the same zoarium with the larger normal ones. 
The ooeciostome is tall, straight, compressed, regularly increasing in size 
toward the tip, usually a little flared at the top, the pore elongated in the 
direction of the fascial axis ; in typical tuba the ooeciostome arises at the 
side of the first tube of a fascicle and is usually free for most of its 
length. The variations are discussed under the variety fasciculifera 
(Hincks). 

Gabb and Horn described the species from the Pleistocene of Santa 
Barbara, California, and while their description and figure are incom- 
plete, there can be no doubt, I have compared abundant recent and 
numerous Pleistocene specimens. Canu and Bassler listed both tuba and 
fasciculifera from the Pleistocene of California. Robertson described 
occidentalis (=^ both tuba and fasciculifera) and listed it from southern 
California to Puget Sound, and O'Donoghue recorded both from numer- 
ous localities in British Columbia. 

In the Hancock Collections it is by far the most abundant species of 
the genus, taken at shore stations and dredged down to a depth of 117 
fms. It is evidently a species of cooler waters as the most southerly 
record is that of Station 275, Raza Island, Gulf of California, 28°48'N, 
113°W. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 651 

Tubulipora tuba van fasciculifera (Hincks), 1884 

Plate 68, fig. 10 

Tubulipora fasciculifera Hincks, 1884:206. 
Tubulipora occidentalis Robertson, 1910:249 (in part). 
Tubulipora fasciculifera, Canu and Bassler, 1923:197. 
Tubulipora fasciculifera, O'Donoghue, 1923:8; 1926:24. 

The zoarium is very similar to that of T. tuba, presenting the same 
variations in form. The zooecia are also similar, the free portions of the 
tubules varying much in length and having the same diameter (0.12 
mm). The only zoarial difference is that made use of by Canu and 
Bassler, the fascicles "never composed of more than 6 tubules," while 
in tuba there may be "from 6 to 20." 

The ovicell, like that of tuba, is expanded into lobes w^hich extend 
between the fascicles, and here also there is the occasional occurrence 
of simpler ooecia. The ooeciostomes are usually situated proximal to the 
first tubule of a fascicle, connate with it for a short distance, with the 
flattened ooeciopore transverse to the axis of a fascicle, the top of the 
ooeciostome sometimes a little flared. 

After studying more than a hundred specimens from various locali- 
ties, I am unable to distinguish sharply between tuba and fasciculifera. 
The above diagnoses are for well-marked specimens, but intermediate 
conditions occur in all of the diagnostic characters. Many colonies have 
only the smaller fascicles, others mostly small ones with a few larger 
fascicles, and still others have chiefly the larger numbers. Occasionally 
the long fascicles arise near the center of the zoarium, while in other 
specimens they are nearer the edge. The ooeciostomes of tuba are usually 
lateral to a tubule with the pore in line with the fascicle, but may be 
proximal to a tubule with the pore transverse, and in fasciculifera both 
of these conditions may sometimes be seen on the same zoarium. The 
form of the ooeciostome is variable though it is always more or less 
compressed ; sometimes it is slightly flared at the tip, or it may be per- 
fectly straight (possibly those in the latter condition have not quite 
completed their growth). If the size of the fascicles and the position 
of the ooeciostome were constant they would be considered good specific 
characters, but I do not find them so. 

Hincks described T. fasciculifera from British Columbia, the exact 
locality not stated. Robertson mentions it under her description of T. 
occidentalis, which embodies some of the characters, and lists occiden- 



652 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

talis from Puget Sound to southern California. O'Donoghue recorded 
it along with occidentalis from numerous places in British Columbia and 
Puget Sound. 

The variety, if the varietal distinction is really worthwhile in this 
case, appears to occur throughout the range of tuba, in the same habitat, 
and they are found together in the Pleistocene at a number of places in 
southern California. 

Tubulipora pacifica Robertson, 1910 
Plate 68, fig. 1 

Tubulipora pacifica Robertson, 1910:248. 
Tubulipora pacifica, O'Donoghue, 1923:8; 1926:25. 

The zoarium is encrusting, usually on algae ; small ( rarely more than 
3 mm across), white and rather delicate; fan-shaped to nearly circular, 
or occasionally with lobes of the same form. The immersed zooecial 
tubules are long and slender, transversely arched and thickly punctate. 
The peristomes are moderately high, about 0.12 mm in diameter; near 
the center of the colony they are single but farther out they are usually 
fasciculate, with one or two rows of peristomes which are connate with 
the tips divergent ; there is a tendency for them to be distributed biradi- 
ately, on either side of the zoarial axis. 

The fully developed ovicell appears to be considerably larger than 
in T. pulchra, with as many as 3 or 4 lobes between the fascicles, but 
frequently they are much simpler, pyriform, and resemble those of 
Crisia, only more immersed, and all the intermediate conditions may be 
observed. The ooeciostome is comparatively short and is very briefly 
connate with the succeeding tubule at its base, sharply diverging proxi- 
mally, or as Robertson expressed it, "It seems to emerge from the side 
of a zooecium at right angles to it." It is flared outward at the tip, 
compressed, the ooeciopore elliptical and about 0.18 mm in its greatest 
diameter. 

At first glance the species has much the appearance of pulchra but the 
dorsal side is smooth without attachment processes, there is no serration 
of the margin at the base, the peristomes are larger, stiffer-looking and 
in adult colonies there is always some fusion of the peristomes into small 
fascicles. Robertson described and listed it from various shorewise locali- 
ties in southern California, and O'Donoghue recorded it from numerous 
places in British Columbia. It has a wide distribution along the coast to 
as far south as Colombia ; apparently a shallow-water species, but dredged 
down to 47 fms. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 653 

Hancock Collections: numerous shore stations and shallow-water 
dredgings about the off-shore islands of southern California; Station 
225-34, Gorgona, Colombia, 2°58'55"N, the most southern record. Also 
Albatross collections, 1911 cruise, at San Francisquito Bay and San 
Esteban Bay, Lower California. 

Tubulipora pulchra MacGillivray, 1885 
Plate 68, figs. 2, 3, and 4 

Tubulipora pulchra MacGillivray, 1885:95. 
Tubulipora fimbria forma pulchra. Waters, 1887:258. 
Tubulipora pulchra, Robertson, 1910:250. 
Tubulipora pulchra, O'Donoghue, 1923:8; 1926:25. 

A beautiful small, white, dehcate species which adheres loosely to the 
substratum, usually a kelp, but frequently to shells. The zoaria are 
small, usually only 2 or 3 mm in extent, more or less fan-shaped or ovate, 
sometimes with lobes of the same size and form. A peculiarity of the 
dorsal surface is the presence of numerous short attachment processes 
which support the zoarium "on tiptoe," as Miss Robertson suggests; on 
the marginal zoids these are larger and project laterally to give the 
border a serrated appearance, especially near the ancestrula. The tubules 
are small, elongate and very slender ; the peristomes are long and slender 
(0.08 to 0.09 mm in diameter), not connate and not seriated. 

The ovicell or gonozoid is simple and little expanded, its form fre- 
quently resembling that of a Crisia but more embedded; at its fullest 
development there are 2 or 3 short lobes extending laterally between 
the peristomes. The ooeciostome is erect and moderately high ; at the base 
it is about as wide as a peristome and at the tip it flares out into a com- 
pressed trumpet shape about twice the width of the base; it is never 
connate with another tubule. 

MacGillivray described the species from Australia, Robertson re- 
corded it from the southern California coast, and O'Donoghue found 
it at a number of localities in British Columbia. 

Hancock Stations: numerous stations along the coast of California 
and among the Channel Islands; Station 72, Guadalupe Island, and 
136-34, Clarion Island, west of Mexico; 468-35, Port Parker, Costa 
Rica; 462, James Island, Galapagos. Shore to a depth of 35 fms. 

Tubulipora flexuosa (Pourtales), 1867 

Plate 71, fig. 11 

I dmonea flexuosa Pourtales, 1867:111. 
Idmonea atlantica var. flexuosa, Smitt, 1872:6. 



654 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

1 Idmonea atlantica van tenuis, Hincks, 1880:452. 
Tubulipora atlantica var. flexuosa, Harmer, 1915:127. 
Idmonea atlantica var. flexuosa, Osburn, 1940:333. 
Idmidronea atlantica var. flexuosa Osburn, 1947:5. 

Pourtales gave only a brief description, in which the most important 
points are the slender, flexuous and round branches. Smitt re-worked 
Pourtales' material and gave a good description and figures. Unfor- 
tunately, to the present time, no one has observed the complete ovicell 
with ooeciostome. It is on the basis of the latter character, chiefly, that 
1 am elevating it once more to full specific standing. 

The zoarium presents the same general characters as the well-known 
atlantica, erect and branching from a small base, but the branches are 
very slender, much flexed pnd sinuous, and in cross-section they are 
round instead of being flattened on the dorsal surface. The fascicles are 
short, the tubules 2 or 3 in series (rarely 1 or 4), while in atlantica they 
are 3 or 4 to as many as 6, and they average a trifle smaller in diameter, 
connate to the tips and slightly narrowed upward from the base. 

The ooecium and the ooeciostome (which is here described for the first 
time) are quite different from those of atlantica. The ooecium is short, 
usually occupying only two interfascicular areas, into which it spreads 
more or less, while that of atlantica is usually very elongate and is 
limited to the axis of the branches and not lobed laterally; the perfora- 
tions of the ovicell wall also appear to be more minute and more numer- 
ous. The ooeciostome presents the most striking difference, as it is very 
short, erect, with a widely flared and rather thick border; it is located 
just medial to the first tubule of a fascicle and slightly separated from it. 
In atlantica the ooeciostome is about as tall as the tubules, curved distally, 
expanded gradually, situated on the distal side at about the second tubule 
of the fascicle, and its base connate with a tubule for a short distance ; 
in the several ooeciostomes I have observed there is no intergradation. 

This form was described by Pourtales and by Smitt from north of 
Cuba and later recovered by Osburn from Porto Rico and the southern 
shore of the Caribbean Sea. Harmer's reference (1915:127) from the 
Netherlands East Indies appears undoubtedly to be the same, for, while 
he did not have a complete ooeciostome, his fig. 1, plate 10, shows the 
base and pore in the characteristic position. Any attempt at a complete 
synonjany would be useless, and it will even be uncertain whether Busk's 
variety tenuis is the same as flexuosa until the ovicells are carefully 
studied. Also, there is no certain record of Tubulipora (Idmonea) at- 
lantica from the eastern Pacific. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 655 

Hancock Stations: Raza Island, Gulf of California, 28°48'N, 113° 
W, at 40 fms, numerous colonies in reproduction ; also one colony from 
James Island, Galapagos, at 54 fms. 

Tubulipora concinna MacGillivray, 1885 
Plate 67, fig. 5 

Tubulipora concinna MacGillivray, 1885:94. 
Tubulipora concinna, Harmer, 1915:123. 

The zoarium is entirely encrusting on erect stems and on flat surfaces, 
the branches narrowly lobate and curved laterally, a small and delicate 
species. The slender peristomes are very elongate, 0.40 to 0.75 mm, 
strongly curved and often sinuate, sparsely punctate, about 0.09 mm in 
diameter and the aperture about 0.07 mm. On flat surfaces the peri- 
stomes are usually directed somewhat outward from the midline of the 
lobes, but on small stems they are very irregular in arrangement; for 
the most part they are distinct, but on the broader portion of the lobe 
they are frequently in series of 2 to 4 and connate to the tips. 

The ovicells are small, almost as simple as in Crisia, narrow proximally 
and gradually expanded and sometimes slightly lobed between the peri- 
stomes, the frontal surface inflated and thickly punctate with very small 
pores. The ooeciostome is nearly terminal, free or in contact with a 
peristome, short, the aperture expanded and ovate in form, transverse 
and about 0.12 mm wide by 0.07 mm long. 

Hitherto recorded only from Australia and the East Indies. Our 
specimens appear to agree in every detail with the description and with 
Harmer's beautiful illustration (plate 10, fig. 10), except that the ovi- 
cells are even simpler and less lobate ; the ooeciostome is an exact counter- 
part. 

Hancock Stations: 1924-49, of? Guadalupe Island, west of Lower 
California, 28°54'08"N, 118°15'36''W, 25-30 fms, on algae, several 
colonies. Also on a sunken buoy brought up from 45 fms at Rocky Point 
(Earl Fox, collector), several colonies; one colony from "ofif San Pedro," 
without other data; and 12 colonies on a kelp stem washed up on shore 
at Palos Verdes (R. C. Osburn, collector), all from southern California. 

Tubulipora egregia new species 
Plate 67, figs. 6 and 7 

The zoaria are encrusting, surrounding the stems of a coralline alga, 
in one case spreading across free from one branch to another; usually 
rough and irregular but one specimen has two flabellate lobes. The most 



656 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

unusual feature is the size of the peristomes, which reach a length of 
as much as 0.75 mm (usually 0.40 to 0.60), and a diameter of 0.26 mm 
(0.30 at the base). The peristomes are entirely free at the tips and only 
rarely connate at the base, nearly erect, perforated with small pores 
nearly to the tips and the basal half or more transversely corrugated. 
The zooecial tubes are correspondingly large, 0.30 to 0.40 mm wide, 
arched in cross-section, thickly perforated and transversely corrugated. 

The ovicell is irregularly lobate, rather flat and its surface thickly 
punctured, enclosing a few peristomes ; the ooeciostome is an erect tube, 
distant from and smaller than the peristomes, slightly enlarging upward, 
the aperture ovate in form and its longest dimension about equal to that 
of the peristomes, finely wrinkled, thin-walled and not punctate. 

The large dimensions of the non-connate and non-seriate peristomes, 
and the nature of the ovicell and ooeciostome easily distinguish this 
species. 

Type, AHFno. 115. 

Type locality, Hancock Station 22-33, La Plata Island, Ecuador, 
1°16'S, SPOS'lO^'W, shore collecting, four colonies all with ovicells, 
Jan. 22, 1933. Another colony, with ovicell, from Hancock Station 
136-34, Clarion Island, west of Mexico, at 32 fms. 

Tubulipora admiranda new species 
Plate 68, figs. 5, 6, and 7 

The zoarium is rounded, slightly irregularly lobate, 10 mm broad, 
attached over most of its dorsal side but with the edges free. The center 
of the zoarium over a width of 4 mm bears only 5 free peristomes, due 
to the great length of the embedded tubules ; outside of this area the 
peristomes rise in clusters of varying size, giving the surface a lobate 
appearance, though there are many free peristomes between the clusters. 
In the clusters the peristomes are connate for most of their length, but 
usually free at the tips. 

The pro-ancestrula is round and measures 0.40 mm in width ; the first 
tubule, which arises from its side, is 0.26 mm wide and 0.78 mm long, 
its peristome 0.55 mm in height. The succeeding tubules are remarkable 
for their length, the embedded portion 1.0 to 2 mm in length and the 
more or less erect peristomes usually about 1.0 mm but may be as much 
as 2 mm. The embedded tubules average 0.40 mm in width, slightly 
arched in cross-section, and are thickly punctate. The peristomes are 
about 0.25 mm in diameter, the pores extending nearly to the tips, 
slightly wrinkled, the apertures round and 0.20 mm in diameter. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 657 

The ovicells, three of which are complete, vary In size, with lobes 
extending between the fascicles, their surface rather flat and very thickly 
punctate. The ooeciostome is a large erect tube, about as wide as a 
peristome at the base, connate, tall, punctate to its tip, somewhat com- 
pressed and broader at the tip, its aperture about 0.30 mm wide by 0.18 
mm long, and the edges of the long sides slightly inflexed ; arising proxi- 
mal to the base of a peristome; in one case the ooeciostome curves 
around the side of the adjacent peristome to open on its distal side. 

The nearly free central area, the great length of the large tubules 
and the semi-erect clustered connate peristomes give this species an 
unusual appearance. The ooeciostome with its broad base resembles 
somewhat that of T. phalangea, but it is much larger and is not hooded 
as in that species. 

Type, AHFno. 114. 

Type locality. Corona del Mar, southern California, 33°36'N, one 
colony on the broad hold-fast of a kelp, washed up on the beach, R. C. 
Osburn collector. 

Tubulipora flabellaris (Fabricius), 1780 
Plate 68, fig. 8 

Tubipora flabellaris Fabricius, 1780:430. 
Tubulipora flabellaris, Harmer, 1899:99 (synonymy). 
Tubulipora flabellaris, Robertson, 1910:247. 
Tubulipora flabellaris, O'Donoghue, 1923:8; 1926:24. 

The zoarium ranges in form from flabellate in younger colonies to 
round in older ones, completely adnate and attached to shells, stems, 
worm tubes, algae, etc., on flat surfaces usually very symmetrical and 
reaching as much as 8 mm in diameter. The central part of the colony 
is rather small, the first few peristomes free and often sinuate; beyond 
this area the peristomes are arranged in hnear series (occasionally in 
small groups) of 2 or 3 or longer, irregularly radiating. The peristomes 
are high, slender, the apertures about 0.12 mm, connate for most of 
their length but the tips usually free. 

The ovicell is more or less lobate, usually spreading between 3 or 4 
fascicles, its surface rather coarsely punctate, and there is rarely any 
evidence of striation; the ooeciostome is a tall slender tube at the side 
of and partially connate with a peristome, much compressed toward the 
top and the aperture slit-like, measuring about 0.13 mm long by 0.04 
or 0.05 mm wide. 



658 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

It is a common species in the northern Atlantic, on the European 
coast, the American coast as far south as Cape Cod, and in the Arctic 
area from Spitsbergen west to Icy Cape, Alaska. Robertson recorded it 
from Puget Sound to southern California, and O'Donoghue from several 
localities in British Columbia. 

Hancock Collections: dredged only once. Station 1122-40, of? San 
Nicolas Island, southern California, 33°18'N, 119°24'l(y'W, at 30 
fms. Also from Point Barrow, Alaska, Arctic Research Laboratory, 
G. E. MacGinitie, collector. 

Genus BATHYSOEGIA new genus. 

Ovicell depressed between the erect tips of the zooecial tubules, 
irregularly arcuate in form vvith the ends of the arc prolonged distally 
into narrow lobes between the fascicles; ooeciostome at the distal border 
of the arc, median, small, erect and connate to a tubule only at its base. 
Zoarium rounded or lobate; the ancestrula and first few single tubules 
tubulipora-like ; then the tubules become more or less erect in small 
groups, connate to their tips. Peristomes are often wanting except in 
older stages, when they arise around the aperture of the partially closed 
end of the tubules. Genotype, Bathysoecia bassleri Osburn, new species. 

In the genotype the erect tubules are so closely connate that there is 
no exposure of the tubules except at their tips, where they produce a 
reticulum. In younger zooecia, near the zoarial margin the tubules are 
thin-walled and wide open ; later the walls become thick at the tips and 
form an infundibular depression with a rounded aperture; still later 
around the aperture there rises a thin-walled peristome which projects 
upward from the bottom of the funnel. 

The ovicell appears to be different from that of any other form among 
the Tubuliporidae. It is developed directly on the basal lamina before 
the tubules distal to it are formed. Later the connate tubules rise high 
around it on all sides of the ovicell, which appears as a depressed and 
irregularly arcuate area with a flat thin-walled surface. The ooecio- 
stome is distal, median and connate with a tubule or between two of 
them, narrow and moderately high. 

The only other species with similar tubules and ooecia that has come 
to my attention is the "? Tubulipora (Tubularia by error) lohulata" 
Osburn, 1933:16, from the Atlantic Coast of North America, which 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 659 

appears to be congeneric ; described below as Bathysoecia hastingsae new- 
species. The status of Tubulipora lobulata Hassall and T. lobulata 
Hincks is also discussed under that species, 

Bathysoecia bassleri new species 
Plate 69, figs. 4, 5, and 6 

Zoarium encrusting on shells, irregularly rounded or with short lobes ; 
a narrow basal lamina; surface reticulated. The zooecial tubules are 
completely connate to their tips, nearly vertical, so closely set that their 
exposed ends occupy all of the frontal surface; more or less hexagonal, 
separated by strong ridges and their apertures widely funnel-shaped over 
most of the zoarium. The tubules arise from the basal lamina, at first 
prone but immeoiately curving upward to become more or less erect. 
They are completely connate from their origin and are so closely united 
that no line of separation is visible. As they approach maturity the distal 
exposed ends become partly closed by a funnel-shaped thickening which 
leaves a large rounded aperture at the bottom of the funnel. Most of 
the tubules remain in this condition, but in older areas some of them 
develop short, cylindrical peristomes inside of the funnel and may pro- 
ject slightly above it. The exposed ends of the zooecia, from ridge to 
ridge, measure 0.20 to 0.35 mm across, the apertures 0.18 to 0.20 mm, 
and the cylindrical peristomes of older zooecia 0.13 to 0.15 mm in 
diameter. 

The ovicell is irregularly arcuate in form and so deeply submerged 
between the high walls formed by the connate tubules that it has none 
of the usual appearance of an ovicell. The frontal layer is thin in com- 
parison with the wall of the tubules, and is perforated by minute pores 
(the only pores visible on the whole zoarium). The ooeciostome is located 
at the distal border in the middle of the arc, compressed and nearly as 
high as the tubules, connate with a tubule or often between two tubules ; 
the ooeciostome slit-like, its long diameter paralled to the zoarial radius, 
the pore 0.15 to 0.20 mm long by about 0.06 mm wide. 

The species is dedicated to Dr. Ray S. Bassler, whose extensive studies 
of the Bryozoa have been of great service to the author. 

Type, U. S. Nat. Mus. no. 11050; paratype, AHF no. 116. 

Type Locality, Lenard Harbor, Alaska, a branch of Cold Bay, Alaska 
Crab Investigation Sta. 60-40, 55°10'N, 163°30'W, at 25 fms, 4 colonies 
on shells. Also from Hein Bank, near Friday Harbor, Puget Sound, 
Washington, one colony, Dr. J. L. Mohr, collector. 



660 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Bathysoecia hastingsae new species 

Plate 69, fig. 7 

Tubulipora (Tubularia by error) lobulata, Osburn, 1933:16. 
? Tubulipora lobulata, Whiteaves, 1874:215; 1901:111. 

The zoarium is irregularly fan-shaped or lobed, completely adnate on 
stones and shells ; thick, especially so near the middle, and sloping down- 
ward to a narrow basal lamina. The zooecial walls are heavily calcified, 
the tubules thick and the only exposed areas (near the ancestrula) trans- 
versely ribbed. There is no evidence of pores except slightly in the pri- 
mary zooecial area. The peristomes are moderately high, erect, single, 
especially near the primary area; farther out they may be single, or 
connate in short lines or small groups. The erect portion appears to 
consist partly of the upturned distal end of the tubule, as in B. bassleri, 
new species, but the condition is not so striking; the remaining portion 
is the peristome, which is considerably smaller than the base on which 
it arises, diameter 0.16 mm; the peristome is present on most of the 
tubules (wherein it differs from B. bassleri, in which most of the tubules 
bear no peristomes). 

The ovicell is similar to that of the genotype but its surface is less 
depressed, a flat, white, finely perforated layer; the chamber extends 
downward to the basal lamina, as it does in B. bassleri. The form of the 
ovicell is like that of bassleri, usually beginning with an arcuate portion 
and extending into narrow lobes which ramify more or less between the 
fascicles; in one case a fascicle has been completely surrounded. The 
ooeciostome differs sharply from that of B. bassleri, as it is a short erect 
cylindrical tube, not at all compressed, connate only at its base, and its 
tip circular and noticeably flared, 0.12 mm across and the pore 0.07 mm 
in diameter. 

Twenty years ago Dr. Anna B. Hastings, after examining a specimen 
from Mount Desert, Maine, wrote me that "it is likely that it is T. 
lobulata Hassall," (Osburn, 1933:16). Now, with very mature judg- 
ment, Dr. Hastings has re-examined the whole problem and writes again 
(March 8, 1952) in part as follows: 

"This time I say with confidence that three species are involved. 

1, T. lobulata Hincks (not Hassall). Excellently described by 
Hincks. I need only to add that the ooeciostome is of similar diameter 
to the zooecial tubes, but shorter, and is attached to the side of one of 
them. It is widely open, directed upwards or a little obliquely with a 
slight out-turned rim. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 661 

2. T. lobulata Hassall. Differs from T. lobulata HIncks in having 
long oblique series of connate zooecia. (Hassall, 1841, pi, X, fig. 2). 
I think Norman may well have been right (MS. note) in referring it 
to T. serpens (T. liliacea Pallas). 

3. T. lobulata ? Osburn. Resembles T. lobulata Hincks in its stout 
zooecia with thick transversely striated walls, and in the arrangement 
of the zooecia in the colony, separately or in small groups, but not in 
connate series ; and in the depressed ooecia. Differs in its less ramified 
ovicell with the ooeciopore placed more or less symmetrically at the 
center of the distal border of the ooecium closer to, and behind rather 
than beside, a zooecial tube ; in the small size of the ooeciostome and in 
the absence of an out-turned rim to the ooeciostome." (The last item 
must now be corrected as I have a complete ooeciostome with a slightly 
out-turned rim. R.C.O.) 

It is a pleasure to be able at last to solve the long-standing problem 
of the position of the West Atlantic specimens of "T. lobulata" which 
could not have been done without the careful analysis by Dr. Hastings, 
to whom I gratefully dedicate the species. 

The species is now known to be distributed on the Atlantic coast from 
Mount Desert Island, Maine (Osburn, 1933:16); Gaspe (Canada), 
Hincks Collection (Hastings, in litt., British Museum), and Green- 
land, "Valorous," 1875, Norman Collection, (Hastings, in litt., British 
Museum). Now I am able to add the Bering Sea, and the species is 
certainly high northern and possibly circumpolar in its distribution. 

Type, AHFno. 117. 

Type Locality, Nunivak Island, Bering Sea (a large island off the 
west coast of Alaska, about 60°N, and 116° W) at 8 to 10 fms, on 
shell, 4 colonies. Another specimen is marked merely "Behring Sea," 
on shell. 

Genus PLATONEA Canu and Bassler, 1920 

Platonea Cznn and Bassler, 1920:759; 1929:548. 
Reptotubigera A'Orhigay, 1853:751 (in part). 
Reptotubigera, Calvet, 1911:4. 
Reptotubigera, Harmer, 1915:119. 
Reptotubigera, Okada, 1928:492. 
Reptotubigera, Borg, 1944:26. 

This genus has been accepted by the above authors to include narrow, 
fasciculate species that are entirely adnate to the substratum, as described 
by d'Orbigny. But d'Orbigny made no reference to the ovicell, the first 



662 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

species which he discussed, R. neocomiensis, does not appear to belong 
to the genus, and his fourth species, R. ramosa d'Orbigny, 1853:754, 
has been selected as the genotype of Reptotubigera. Since there must 
always be uncertainty when based only on zoarial characters, Canu and 
Bassler erected the genus Platonea with the description of the ovicell 
and ooeciostome and with Reptotubigera philippsae Harmer as the geno- 
type. The ovicell is broadly lobed between the fascicles and extends to 
the borders of the zoarium on both sides and the ooeciostome is short, 
erect, more or less expanded and is located proximal to the first peristome 
of a fascicle and separated from it. 

If we are to accept this type of ovicell as distinctive of the genus, the 
zoarial description must be modified to include several species which are 
only partly adnate or semierect, but which have the special characters 
of a broad ovicell extending between the fascicles and an ooeciostome 
which is short, isolated and located near a first peristome of a fascicle. 
This last character, however, is shared by some species of Tubulipora. 
Also the marginal basal lamina, which is characteristic of the adnate 
species, is wanting in the suberect species. 

As with many of the Tubuliporidae it is difficult to draw sharp dis- 
tinctions in all cases but, dismissing the other characters, the idmoneiform 
arrangement of the fascicles and the nature of the ovicell and ooeciostome, 
together present a very characteristic facies sufficient to give this group 
a distinct place among the genera of the Tubuliporidae. 

While it is possible that some of the fossil species included by 
d'Orbigny under Reptotubigera will be found to agree with Platonea, 
there is no indication in his description or figures of an ovicell and all 
of his figures represent the peristomes as short and non-connate. I am 
therefore following Canu and Bassler in the use of Platonea for species 
in which the reproductive characters are present. 

Platonea veleronis new species 
Plate 69, fig. 2 

Zoaria adnate, apparently attached to algae; usually consist of a 
single lobe but occasionally a single branch occurs; maximum length 
5 mm, width 1.5 to 2 mm; the dorsal side not expanded beyond the 
lateral peristomes. The peristomes are arranged in series of 4 to 7, 
closely connate, except for one or two at the outer ends of the series; 
the aperture more or less quadrangular, measuring about 0.09 mm wide 
by 0.12 mm long. At the proximal end the peristomes are single or in 
short series. The fascicles are high, 0.50 to 0.70 mm, very regular, 
usually alternating on the opposite sides of the midline, the interspaces 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 663 

The ovicell usually occupies 3 (sometimes 2, or 2 on one side with 3 
on the other) of the interfascicular areas on each side toward the distal 
end and extends laterally the full width of the lobe, smooth or slightly 
wrinkled and perforated with very numerous minute pores. The ooecio- 
stome is short, erect, about as wide as a peristome, the aperture suddenly 
expanded transversely and 0.09 mm long by 0.18 mm wide; located 
slightly proximal and medial to the first peristome of a fascicle and not 
connate with it. The ovicell is usually situated near the tip of the lobe, 
but in two cases there is an additional one situated more proximally, 
smaller and separated from the distal one by 2 or 3 fascicles. 

Type, AHFno. 118. 

Type locality, Hancock Station 450, Albemarle Island, Galapagos, 
0°55'S, 90°30'W, at 60 fms. Also dredged at stations 190-34, Albemarle 
Island ; 201-34 and 473, Hood Island ; 41 1, Duncan Island ; 452, Charles 
Island ; 453, Gardner Island, and 484, Barrington Island, all from the 
Galapagos, at 25 to 75 fms. It is evidently well distributed among the 
Galapagos Islands but has not been noted elsewhere. 

Platonea expansa new species 
Plate 69, fig. 3 

The zoarium consists of somewhat clavate lobes about 2 mm in width 
near the extremity, the tip rounded, loosely attached, apparently always 
on algae. The fascicles are unusually long, very regular and alternating 
in arrangement on each side of the midline of the lobe ; the fascicles near 
the proximal end with 4 to 6 tubules, the more distal ones with about 
7, closely connate to their tips, except for 1 or 2 at the outer ends which 
are either connate only at the bases or are entirely free. There is a 
moderately broad basal lamina. The apertures of the tubules are quad- 
rangular in the connate series, round in the free ones, and measure about 
0.14 mm in diameter. Distance between the fascicles averages about 
0.24 mm. 

The ovicell is very broad, extending to the outer ends of the fascicles, 
and occupying three interfascicular areas on one side and two on the 
other, smooth with numerous very small pores. The ooeciostome, proxi- 
mal to the first peristome of a fascicle, is short, not connate, its diameter 
at the base noticeably wider than a peristome, directed somewhat proxi- 
mally, flared widely at the tip, the opening transversely elliptical. 

The length and regularity of the high fascicles is the most striking 
feature of the species. 

Type, AHF no. 120. 



664 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Type locality, Hancock Station 190-34, Albemarle Island, Galapagos, 
0°55'S, 90°30'W, at 58 fms. Also at Station 201-34, Hood Island; and 
453, Gardner Island, Galapagos, 35 to 65 fms. 

Platonea elongata new species 
Plate 69, fig. 1 

Zoarium adnate, slender, unbranched ; the dorsal lamina narrow, 0.95 
mm at its widest part, while the high fascicles project much beyond it 
to a width of 1.50 mm. The fascicles are regular and alternating on each 
side of a distinct midline, moderately high (0.40 to 0.50 mm) ; the peri- 
stomes all connate in series of 6 (5 to 7), their apertures rounded or 
slightly quadrate, 0.09 to 0.10 mm in diameter. There are no free 
peristomes or vestigial tubules at the lateral margin. 

The ovicell is very elongate, its lobes occupying 7 interfascicular areas 
on each side and extending laterally the width of the lobe. The ooecio- 
pore is located in the usual position beside the first peristome of a fascicle, 
but unfortunately the ooeciostome is broken away. 

The zoarial characters are much like those of P. veleronis n. sp., but 
the lobe is much narrower. The most important character is the great 
length of the ovicell, covering 7 interfascicular areas on each side, while 
in the 11 specimens of veleronis from the Galapagos Islands there are 
never more than 3. There are no intermediate conditions presented and, 
until contradictory evidence is discovered, this must be considered a 
different species. 

Type, AHFno. 119. 

Type locality, Hancock Station 1064, off Santa Barbara Island, 
southern California, 33°30'01"N, 119°02'20"W, at 27 fms, one colony. 
Also at Station 1143-40, off Portuguese Point, southern California, 
33°44'59"N, 118°22'35''W, at 16 fms, one colony. 

Genus FASGIGULIPORA d'Orbigny, 1847 

Like most of d'Orbigny's descriptions, this one is very brief, "well 
characterized by its shell-like (testaces) branches, smooth exteriorly, 
terminating at the upper extremity in a fascicle of rounded, open cells." 
(Transl.) 

Canu and Bassler, 1920:808, add the following: "This genus differs 
from Frondipora in its long fascicles not arranged on a single side of 
the zoarium." 



NO. 3 OSBURN : EASTERN PACIFIC RRYOZOA — CYCLOSTOMATA 665 

Borg, 1926:303 and 382, gives a complete description of the genotype, 
F. ramosa, including the first information about the ovicell, which had 
not previously been noticed, no doubt for the reason that it is but little 
differentiated from the zooecial tubules. It is very elongate and slender, 
slightly expanded on the side of a fascicle, the aperture terminal and 
directed forward. 

Borg is quite justified in removing this genus from the Frondiporidae, 
as the position and nature of the ovicells are very different. However 
there is no justification for Borg's resurrection of d'Orbigny's "Family 
Fascigeridae," since there appears to be no genus Fascigera, and the pro- 
ancestrula and early development, which were hitherto unknown, are 
similar to those of the Tubuliporidae. 

Fasciculipora pacifica new species 
Plate 70, figs. 1, 2, 3, and 4 

The zoarium is fungiform from a narrow base, the largest colony in 
my possession (somewhat broken) measures about 25 mm in height by 
45 mm in the longest diameter, and the longest fascicles are 30 mm. 
The base, broken away, is evidently small. The primary branches are 
comparatively narrow at the base and gradually enlarge, either branch- 
ing or becoming flabellate or folded into contorted fascicles which fre- 
quently coalesce at their tips or are bridged by small flabellate horizontal 
branches consisting of a few zoids. The surface of the adult colony 
resembles the meandering contortions of the human cerebrum. 

The tubules are excessively elongate, 0.30 to sometimes 0.40 mm in 
diameter, in cross-section compressed and hexagonal, on the surface of 
the fascicles rounded and more or less indicated by separating grooves. 
At the tops of the fascicles the tubes do not project, but on the sides the 
occasional tubes which appear to be left behind in the elongation of the 
branch usually show a definite short peristome which is more or less 
erected and with a round aperture. The walls of the tubules are thickly 
perforated by small pores, but on the bases of the older fascia these are 
obscured by a secondary thickening which is more or less ribbed trans- 
versely. 

The ovicells are little modified and resemble the ordinary tubules so 
much that they are easily overlooked. As they emerge on the lateral 
surface of a fascia they take their place among the normal tubules and 
are only slightly larger. They continue upward on a level with the 
tubules, becoming gradually wider until they are twice or three times 
as wide and with a nearly flat frontal surface. The ooeciostome is a 



666 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

little sub-terminal and consists of a short tube more erect than the 
lateral peristomes and somewhat larger; the aperture is rounded but 
modified by irregular folding of the rim. The terminal portion of the 
ovicell usually extends a short distance beyond the ooeciostome and may 
be slightly lobate, and in one case it even surrounds the peristome of a 
neighboring tubule. Borg (1926:383, text figs. 83 and 84) shows the 
ovicell bifurcating at a branch in F. ramosa, with the ooeciostome 
terminal on one of the branches, but I have not observed this condition 
in F. pacifica. 

Fortunately there are several stages in the development of young 
colonies, two of which show the ancestrula which has not been previously 
observed in this genus. These are typically tubuliporoid, with the first 
zoid emerging from the side and the several succeeding generations of 
zoids encrusting fan-shaped, as in Tubulipora. I would undoubtedly 
have mistaken them for young stages of that genus if I had not had a 
continuous succession of stages as well as the adult condition, from the 
same collection (Station 1193-40), for comparison. The tubules of the 
first few zoids are at first encrusting, then become semierect with elon- 
gate peristomes as in Tubulipora. After 3 or 4 generations of zoids, the 
fascicles begin to make their appearance and the zoarium becomes very 
irregular. D'Orbigny was not far wrong in his belief that this genus 
should be "partie de la meme famille que les Tubulipores" (1847:20). 

Type, AHF no. 121. 

Type locality, Hancock Station 1193-40, Santa Cruz Island, southern 
California, 34°N. Lat., shore collection at low tide, numerous young 
stages encrusting stems, and fragments of the adult stage. Also a large 
colony from San Felipe, Mexico, 31°N. Lat., near the head of the Gulf 
of California, shallow water, presented by Dr. A. E. Noble. 

Family Entalophoridae Reuss, 1869 

Zoarium erect, branched, without joints; zooecial tubes elongate, 
opening on all sides of the rounded stem and branches ; gonozoids usually 
situated near the tip of a branch or below a bifurcation, simple and 
elongate or swollen and perforated by zooecial tubes. The arrangement 
of the tubules on all sides of the cylindrical stem is the easiest diagnostic 
character. 

The first stage of development is encrusting and tubuliporoid, and 
from this small base the erect portion of the zoarium arises. Owing to 
the mode of development there has been much difference of opinion as 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 667 

to whether the family Entalophoridae should be maintained or whether 
Entalophora should be considered a genus of the Tubuliporidae. On 
the basis of the simple ovicell, alone, Entalophora would go very nicely 
under Oncousoeciidae, but Bientalophora Borg, with an expanded ovicell 
surrounding some peristomes, would necessarily be synonymous with 
Diaperoecia Canu. As so few species are known perfectly I am leaving 
the family to stand on the basis of the special zoarial character of a 
round, erect stem, with peristomes opening on all sides. 

Genus ENTALOPHORA Lamouroux, 1821 

Zoarium slender, erect, usually only 4 to 8 series of zooecial tubes 
constitute the stems and branches ; the embedded tubes are very elongate, 
parallel, their peristomes curved sharply outward. Gonozoid simple, 
elongate, sometimes only a little wider than the zooecial tubes, located 
usually just below a bifurcation or near the end of a branch ; the ooecio- 
pore terminal. Genotype E. cellarioides Lamouroux, 1821:81. 

Entalophora symmetrica new species 
Plate 70, figs. 6 and 7 

The zoarium is thick-stemmed, about 1.50 mm in diameter, branched 
twice dichotomously at about 90 degrees, the branches as thick as the 
main stem. The secondary branches are at right angles to the primary 
ones. The tubules are elongate and distinct, with well marked separating 
grooves, the pores numerous and conspicuous; about 12 tubules surround 
the stem equally on all sides and arranged more or less in quincunx. The 
peristomes are moderately high, inclined distally, somewhat tapered 
toward the tips and porous like the frontal for most of their length; 
diameter of the aperture 0.15 to 0.17 mm. 

The ovicell is simple, pyriform, not extended between the peristomes, 
moderately inflated, its surface smooth; the ooeciostome is terminal, 
slightly elevated, more erect than the peristomes and slightly smaller, 
its aperture rounded and 0.12 mm in diameter. 

This species has the zoarial form of a Bientalophora, but there are 
no covering kenozooecia, the tubules are evident throughout their length, 
and the gonozoid is of the simple type characteristic of Entalophora. 

Type, AHF no. 122. 

Type locality, Hancock Station 170-34, Stephens Bay, Chatham 
Island, Galapagos, 0°47'30''S, 89°31'W, at 32 fms, one colony without 
base. 



668 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Entalophora proboscideoides Smitt, 1872 

Plate 70, figs. 8 and 9 

Entalophora proboscideoides Smitt, 1872:11. 

Entalophora proboscideoides, Canu and Bassler, 1928:160. 

Entalophora proboscideoides, Osburn, 1947:4. 

Zoarium erect, slender, branching widely, the stem composed of 6 to 
8 very elongate tubules; the embedded tubules about 0.13 mm in di- 
ameter and the peristomes, which open on all sides of the stalk, about 
0.10 mm. The longest peristomes are about 0.50 mm, perforated and 
lightly wrinkled like the embedded tubules. 

The ovicell is simple, a distinct elliptical swelling of the distal end of 
a long tubule, 0.55 mm long by 0.35 mm wide, thickly perforated. The 
peristome is terminal, bent I'orward sharply, the aperture transversely 
elliptical, 0.13 by 0.06 mm. 

Described by Smitt from west of the Tortugas Islands, Florida, at 
68 fms. Recorded by Osburn (1947:4) from 8 stations along the 
southern shore of the Caribbean Sea (Hancock Atlantic Expedition, 
1939), and by Canu and Bassler (1928:160) from the Pliocene of 
Bocas Island, Panama. Our one ovicelled specimen appears to agree in 
all details with those from the Caribbean Sea. 

Hancock Station 457-35, Secas Islands, Panama, 12 fms. 

Entalophora capitata Robertson, 1900 

Entalophora capitata Robertson, 1900:328 (Plate 21, fig. 12 only); 

1910:257. 
Entalophora capitata, O'Donoghue, 1923:13. 
Diaperoecia capitata, O'Donoghue, 1926:22. 

Dr. Robertson's 1900 description is practically worthless as she con- 
fused this form with another species which Borg has since described 
(1933:325) as Heteropora pacifica alaskensis. In 1910 Robertson cor- 
rected the error and based her re-description on the specimen from which 
figure 12 of her former account was taken. O'Donoghue in 1926 placed 
the species under Diaperoecia, where, if only the nature of the ovicell 
is considered, it would seem to belong. The species has not appeared in 
the Hancock collections and I am unable to form a definite opinion. 

Robertson's 1910 description is as follows: 
"Zoarium arising from a flattened or encrusting base and growing from 
5 to 8 mm in height. Zooecia tubular, uniting in a short, stout column 
terminating in a broad somewhat rounded head ; distal ends free, usually 
extending for a considerable distance beyond the surface of the colony. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 669 

both of the supporting column and of the head. Ooecium an inflation 
of the surface of the head. Ooeciostome and ooeciopore slightly com- 
pressed, opening beside the zooecial aperture." Orca, Prince William 
Sound and Sitka, Alaska. 

O'Donoghue listed it from several localities in British Columbia. 

Entalophora sp. 

Zoarium slender, nearly straight, 4 or 6 tubules constituting the stem, 
width 0.75 to 0.90 mm ; the peristomes elongate, nearly at right angles 
to the stem axis, perforated like the tubules nearly to their tips. On the 
surface the tubules are more or less distinct, the whole surface trans- 
versely wrinkled and perforated with small pores ; on the older part of 
the stem the peristomes also are wrinkled on the basal portion. There 
is a tendency toward spiral arrangement, though 2 or 3 peristomes may 
arise at nearly the same level. Width of stem 0.75 to 0.90 mm; width 
of tubules on the stem 0.30 mm; width of apertures 0.16 to 0.20 mm; 
longest peristome 0.65 mm but the average about 0.25 mm. 

The specimen consists of part of a stem 20 mm in length, both base 
and tip wanting and without an ovicell. It has some resemblance to 
E. proboscideoides Smitt, 1872:11, but it is much larger, the apertures 
nearly twice as broad. The large size of the tubules, the width and 
length of the peristomes and the coarse transverse striation of the stem 
seem to indicate it as an undescribed species, but in the absence of an 
ooecium I hesitate to give it a name. 

Hancock Station 450, Cartago Bay, Albemarle Island, Galapagos, 
0°5yS, 90°30^W, at 70 fms. 

Entalophora raripora d'Orbigny is listed by Robertson 1910:256 from 
Monterey, California, and by O'Donoghue, 1923:13, from several 
places in British Columbia. 

Entalophora clavata Busk is also recorded by O'Donoghue, 1923:13, 
from several British Columbia localities. 

Entalophora vancouverensis O'Donoghue, 1923:13, is described and 
recorded for Cardale Point, Round Island, British Columbia. From its 
appearance as judged by figure 7 (plate 1) it may be a species of Bi- 
entalophora, but O'Donoghue does not mention the presence of kenozoids 
on the stalk. 

The Entalophoras are evidently much in need of a thorough restudy. 



670 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Genus BIENTALOPHORA Borg, 1944 

"Zoarium erect, branching repeatedly, branches originating through 
forking of the stem ; zoarium composed of autozoids, kenozoids and 
gonozoids. Kenozoids smaller and shorter than autozoids, always closed, 
numerous, forming greatest part of surface of zoarium. Autozoids pro- 
truding through layer of kenozoids, distal portions of their cystids ar- 
ranged in quincunx or spirally, opening all around the stem. Gonozoids 
with middle portion large, strongly dilated, traversed by numerous auto- 
cystids; distal portion seemingly not terminal." (Borg, 1944:114). 
Genotype Entalophora regularis MacGillivray, 1887:219. 

The two striking characters which distinguish this genus are: (1) 
the greater development of the ovicell which extends over a capitulum 
or broader area of the stem and surrounds some peristomes, and (2) 
the presence of a thin layer of completely closed small kenozoids over 
the zoarium between the peristomes. 

The nature of the ovicell, enclosing peristomes, would place the mem- 
bers of this genus in Diaperoecia Canu, but as this character appears in 
a number of other genera which are zoarially quite distinct, I have come 
to the conclusion that parallel evolution may apply to the development 
of the ovicell as well as to the zoarium. 

Bientalophora cylindrica new species 
Plate 70, figs. 10 and 11 

The zoaria are erect, with round straight stems and branches 1.50 
to 1.60 mm in diameter; dichotomous, the branches diverging at an 
angle of about 60 degrees, the distance between branches about 1.50 
cm; in one case two branches have fused where they came in contact. 
The branches are very slightly widened toward the tip. The bases of 
both of our colonies are broken away, but the remaining longest portion 
measures 17 mm in length. The general appearance is much like that 
of a Myriozoum. 

The zooecia are distributed all around the stem, irregularly quin- 
cuncial, their tubes not visible on the surface. The peristomes are very 
short, the longest not more than 0.10 mm high, 0.15 to 0.17 mm in 
diameter, the apertures 0.12 or 0.13 mm. There are frequent small areas 
which are free from peristomes. The whole surface between the peri- 
stomes, clear up to the growing tips, is covered by a layer of small 
kenozooecia which are thickly perforated by small pores and their out- 
lines marked by slightly raised lines. These kenozooecia form the 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 671 

"lamina" which Waters, 1914:842, described under B. (Entalophora) 
regularis. The kenozooecia are smaller than the autozooecia, irregular 
in size and form but usually somewhat diamond-shaped. 

There are no ovicells on our specimens so complete identification is 
impossible. There is considerable resemblance to B. regularis (Mac- 
Gillivray), the genotype, as figured by Borg (1944, plate 11, figs. 3 
and 4), but the diameter of the apertures is distinctly smaller, the 
diameter of the stems somewhat greater, and the peristomes noticeably 
shorter. Since B. regularis is known only from the Australian area, it 
seems preferable to give this California form a name. 

Type, AHF no. 123. 

Type locality, Monterey Bay, California, 36°N, 122°W, at 40 fms, 
F. P. Shepard, collector, two fragments. 

Family Frondiporidae Busk, 1875 

The zoarium consists of an encrusting, branched, ramifying base from 
which arise erect cylindrical fascicles which are usually separated by 
well-marked interfascicular spaces. The ovicell is developed between 
the fascicles, either simple or lobate, and is sometimes perforated by 
one or more tubules ; the ooeciostome is but little elevated, in Filifascigera 
remote from any of the zoecial tubules, while in Frondipora Borg (1926, 
text fig. 81) shows it as a crescentic pore adjacent to the base of a tubule. 

Genus FILIFASCIGERA d'Orbigny, 1852 

"The colonies are creeping, narrow, linear, or curved. The tubes are 
grouped in salient, orbicular, or elliptical fascicles, regularly spaced. 
The orifices are polygonal. The ovicell is a vesicle placed between the 
fascicles and perforated by closed tubes." (Canu and Bassler, 1929:523). 
Genotype, Filifascigera dichotoma d'Orbigny, 1852. 

This genus has been much neglected since d'Orbigny's time and until 
very recently has been known only as a fossil. Canu and Bassler (1928: 
44) described F. robusta from Hawaii and found the ovicell for the 
first time. Later (1929:524) they described two other recent species, 
F. pluripora and F. parvipora, from the Philippines. 

The above generic description requires a few additions. The orifices 
are not polygonal except in the fascicle and at its tip where the tubules 
are compressed together. The free peristomes, which rise as much as 0.40 
mm above the tips of the fascicles, are cylindrical with circular apertures, 
all separated and curving outward. The fascicles are not always evenly 



672 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

spaced and on a rough background may be quite irregular in distribution, 
anywhere from 0.20 mm to more than 1.00 mm apart. Apparently the 
genus is widely distributed over the central area of the Pacific. 

Filifascigera clarionensis new species 
Plate 69, figs. 8, 9, and 10 

The zoarium is encrusting, tortuous, the branches narrow, averaging 
about 0.60 mm in width between the fascicles. The basal portions of 
the zooecial tubules are completely embedded with no separating grooves, 
the distance between the fascicles varying greatly, from 0.26 to 1.10 
mm and averaging about 0.60 mm. The fascicles, or bundles of tubules, 
are nearly erect to a height of 0.40 to 0.50 mm, round or elliptical in 
cross-section, and consist of 2 to 8 tubules (6 is a characteristic number). 
The outlines of the tubules are evident only on the upper part of the 
fascicles. At the top of the fascicles the tubules end in free peristomes 
which are cylindrical, uniform in diameter or slightly flaring at the tips, 
and curved outward from the center, the longest being as much as 0.40 
mm. The apertures are circular, about 0.11 mm in diameter, and the 
peristomes 0.14 mm in diameter. 

The ovicell is a rather conspicuous non-lobate swelling between the 
fascicles and extending from one fascicle to the next, the peristomial 
tubes rising above its level ; it measures about 0.55 by 0.80 mm. The 
ooeciostome is somewhat off-center, a short tube which flares outward at 
the edges, with a short-elliptical aperture; it is completely dissociated 
from any of the zooecial tubules. 

The description is taken from two colonies from Clarion Island, one 
on a worm tube, the other on a coralline. Another colony from Santa 
Barbara Island, southern California, without an ovicell, appears to be 
the same, as the measurements agree, and it also encrusts a worm tube. 

Type, AHF no. 125. 

Type locality, Hancock Station 137-34, Clarion Island, 18°19'05"N, 
114°44'25''W, at 25 fms. Also at 1067, Santa Barbara Island, southern 
California, 33°22'30''N, 119°03'45"W, at 55 fms; another at 1624-48, 
Santa Catalina Island, 36 fms, on a shell, and still another at 1914-49, 
San Cristobal Bay, Lower California, 27°24'48''N, 114°34'40''W, at 
40 fms. 

Filifascigera fasciculata (Hincks), 1880 

Stomatopora fasciculata Hincks, 1880:441. 
Proboscina fasciculata, O'Donoghue, 1926:17. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 673 

O'Donoghue gave no description of this species, but his illustration 
(plate 2, fig. 12) certainly shows a Filifascigera and it may well be the 
S. fasciculata of Hincks. The description of the species by Hincks is very 
complete, showing the erect arrangement of the tubules in bundles, well 
spaced and elevated, and he also figures the ovicell (plate 59, fig. 4) set 
between the fascicles with the ooeciostome off center and separated from 
the tubules. The ovicell has much the same appearance as that of F. 
clarionensis new species, described above, but the ooeciostome is much 
compressed and its pore almost slit-like. 

The only question is whether O'Donoghue's species is that of Hincks, 
and that cannot be determined here as the species has not appeared in 
the Hancock material. 

O'Donoghue records the species from Northumberland Channel and 
Gabriola Pass, British Columbia, and the San Juan Islands, Puget 
Sound. 

A specimen from southern Alaska, U. S. Fisheries Alaska Crab In- 
vestigation, Sta. 82-40, may belong here, but in the absence of complete 
ovicells the identification is questionable. It is a much larger species 
than the preceding, the apertures measuring 0.16 to 0.18 mm in diameter. 
The one ovicell is properly located for this genus, but is incomplete and 
lacking the ooeciostome. 

? Filifascigera sp. 

Another species which probably belongs to this genus but may be a 
Frondipora was taken at Hancock Station 1914-49, off Guadalupe 
Island, west of Lower California, 28°52'N, 118°19'W, at 5-15 fms. 
The fascicles are larger and higher than those of F. clarionensis (the 
aperture 0.14 mm in diameter), and several fascicles sometimes arise 
from a single base to form a complex fascicle. The peristomes usually 
rise free above the top of the connate portion of the fascicle, as they do 
in F. clarionensis. The ovicell lies in the space between the fascicles of 
a complex fascicle with lobes extending among them. Unfortunately 
there is no evidence of an ooeciostome. 

The material is too imperfect for positive identification but it is cer- 
tainly different from either of the species mentioned above, especially 
in the nature of the complex fascicles and the ovicell. The small fascicles, 
usually of less than 8 tubules, would seem to remove it from Frondipora. 



674 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Division 2. Articulata Busk, 1859 

(Camptostega Borg, 1926) 

The Crisias 

"Primary zoid erect, separated by a chitinous joint from the pro- 
ancestrula; zoarium jointed; rhizoids present. Body wall a gjonnocyst; 
vestibular sphincter present ; brood chamber a gonozoid, moderately 
dilated in its middle part; polypide of gonozoid degenerating before 
having been fullgrown." (Borg 1944:133). 

This division is clearly distinguished by the jointed zoarium, by the 
presence of rhizoids, which are formed of jointed series of kenozoids, 
and by the mode of development from the larva, all of which are differ- 
ent from the other divisions. On attachment the larva forms a dome- 
like structure, on the top of which the first functional zoid is produced 
and from which it is separated by a chitinous joint. There is only one 
family, the Crisiidae, with a number of genera, depending chiefly on 
the structure of the internodes. 

The erect, jointed zoarial form is so strikingly different from any 
other cyclostome type of growth that the members of this Division are 
easily placed at once. 

Family Grisiidae Johnston, 1838 

The zoarium is erect and jointed, the zooecia in a single series or 
alternating in two series, or without definite arrangement in the older 
branches of Crisulipora; the internodes consist of one zooecium to many ; 
attached by jointed radicles, rhizoids, which consist of a series of elon- 
gated, tubular kenozooecia. The ovicell is an enlarged zooecium (gono- 
zoid) more or less pyriform in shape, with the ooeciostome (pore) 
terminal or nearly so. The characters of the ovicell and its ooeciostome 
are essential for the positive determination of most of the species of 
this family. 

Key TO THE Genera of Crisiidae 

1. Internodes of 1 or 2 zooecia ; elongate filiform spines present . . 2 
Internodes with 1 to many zooecia ; no filiform spines .... 3 

2. Only one zooecium to an internode Crisidia 

Two zooecia to an internode, fertile internodes may have 3 

to 5 ; the ovicell is free for much of its length and the 
ooesciostome is on the dorsal side Bicrisia 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 675 

3. Sterile internodes of 1 to 3 zooecia, fertile ones of 3 to 5; 

ovicell adnata for its whole length and its ooeciostome 

terminal Filicrisia 

Internodes of 3 to many zooecia, usually 5 or more except at 
the base ; ovicell adnata for its entire length, the ooeciostome 
more or less terminal 4 

4. Internodes with 2 alternating series of zooecia ; ovicell promi- 

nent, expanded dorso-ventrally Crista 

Internodes with 2 to 8 or more zooecia in cross-section, not 
regularly arranged ; ovicell expanded laterally between the 
tubules Crisulipora 

Genus GRISIDIA Milne-Edwards, 1838 
Genotype, Sertularia cornuta Linnaeus, 1758. 

Crisidia cornuta (Linnaeus), 1758. 
Plate 71, fig. 1 

Sertularia cornuta Linnaeus 1758:810. 
Crisia cornuta, Hincks, 1884:203. 
Crisia cornuta, O'Donoghue, 1923:7. 
Crisidia cornuta, O'Donoghue, 1926:18. 
Crisidia cornuta, Borg, 1926:260 and 349. 

The zoarium is delicate, branching dichotomously, each branch con- 
sisting of a single series of zooecia, each zooecium constituting an inter- 
node. The zooecium is slender, elongate (0.60 to 0.80 mm long), dis- 
tinctly arcuate ; the succeeding zooecium arises on the dorsal side toward 
the distal end, paired when branching. Long filiform processes (1.0 
mm or longer) often arise beside the zooecial base, jointed at the base 
and usually twice more ; from their position and mode of origin these 
processes would seem to be vestigial zooecia; at any rate they are not 
homologous with the spines of other bryozoan orders. 

The ovicell is a gonozoid, free with a terminal ooeciostome, and rep- 
resents an internode ; it never bears a spinous process. 

Hincks and O'Donoghue have listed this species from several localities 
in British Columbia waters ; otherwise it has not been noticed previously 
on the Pacific coast. 

Hancock Stations: 1269-41, off Anacapa Island, at 41 fms; 1279-41, 
off San Miguel Island, at 40 fms-; and 2042-51, off Long Beach Light, 
at 14 fms; all from southern California. Also off Pescadero Point, 
outside of Monterey Bay, California, A. E. Blagg, collector. 



676 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Genus BIGRISIA d'Orbigny, 1853 
Genotype, Crisidia edwardsiana d'Orbigny, 1839. 

Bicrisia edwardsiana (d'Orbigny), 1839. 
Plate 71, fig. 2 

Crisidia edwardsiana d'Orbigny 1839:8. 
Crisia cornuta, Robertson, 1900:328, 
Crisia edwardsiana, Robertson, 1910:237. 
Crisidia edwardsiana, O'Donoghue, 1926:18. 
Bicrisia edwardsiana, Borg, 1926:260 and 351. 

The zoarium is usually bushy, much branched, reaching a height of 
50 to 75 mm, the tips of the branches curved forward. The zooecia are 
tubular, 0.50 to 0.70 mm long, somewhat arcuated, usually 3 to 5 to 
an internode (basally 1 or 2). Jointed "spinous processes," 1.0 mm or 
more in length, similar to those of Crisidia, are apparently vestigial 
zooecia. 

The ovicell is somewhat elliptical in form and is free for nearly its 
entire length ; a characteristic feature is the position of the ooeciostome 
on the dorsal side near the distal end. 

Robertson evidently confused this species with Crisidia cornuta. Her 
specimens with "Internodes consisting typically of a single zooecium" 
must have been cornuta, while those of "two, three, four or five zooecia" 
are undoubtedly edwardsiana. Her figure of the ovicell, 1910, plate 19, 
fig. 10, is definitely edwardsiana. 

Robertson listed this species (and cornuta) from Alaska to San Diego, 
California. O'Donoghue recorded it from the San Juan Islands, Puget 
Sound. It appears to be more common than cornuta on the California 
coast and extends farther southward. 

Hancock Stations: 1320-41 and 1370-41, off Santa Catalina Island, 
shore to 18 fms; 1210, at La Jolla, shallow water; San Diego Jetty, 
shore (Dr. H. R. Hill) ; Newport Harbor on floats (R. C. Osburn) ; 
all from southern California. Also at 843-38, Lobos de Afuera Islands, 
Peru, 6°53'50"S, 80°43'30"W, at 25 fms. This beautiful but incon- 
spicuous little species is probably more common than the number of 
stations indicates and possibly extends along the whole Pacific coast. 

Genus FILIGRISIA d'Orbigny, 1853 
Genotype, Crisia genie ulat a Milne-Edwards, 1838 

In younger stages of growth this genus resembles Bicrisia except for 
the absence of the soinous orocesses. but older soecimens are readilv 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 677 

distinguishable by the conspicuous black joints and by the ovicell, which 
is more slender, adnate to the internode for its full length, and with a 
terminal ooeciostome. 

Filicrisia geniculata (Milne-Edwards), 1838 
Plate 72, fig. 5 

? Crisidia gracilis Trask, 1857:113. 

Crista geniculata, Robertson, 1910:235. 

Crisia geniculata, O'Donoghue, 1923:7; 1926:18. 

Filicrisia geniculata, Borg, 1926:263 and 351. 

Zoarium bushy, the branches nearly straight, reaching a height of 15 
to 25 mm. The zooecia are tubular, straight, 0.60 to 0.90 mm long, 
3 to 5 to an internode (except basally). In the older zooecia the joints 
are black and conspicuous. 

The ovicell is long and but little inflated, adnate for its entire length, 
the ooeciostome situated near the dorsal border with its tube bent for- 
ward (sometimes straight). In the absence of the ovicell it is impossible 
to distinguish positively between this species and franciscana. For this 
reason the Crisidia gracilis of Trask must remain in doubt, though it 
appears to be one of these two species. 

Robertson first noted the presence of this species on the Pacific coast 
and recorded it from Dillon Beach, north of San Francisco, to San 
Pedro, California. O'Donoghue listed it from numerous localities in 
British Columbia. 

Hancock Stations: Not taken in dredging, but found at numerous 
shore stations in shallow water about the islands of southern California 
and along shore from Monterey Bay south to San Pedro, California; 
well distributed in this area, but never abundant. It appears to be more 
common farther northward. 

Filicrisia franciscana (Robertson), 1910 

Plate 72, fig. 4 

Crisia franciscana Robertson, 1910:233. 
Crisia occidentalis, Robertson, 1903:116. 
? Crisidia gracilis Trask, 1857:113. 
Crisia franciscana, Okada, 1917:338. 
Crisia franciscana, O'Donoghue, 1923:7. 
Crisidia franciscana, O'Donoghue, 1926:19. 

This species resembles C. geniculata in practically all respects except 
for the ovicell. The zoarial form and the number of zooecia to the 



678 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

internode are the same, and both have the characteristic black joints. 
The zooecia of franciscana are slightly larger and are usually more 
expanded toward the distal end. 

The ovicell is more inflated than in geniculata and the ooeciostome 
is situated on the frontal border with its tube curved backward. 

Reported by Robertson from Orca, Alaska, to southern California at 
numerous places, and by O'Donoghue from British Columbia. It is 
one of the most common crisias along the California coast from low 
tide to 25 fms. Okada also found it common in Japanese waters. 

Hancock Stations: Numerous stations about the islands and along 
the coast of southern California from San Francisco southward to San 
Diego; Dillon Beach, north of San Francisco (R. J. Menzies) ; Mussel 
Point, northern California (A. E. Blagg) ; San Juan Islands, Puget 
Sound (J. L. Mohr) ; much more abundant than C. geniculata, often 
occurring in considerable masses on piles and floats ; low tide to 50 fms. 

Genus CRISIA Lamouroux, 1812 

Genotype, Sertularia eburnea Linnaeus, 1758 :810. In this well-known 
genus the internodes are longer, with 5 to as many as 30 or more zooecia 
in some of the species. These are arranged very symmetrically in two 
alternating series, the short projecting peristomes giving the edges a 
serrated appearance. The ovicells or gonozoids are usually in the mid- 
line of the frontal surface, between the rows of zooecia. It is unfor- 
tunate that the ovicells, which are often lacking, are necessary for the 
positive determination of most of the species. Dr. Alice Robertson gave 
an excellent account (1910:229-245) of the Pacific coast species known 
to her and her key is used here, with additions and slight modifications. 

The little shrub-like colonies of the crisias are often abundant in 
shallow water, attached to anything that may afford a lodging place 
and conspicuous because of their chalky whiteness. 

Key to the Species of Crisia 

1. Ooeciostome with a cap-like flap extending forward above 

the aperture operculata 

Aperture of ooeciostome without covering flap 2 

2. Ovicell very short and wide, ooeciostome almost wanting, pore 

round ; internodes long and slender elongate 

Ovicell elongate and gradually expanding 3 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 679 

3. Ooeciostome curved or bent forward 4 

Ooeciostome straight, though the opening may be directed 

somewhat forward 6 

4. Branches of zoarium strongly curved forward; ooeciostome 

curved forward, its aperture elh'ptical and its proximal lip 

somewhat inflected eburnea 

Branches straight and more divergent 5 

5. Ooeciostome long, slender, conspicuously bent forward, its 

pore round pugeti 

Ooeciostome short, pore elliptical; internodes long . . serrulata 

6. Branches of zoarium curved inward or spicate at the tips; 

ooeciostome short, pore round or short elliptical . . occidentalis 
Branches straight and stiff, internodes long 7 

7. Ooeciostome distinctly flared at the tip, transversely long- 

elliptical; a northern species cribraria 

Ooeciostome not flared, short and inconspicuous, pore round 

and opening forward maxima 

Grisia serrulata, new name 
Plate 72, fig. 2 

Crisina serrata Gabb and Horn, 1862:174. (Preoccupied by d'Orbigny, 

1853:598). 
? Crisia denticulata, Hlncks, 1884:203. 
Crista pacifica Robertson, 1910:242. 
Crisia pacifica, O'Donoghue, 1923:7. 
Crisia serrata, Canu and Bassler, 1923:196. 
Crisia serrata, O'Donoghue, 1926:18. 

Zoarium forming bushy tufts reaching a height of 25 mm. The inter- 
nodes are long, ranging from 12 to more than 30 zooecia, the longer 
ones slightly sinuate and not inflected ; joints yellow to brownish ; basis 
rami of a branch short and wedged in between the zooecia without dis- 
turbing their position. The zooecia are connate nearly to their tips which 
turn forward sharply, the aperture facing frontally; the dorsal lip of 
the aperture sometimes with a low point. The frontal surface of the 
branch bears a median keel and the distance between the zooecial aper- 
tures is less than the width of the branch. 

The ovicell is large, a little flattened, inclined in the axis of the branch 
and adnate for its whole length; the tube of the ooeciostome is short, 
opening either ventrally or distally, the aperture more or less elliptical. 



680 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Gabb and Horn described the species from the Pleistocene of Santa 
Barbara, California, overlooking d'Orbigny's previous use of serrata; 
Canu and Bassler listed it from the Pleistocene of Santa Barbara and 
Santa Monica, and the writer has found it in the Pleistocene of San 
Pedro and Newport Harbor, California. Robertson described pacifica 
from the "San Diego region only." O'Donoghue recorded it from numer- 
ous British Columbia localities, and it is a common species all along the 
coast from British Columbia southward to Cedros Island (28°N), 
and less commonly to the Galapagos Islands. 

Hancock Stations: Dredged at more than 30 stations and taken at 
numerous shore stations, most abundant about the island region off 
southern California. Galapagos Islands, 5 stations: 152-34, Albemarle 
Island, shallow water; 170-34, Chatham Island, 32 fms; 193-34 and 
198-34, Charles Island, 10-65 fms, and 804-38, Onslow Island. Also 
at 1051-40, Angel de la Guardia Island, Gulf of California, 21 fms. 

Crisia occidentalis Trask, 1857 
Plate 71, figs. 3, 4, and 5 

Crisia o ccidentalis Trask, 1857:113. 

Crista eburnea, Robertson, 1903:116. 

Crisia occidentalis, Robertson, 1910:239. 

Crisia occidentalis, O'Donoghue, 1923:7; 1926:18. 

Zoaria forming dense tufts reaching a height of 25 mm, the tips of 
the branches often inflected, especially in ovigerous colonies. The inter- 
nodes consist of 3 to 5 zooecia near the base, the more terminal ones 
from 7 to 12; joints white to yellow; basis rami not wedged in between 
the zooecia but extending along the outer side of its mother zooecium, 
though there is some variation in this respect. The frontal surface of 
the internode is slightly keeled and the distance between zooecial aper- 
tures is about equal to the width of the branch. The zooecia are connate 
for their entire length, the tips directed forward ; frequently there is a 
short point back of the dorsal lip of the tube, and the tips of the terminal 
branches often end in spinous points. 

The ovicell is moderately large, elongate pyriform, inclined in the 
axis of the internode ; the ooeciostome is situated a little back of the 
summit of the ovicell, with a short, straight or slightly curved tube, the 
circular aperture opening more or less upward. 

One might conclude from Robertson's discussion that ovigerous 
colonies have only inflected branches and that only the male colonies 
have the terminal spinous points. This is not the case, however, as 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 681 

ovigerous colonies frequently bear the pointed tips and their branches 
may be perfectly straight. Sex differentiation does not seem to be the 
answer to these variations. 

Trask described the species from San Francisco, very inadequately. 
Miss Robertson accepted the name and redescribed the species, recording 
it from Puget Sound, Washington, to San Pedro, California. O'Donog- 
hue listed it from Banks Island and Gabriola Pass, British Columbia. It 
is a common species along the California coast from low water to 30 
fms, and south rarely to the Galapagos Islands. 

Hancock Stations: Dredged at 12 stations, mostly about the islands 
ofl southern California. Station 470-35, Port Parker, Costa Rica, 5 
fms, and 85-33, North Seymour Island, Galapagos, shore collection. 
The number of the dredging stations does not indicate the abundance 
of the species, as it is much more common in shallow water near shore. 

Grisia operculata Robertson, 1910 
Plate 71, figs. 6 and 7 

Crisia operculata Robertson, 1910:240. 
Crista operculatdj O'Donoghue, 1923:7. 

The zoarium is fragile, with irregular tufts reaching a height of 
about 20 mm; internodes consist of about 10 to 20 zooecia, though the 
number may reach 30 or more; the frontal surface rounded but not 
keeled, the basis rami exposed for most of its length. The zooecia are 
very slender, connate for most of their length, though the free tips are 
longer than in most crisias. The distance between the zooecial apertures 
is considerably greater than the width of the internode. 

Ovicell elongate pyriform, inclined to one side of the internodal axis; 
"the dorsal wall of the ooecium extending upward and forward covering 
the ooeciostome as with a lid or cap, the operculum" (Robertson). The 
ooeciopore is a semicircular slit beneath the cap. 

The species was described from "one station on the southern Cali- 
fornia coast, depth not known." O'Donoghue recorded it from Houston 
Passage, British Columbia, 15 fms. 

Hancock Stations: dredged at only 4 stations: 1378-41, Santa Cata- 
lina Island, southern California, 2-3 fms; 1049-40, Angel de la Guardia 
Island, Gulf of California, shore ; 675-37, Pulpito Rock, Gulf of Cali- 
fornia, 55 fms, and San Francisco Island, Gulf of California, 47 fms, 
24°47'35"N, 110°35'55'''W, the most southern record. Apparently it 
is not a very common species. 



682 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Crisia maxima Robertson, 1910 
Plate 72, fig. 3 

Crisia maxima Kohcrtson, 1910:243. 

Crista maxima, O'Donoghue, 1923:7, 1926:18. 

Zoarium coarse, stiff, with straight, long internodes, resembling C. 
serrulata in its manner of growth, but coarser and larger, occasionally 
more than 50 mm in height. The internodes are elongate, usually with 
from 12 to 20 zooecia but may contain more than 40; older joints dark 
brown; basis rami not wedged in between the zooecia. The zooecia are 
closely connate to their tips, which are turned sharply forward; the 
distance between their apertures is greater than in serrulata, usually dis- 
tinctly greater than the internodal width. The front of the internode is 
slightly arcuate in cross-section and not keeled. 

The ovicell is large, the frontal surface prominent, the distal end 
more or less truncate; the tube of the ooeciostome is short, straight, 
slightly tapered and opens more or less ventrally. 

Recorded by Robertson on the southern coast of California from be- 
tween tide marks at Escondido, Deadmans Island (San Pedro), and 
White's Point, and dredged from San Pedro to Coronado Island down 
to 40 fms. O'Donoghue lists it from several British Columbia localities 
down to 25 fms. 

Hancock Stations: 31-33 and 362-35, Hood Island, Galapagos, 1°22' 
52"S, 89°39'15"W, at 4 to 20 fms (the most southerly record) ; also 
at 352-35, Chatham Island, 35 fms; and 810-38, Barrington Island, 48 
fms, Galapagos. Station 1051-40, Angel de la Guardia Island, Gulf of 
California, 21 fms; 870-38, Isabel Island, west of Mexico, 10-15 fms; 
and 894-38, San Miguel Island, 5-15 fms, 1238-47, San Clemente 
Island, 14 fms, and 1232-47, off San Pedro, 18 fms, southern California. 
Also off Pescadero Point, near Monterey Bay, California (A. E. Blagg, 
collector) . 

Crisia ebumea (Linnaeus), 1758 
Plate 71, fig, 10 

Sertularia eburnea Linnaeus, 1758:810. 
Crisia eburnea, Hincks, 1880:420; 1884:203. 
Crisia eburnea, Osburn, 1912:215; 1923 :5D. 

Zoarium forming dense bushy tufts, the branches characteristically 
curved forward. The internodes are short, usually 5 to 7 zooecia, the 
joints yellow, sometimes dark near the base. Zooecia almost entirely 
connate, the short free portions nearly at a right angle to the tubules. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 683 

Ovicell large, curved forward, usually replacing the second zooecium 
of an internode ; the ooeciostome curved forward, widest at its base, the 
pore transversely elliptical and the proximal margin somewhat inflected. 

This is a very common species on the coasts of Europe and North 
America (Atlantic coast) and entering the Arctic Ocean, common in 
Greenland waters. Recorded for Icy Cape and Point Barrow, Alaska 
by Osburn, 1923 :5D. Hincks (1884:203) reported it from Virago 
Sound, British Columbia, but this record appears to be questionable as 
the species has not been recovered south of northern Alaska, and he may 
well have confused it with C. occidentalis Trask, which has the same 
growth form of incurved branches and is common in British Columbia 
waters. 

Point Barrow, Alaska, 18 fms, Arctic Research Laboratory, G. E. 
MacGinitie, collector. 

Crisia cribraria Stimpson, 1853 
Plate 72, fig. 1 

Crisia cribraria Stimpson, 1853:18. 
Crisia eburnea var. cribraria, Verrill, 1879:28. 
Crisia eburnea var. cribraria, Whiteaves, 1901:110. 
Crisia cribraria, Osburn, 1912:215; 1912a :276; 1933:8. 

The zoarium consists of nearly erect, straight and stiff flabellate 
branches rising to a height of 20 to 25 mm. The internodes are long, 
usually about 18 or 20 zooecia, the joints occasionally wanting. The 
zooecia are almost completely fused, with only a very short peristome 
which curves abruptly forward and slightly toward the axis of the 
branch, a sharp projection often present on the outer border of the 
aperture. 

The ovicells are large, elongate, the distal end prominently rounded 
and more or less obscuring the ooeciostome from a frontal view. The 
ooeciostome is short and broad, the aperture almost slit-like, the tip 
somewhat flared outward. 

Stimpson described the species from Grand Manan Island, Maine, 
and it is a fairly common species along the east coast of North America 
as far south as Cape Cod, 

Point Barrow, Alaska, 7 fms, Arctic Research Laboratory, G. E. 
MacGinitie, collector; two colonies in reproduction. 



684 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Grisia elongata Milne-Edwards, 1838 
Plate 71, fig. 9 

Crista elongata Milne-Edwards, 1838:203. 
Crista elongata, Harmer, 1915:96 (synonymy). 
Crisia elongata, Canu and Bassler, 1928:157. 
Crisia elongata, Osburn, 1940:328; 1947:3. 

Zoarium with long, slender, sprawling branches; the internodes elon- 
gate, usually with about 14 to 16 zoids but ranging from 6 to 30 or 
more, the joints of both branches and radicles jet black (brownish in 
younger areas of the colony). The tubules of the zooecia are embedded 
and their outlines scarcely visible, their peristomes short and turned 
sharply forward, with usually a small denticle behind the distal border. 

The ovicell is situated usually near the middle of an internode, short, 
suddenly and broadly inflated ; the ooeciostome is little or not at all 
elevated and its pore is a transverse slit. 

Our rather scanty material agrees well with Harmer's excellent de- 
scription (1915:96) and with specimens from the West Indian region. 
It is my opinion that the C. eburnea forma denticulata of Smitt (1872: 
4), the C. denticulata of Osburn (1914:185) and the C. denticulata of 
Canu and Bassler (1928:156), all from the Gulf of Mexico and the 
West Indian region, should be referred to C. elongata. 

Hancock Station, 277-34, off Isabel Island, Gulf of California, 21° 
5 r3 5'"N, 105°30'W, at 10-25 fms. It is apparently a circumtropical 
species. 

Grisia pugeti Robertson, 1910 
Plate 71, fig. 8 

Crisia pugeti Robertson, 1910:244. 

Crisia pugeti, O'Donoghue, 1923:8; 1926:18. 

The zoarium is rather stiff and straggling in appearance, the inter- 
nodes varying greatly from 7 to more than 30 zooecia. The joints are 
colorless or slightly brownish in older zoaria. The branches are rather 
numerous, 3 or 4 to an internode, the basis rami usually exserted but 
sometimes short, always a branch immediately above the top of the 
ovicell. 

The ovicell is usually situated low in the internode, most frequently 
the third member of the internode; elongate, expanding rapidly near 
the base and maintaining the same width for most of its length ; con- 
siderably inflated and adnate to the internode for its full length. The 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 685 

ooeciostome is longer than usual, turned forward sharply, the aperture 
round or slightly elliptical- when the ooeciostome is fully developed this 
is the most striking character of the species. 

Described from Friday Harbor, Puget Sound. O'Donoghue has also 
recorded it from a number of localities in British Columbia at 10 to 
25 fms. 

Hancock Collections: Clayoquot Sound, British Columbia, at low 
tide, E. F. Ricketts, collector. 

Grisia denticulata (Lamarck), 1853 

Hincks (1884:203) reported this species from Houston-Stewart Chan- 
nel, British Columbia, without description. As it has not been noticed 
since on the Pacific coast it seems probable that he had another species, 
possibly C. serrulata Osburn, which is common in that area and which 
has some of the characters of denticulata. 

Grisia calif ornica d'Orbigny, 1853 

Crista calif ornica d'Orbigny, 1853:599. 
Crisia calif ornica, Busk, 1875:8. 

What this species from "Basse-Californie" may be is altogether prob- 
lematical, as d'Orbigny's description is so indefinite as to be useless and 
is without illustration. Busk merely translates d'Orbigny's description 
and questions whether it may refer to C. denticulataj which is not at all 
likely. The name should be dropped. 

Grisia punctata d'Orbigny, 1853 

Crisia punctata d'Orbigny, 1853:600. 

This species is also entirely unrecognizable from the very short de- 
scription and lack of figures. It was recorded from the Gulf of Cali- 
fornia "He du Venado, mer Vermeille, en Californie." The name should 
be dropped. 

Genus GRISULIPORA Robertson, 1910 

"Zoarium erect, dendroid, composed of internodes united by chitinous 
joints. Zooecia tubular, disposed in several alternate rows. Ooecium 
an inflation of the surface of an internode." (Robertson, 1910:254). 
Genotype, Crisulipora occidentalis Robertson. 



686 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

In addition it should be stated that, similar to Crisia, there are jointed 
rhizoids or radicles consisting of tubular kenozooecia ; the primary disc 
is separated from the primary zoid by a joint; the primary zoid arises 
from the top of the primary disc and not from its side ; the lower inter- 
nodes of the colony and its branches are uniserial or biserial; and the 
rhizoids, which are exactly like those of Crisia, often give rise to 
branches. For these reasons I agree with Borg (1926:475-6) in placing 
Crisulipora in the family Crisiidae. 

On the other hand, the ovicell usually resembles that of the Diaperoe- 
ciidae where Canu and Bassler (1920:749) have placed the genus. The 
gonozoids are more or less embedded between the autozoids and some- 
times they are expanded and surround a few peristomes. In narrower 
branches, however, they are simple, as those of Crisia, the only difference 
being that they are more embedded between the neighboring tubules. 

Crisulipora occidentalis Robertson, 1910 
Plate 72, fig. 6 

Crisulipora occidentalis Robertson, 1910:254. 
Crisulipora occidentalis, Okada, 1917:342. 
Crisulipora occidentalis, Marcus, 1937:21. 

The zoaria form large, stiff, often tangled masses, to a height of 30 
mm, attached by jointed radicles; additional zoaria are often produced 
from creeping radicles. Internodes long, separated by chitinous joints, 
the terminal ones gently curved backward, the more proximal ones 
shorter and straight. In the proximal internodes the zooecia number 
from 1 to 3 or 5, but terminal ones may have 40 or more. The zooecia 
are not symmetrically arranged as they are in Crisia, but are irregularly 
distributed. Cross sections of fertile internodes may show as few as 4 
or 5 zooecia, or as many as 8 to 10 at the widest part. Branching is like 
that of Crisia, with a more or less exserted basis rami. 

The zooecial tubes are slender and elongate, the frontal surface 
rounded in cross-section and the separating grooves quite distinct. The 
peristomes are moderately long, gently curved forward, a little narrower 
than the tubules, the aperture round and about 0.12 mm in diameter; 
in the basal internodes the tubules often show a rather regular alternate 
arrangement. 

The ovicells are elongate, narrowly wedge-shaped proximally and 
widening gradually upward, with much variation in size and form ; and 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 687 

there may be as many as 3 to an internode. In the narrower internodes 
they may be as simple as those of Crista (though more embedded), with 
a terminal ooeciostome which is smaller than a peristome and may 
terminate simply, or the rim may be expanded and slightly bell-shaped, 
the pore round. On broader internodes the ovicells are more expanded 
laterally and sometimes extend beyond the ooeciostome, and occasionally 
a few of the neighboring peristomes are surrounded. 

". . . at low tide almost anywhere on the coast of Southern California. 
... It has been dredged ofif the coast, from San Pedro to San Diego in 
depths ranging from 2 to 17 fathoms." (Robertson). Okada has re- 
corded the species from the Bay of Sagami, Japan, and Marcus reports 
it from Santos Bay, Brazil. 

Hancock Stations: Dredged at 28 stations, all the way from Point 
Conception, California to Peru. Station 844-38, Lobos de Afuera 
Islands, Peru, 6°55'40"S, 80°43'50"W, shore to 30 fms, the most 
southerly record; 31-33, Hood Island, Galapagos, 1°22'52''S, 89°39' 
15"W, at 4 fms; 308, Bahia Honda, Panama; Clarion Island, west of 
Mexico ; 7 stations in the Gulf of California ; Dewey Channel west of 
Lower California; and abundant about the Channel Islands off southern 
California as well as along shore; from low tide mark to a depth of 
47 fms. 

Division 3. Gancellata Gregory, 1896 

(Pachystega Borg, 1926) 

The Horneras etc. 

The primary zoid is erect but not separated from the ancestral disc 
by a joint; the zoarium is not jointed, erect, usually branched like a 
tree. The "wall of zoarium double, consisting of a gymnocyst and cryp- 
tocyst, the latter undergoing a process of secondary calcification, by 
which the zoarium in its older parts becomes very strongly calcified." 
(Borg, 1944:175). The ovicell or brood-chamber is a much expanded 
gonozoid, usually situated on the dorsal side or between two branches. 

Borg (1944:179) included the families Horneridae and Crisinidae, 
and erected three new families, Steghorneridae, Pseudidmoneidae and 
Calvetiidae, but did not discuss the Cytisidae. 

The only families we have to deal with are the Horneridae and 
Cytisidae. 



688 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Family Horneridae Smitt, 1867 

The zoarium is erect, branching like a tree, with rounded stems and 
branches ; the zoids opening on the frontal side only ; the inflation of the 
gonozoid or ovicell on the dorsal side of the zoarium. The family has 
not hitherto been recorded from the Pacific coast of America. 

Genus HORNER A Lamouroux, 1821 

With the characters of the family. Genotype, Hornera frondiculata 
Lamouroux, 1821. The zoarium with a moderate encrusting base, an 
erect round stem which branches like a tree with the successive branches 
diminishing in diameter, and the zooecia all opening on one side, easily 
distinguish the genus. The species are usually highly colored red or 
purple. 



Hornera pectinata Busk, 1861 
Plate 72, figs. 10, 11, and 12 



Hornera pectinata Busk, 1861:79; 1875:18. 
Hornera pectinata, Johnson, 1897:61. 
Hornera pectinata, Norman, 1909:280. 

The zoarium is erect, flabellate, the short main stem rising from a 
slightly expanded base; height 25 mm; the base measures 5 by 7 mm, 
the main stem 3 mm, the larger branches about 2 mm, the terminal 
branches just below the tips 0.50 to 0.60 mm. The branching is in one 
plane, irregular with a tendency toward dichotomy; the main branches 
are rather regularly tapered from base to tip ; all of the branches are 
round even in the youngest stages. Stunted branches rare. Apparently 
purple when living. 

The zooecia are irregularly arranged in more or less transverse rows 
of 2 to 4 tubules, connate or separated and often single. All of the 
peristomes are short, but the outer ones are slightly longer. On older 
branches the apertures are nearly level with the surface, the apertures 
round, 0.20 to 0.24 mm in diameter, the rim of the peristome thin, often 
slightly flared and delicately serrate (never incised), the points being 
the tips of the parallel ridges or thickenings of the peristome. The longest 
marginal peristomes are seldom as much as 0.20 mm in height. The 
sulci are strongly developed on both frontal and dorsal sides and the 
pores are round or slightly elongate. The ridges between the sulci are 
very irregular on the frontal surface, but are continuous and more or 
less parallel on the dorsal side. Complete calcification of the zoarium 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 689 

extends only about one third of the distance above the base and the 
peristomes protrude above the level of the front on the v\^hole upper two 
thirds of the zoarium. Ovicells are not developed on any of our speci- 
mens, and in their absence positive identification is impossible. Hov^^- 
ever, the zoarial characters given by Johnson (1897:61) in his amplifi- 
cation of Busk's description of pectinata apply very well to our speci- 
mens: 

"branches terete . . . ultimate branches tapering . . . Anterior sur- 
face pierced by numerous oval pores, which are sunk in depressions and 
have slightly raised borders. Between the pores the surface is irregularly 
ridged. The pores on the dorsal surface are larger and are partially 
filled up inside. The peristome is minutely dentate." 

Johnson also describes the ovicell, "dorsal, brownish, semiglobular, 
and the surface is thickly set with warts, each of which has a depression 
at the top with a perforation therein." 

The species has been recorded only from the Madeira Islands. 

Hancock Stations: 1397-41, Santa Rosa Island, 33°38'40"N, 119° 
58'30"W, at 77 fms; 1299, off Point Firmin, 33°4r45''N, 118°17'50" 
W, at 18 fms; and off Santa Catalina Island, 33°24a5''N, 118°13'30" 
W, at 228 fms ; all from southern California. 

Hornera pinnata Canu and Bassler, 1929 
Plate 72, figs. 7, 8, and 9 

Hornera pinnata Canu and Bassler, 1929:550. 

The zoarium is erect from a very small base, the branching dicho- 
tomous and irregular, the branches with short pinnules of various sizes. 
There are usually two rows of peristomes on each side of the midline, 
sometimes only one row, irregularly alternating, the outer ones the 
longer. The frontal surface is deeply grooved, the pores conspicuous and 
3 to 5 in number on each tubule. The peristomes measure about 0.12 
mm in diameter and the aperture about 0.10 mm. The pinnules (dwarfed 
branches) vary greatly in size and the number of their tubules varies 
from 3 to 8 ; sometimes a pinnule ends in a blunt point with a peristome 
medially placed at the tip. The dorsal side of the zoarium is deeply 
grooved longitudinally, with conspicuous pores at the bottom of the 
grooves. 

Ovicells are wanting on all of our specimens, so it is impossible to be 
absolutely certain of the species, but the zoarial characters seem to agree 
closely with the description of pinnata. Also I have for comparison an 
ovicelled specimen from Hawaii which is undoubtedly H. pinnata. The 



690 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

diameter of the peristomes is only slightly larger, the arrangement and 
number of the tubules and pores is the same, and the nature of the pin- 
nules corresponds. 

It was described from the Philippines, the China Sea and Borneo. 

Hancock Stations: 1323-41, off Santa Catalina Island, southern Cali- 
fornia, 33°14'40''N, 118°12'15''W, at 152 fms; 2158, Ranger Bank 
off Cedros Island, west of Lower California, 81 fms; and 299, San 
Jose del Cabo, near the southern tip of Lower California, 22°56'15"N, 
109°47'15"W, at 83 fms. 

Family Cytisidae d'Orbigny, 1854 
Genus DISGOGYTIS d'Orbigny, 1854 

D'Orbigny (1854:1061) gave an unusually careful and full descrip- 
tion of this genus, in which he mentioned the attached base upon which 
rises a narrow peduncle expanding upward into a cupuliform head, the 
whole zoarium shaped like a wine-glass ; the upper surface very concave 
or infundibuliform at the center and the margin with numerous simple 
or branched fascicles. He also observed and figured (Plate 798, fig. 8) 
the unusual position of the ovicell on the under side of the cup above 
the peduncle. Genotype, Pelagia eudesii Michelin, 1844:123. 

This genus has been known only as a fossil from the Cretaceous until 
O'Donoghue (1926:26) described D. canadensis as a recent form from 
British Columbia. In the Hancock Collections there are several zoaria 
which are similar and which fill all the requirements of Discocytis. 

Discocytis californica new species 
Plate 69, fig. 11 

The zoarium is attached by a round thin disc, from the center of 
which rises a comparatively thin cylindrical peduncle ; at the upper end 
this widens gradually into a funnel-shaped head or capitulum like an 
inverted cone, the whole structure resembling a minute and moderately 
short-stemmed wine glass. On its upper surface the capitulum is con- 
cave and the whole central area is occupied by rather large cancelli with 
thick walls and rounded apertures. Around the rim of the cup the func- 
tional tubules are arranged in short fascicles, 8 to 10 in number, com- 
pletely connate, the apertures measuring about 0.10 mm in diameter. 
The base and stem show no open tubules and appear as if covered by 
a thin pellicle. The measurements of the various zoarial parts are, on 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 691 

our largest specimen: height 1.75 mm, width of base 1.20 by 1.30 mm, 
width of stem 0.53 mm, length of stem 0.60 mm, height of capitulum 
1.15 mm, width of capitulum 1.57 by 1.70 mm, height of fascicles 
0.25 mm. 

The ovicell is very large, conspicuous, rounded and bulbous and is 
situated on the under (dorsal) side of the capitulum close to the base 
of the fascicles, its width 0.55 by 0.65 mm. A portion of the wall is 
broken away and the ooeciostome is wanting. On another, somewhat 
smaller, specimen there is a smaller ovicell of similar appearance, and 
it also has been injured. 

At first I presumed this to be D. canadensis O'Donoghue, 1926:26, 
but it is much smaller, the ovicell is strikingly different in form, and 
O'Donoghue describes the capitulum as broad and flattened. 

Type, AHF no. 124. 

Type locality, off Rocky Point, southern California, about 33°49'N, 
encrusting on a sunken buoy at a depth of 45 fms, three colonies. 

Discocytis canadensis O'Donoghue, 1926 

Super cytis digitata, O'Donoghue, 1923:16. 
Discocytis canadensis O'Donoghue, 1926:26. 

This species has not been found in the Hancock Collections, but the 
following digest of O'Donoghue's description is here given to indicate 
the differences. Zoarium cupuliform; base small, flat and circular; stalk 
short, narrow, expanding into a broad, flattened funnel-shaped capitulum, 
from the edge of which a number of pinnules radiate outward; each 
pinnule (fascicle) consists of 12 to 20 tubes closely connate. Largest 
specimen 4 mm high, the stalk 1.75 mm thick, and the capitulum 7.25 
mm across. 

Ooecium transversely elongate, sinuous, running up slightly between 
the bases of the fascicles, its breadth one fifth to one fourth of the cir- 
cumference. Ooeciostome a circular aperture surrounded by a flattened 
ring-like margin, sub-terminal near the middle of the ooecium. 

Recorded by O'Donoghue from a number of localities on the British 
Columbia coast and south to the San Juan Islands in Puget Sound. 



692 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Division 4. GeriopoHna Hagenow, 1851 

(Heteroporina Borg, 1933) 

The Heteropores 

"Primary zoid adnate or partially erect; zoarium varying in shape, 
adnate, suberect or erect, composed of zoids of two kinds, autozoids and 
kenozoids, the latter at least as numerous as the former, both autozoids 
and kenozoids opening at about right angles to the surface of the 
zoarium ; wall of the zoarium double, consisting of gymnocyst and 
cryptocyst; brood chamber a coelomic space, formed by the absorption 
of the subdistal portions of some autozoids and numerous kenozoids 
outside the fertile, ovigerous zoid (zoarial brood-chamber)." (Borg, 
1944:208). 

Family Heteroporidae Waters, 1880 

"Zoarium erect, pedunculate and capitate, or arborescent ; autozoids 
and kenozoids about equally numerous or the former less in number; 
apertures of both kinds of zoids scattered over the surface of the zoarium, 
not forming clusters, circular or polygonal in shape; brood chamber 
zoarial, not visible from surface except in form of a slight swelling of 
that part of the zoarium." (Borg, 1944:209). 

Two genera are represented in our material, Heteropora Blainville, 
1830, and Borgiola Strand, 1933 (Canuella Borg, 1933, preoccupied), 
the latter having the autozoids often forming small clusters instead of 
being more regularly distributed. One species of this genus forms a 
heavy incrustation without any erect branches. 

Genus HETEROPORA Blainville, 1830 

"The zoarium is erect, arborescent, its surface smooth or slightly 
rugose, honeycomb-like when the cystids are open ; the kenozoids much 
more numerous than the autozoids, located between them and thus 
separating them, aperture circular or polygonal." (Borg, 1944:210). 
Genotype, Ceriopora cryptopora Goldfuss, 1827. 

Robertson (1910:258) gave a very clear statement of certain zoarial 
details: "If one examines the growing tips of a branch, the tubular open- 
ings found there are for the most part those of zooecia in various stages 
of maturity. Between them, formed by minute triangular spaces where 
the walls of zooecia do not come into contact, are the interstitial spaces 
(kenozooecia). As growth proceeds, both zooecia and interstitial canals 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 693 

curve outwards, and although at the growing tip these tubes are parallel 
to the axis of the branch, when adult they curve almost at right angles 
with the axis of the branch and the apertures open laterally, the larger 
zooecial apertures being surrounded with a circle of small interstitial 
openings." 

Borg (1932, 1933 and 1944) has made a very detailed study of the 
anatomy, development and the brood-chambers of recent species of 
Heteropora and related genera. He rejects the genus Tretocycloecia 
Canu, 1918a :346, erected to include the species with ovicells and leaving 
the old genus Heteropora for those in which the ovicells are unknown, 
as "inadmissible." 

Heteropora magna O'Donoghue, 1923 
Plate 73, fig. 13 

Heteropora magna O'Donoghue, 1923:14. 
Tretocycloecia magna, O'Donoghue, 1926:29. 
Heteropora pelliculata, Robertson, 1910:258 (part). 
Heteropora magna, Borg, 1933:326. 
Tretocycloecia pelliculata, Canu and Bassler, 1922:110. 

"Zoarium stout, densely branched, more or less spherical in outline; 
distal ends of autozoids not or only slightly protruding; apertures of 
kenozoids mostly open, but sometimes, in older portions of the zoarium, 
closed." (From Borg's key, 1933:284). 

The encrusting base gives rise to erect cylindrical stems, 3 to 5 mm in 
diameter, which branch dichotomously while retaining their original 
thickness, but a little swollen at the tips. The zoarium thus has a more 
or less spherical form, except when modified by the substratum. There 
is occasional anastomosis of the branches, "but not so frequently as in 
H. pelliculata (now H. pacifica Borg, q.v.) and the colony as a whole 
has a much more stout and compact appearance . . . and may measure 
100 by 70 mm." (After O'Donoghue). Zoarium purplish-brown in 
color. 

Ovicells were not observed by either O'Donoghue or Borg. They are 
large irregular areas on the sides of the branches near the tips and par- 
tially surrounding the branches, only slightly elevated, but conspicuous 
because of the closure of the kenozoids ; appearing as a complete, calcified, 
thin, whitish cover of the brood-chamber, with the exception of the 
peristomes of the autozoids which are not displaced. The peristomes 
are more prominent than elsewhere on the zoarium and are often slightly 



694 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

flared at the tips and thus appear somewhat larger than the ordinary 
ones. I have not been able to find any aperture on the four brood- 
chambers in my material which I can positively identify with ooeciopores, 
and it is therefore probable that they are similar to the ordinary aper- 
tures. On the removal of the roof of the brood-chamber an extensive, 
broad cavity is revealed, with the bases of the absorbed kenozoids at the 
bottom. The peristomes of the autozoids usually traverse the cavity 
without modification, but I have found a few which have become closed, 
within the chamber, by a membrane with a central raised pore similar 
to those of Diastopora. The apertures of the autozoids average about 
0.18 mm in diameter. 

This species did not appear in the Hancock dredgings, but I have a 
fine specimen from Friday Harbor, Puget Sound, 70 mm long by 45 
mm wide and 40 mm high, without further data, but evidently dredged 
locally. The encrusting base is 7 by 9 mm across, and there is a secondary 
attachment of similar appearance by a branch, 5 mm across. Another 
portion of a colony, loaned by Dr. R. E. Foerster, Director of the Pacific 
Biological Station, Nanaimo, British Columbia, and bearing O'Donog- 
hue's identification, is from Gabriola Pass, B. C. This specimen also has 
a brood-chamber. 

Heteropora pacifica Borg, 1933 

Heteropora pacifica Borg, 1933:317. 
1 Heteropora sp. Whiteaves, 1882:279. 
Heteropora pelliculata, Robertson, 1910:258 (part). 
Heteropora pelliculata, O'Donoghue, 1923:14 (part). 
Tretocycloecia pelliculata, O'Donoghue, 1925:96; 1926:28 (part). 

(The synonymy according to Borg). 

Borg separated the more slender, more intricately branched and more 
highly anastomosed form mentioned by Robertson and O'Donoghue as 
a distinct species. As I have not had an opportunity to study H. pacifica, 
I can only indicate the essential points of difference given in Borg's 
description. 

The zoarium is erect, branching profusely and dichotomously or in 
an irregular way. The branches frequently anastomose, giving the whole 
colony a complexly reticulated appearance which is highly characteristic 
of the species. The diameter of old stems is about 5 mm, of younger 
ones 3 mm on an average. Color of dried zoaria grayish, the tips pink. 
Autozoids with the apertures usually on a level with the surface, but 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 695 

in more sheltered areas the peristomes are very evident. The apertures 
are about 0.17 mm in diameter, in older parts of the colony they are 
usually closed. The kenozoids never project above the surface and are 
usually closed except tow^ard the growing ends of the branches. Brood- 
chamber not knov^^n. 

While this species may have a considerable range along the Pacific 
coast from Alaska to California, as suggested by O'Donoghue, the only 
positive records are those of the material studied by Borg, Vancouver 
Island region to Middleton Island, southern Alaska, down to a depth 
of 25 meters. 

There is in the Hancock collection a very small fragment labelled 
Heteropora pelliculata Waters by Robertson, from "San Diego, Cali- 
fornica," which may belong to H. pacifica as the zoids have the same 
measurement, 0.17 mm. There are also several much worn fragments 
from Hancock Station 1278-41, off San Miguel Island, southern Cali- 
fornia, at 35 fms, which present the same zooecial measurements and 
with stems 2 to 3 mm in diameter, which may belong here. But none 
of these specimens is in condition for determination beyond the genus. 

There are also several much worn fragments from Hancock Stations 
143-34, Wenman Island, and 170-34, Chatham Island, Galapagos 
Islands, which present about the same measurements, but which are not 
identifiable beyond the genus Heteropora. 

Heteropora alaskensis (Borg), 1933 

Plate 73, figs. 10, 11, and 12 

Heteropora pacifica var. alaskensis Borg, 1933:325. 
Heteropora pelliculata, Robertson, 1910:258 (part). 
"i Heteropora pelliculata, O'Donoghue, 1923:14 (part). 

The zoarium is very irregular and much smaller than that of H. 
magna or H. pacifica. Our largest colony measures 16 mm in height by 
25 mm in width, with 16 branches which average about 2 mm in 
diameter, but most of the 10 colonies are shorter and more compact; 
the branches, beyond a bifurcation, are 2 to 4 mm long (in one case 
7 mm) ; only a few cases of anastomosis occur. 

The essential characters which differentiate this form from other 
species of Heteropora are: (1) the peristomes of the autozoids project 
to a marked degree on the branches all the way up to the margins of 
the cancellated tops, and on older basal branches they still rise slightly 
above the level of the zoarial surface; diameter of apertures 0.14 (0.13 
to 0.17) mm; (2) the kenozoids are covered and closed over the whole 



696 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

surface of the colony up to the margin of the cancellated tops of the 
branches. In these characters our specimens agree with Borg's descrip- 
tion of H. pacifica var. alaskensis, and the differences are so striking and 
constant as to warrant the elevation of the "variety" to specific standing. 

On protected areas the peristomes may be as much as 0.25 to 0.30 
mm long, though usually they are much shorter. The tips of the branches 
are sometimes evenly rounded, but more frequently they are somewhat 
clavate or spatulate, slightly excavated on one side where a brood- 
chamber is located, and the rim may extend beyond and give off two 
or three branches, thus leaving the brood-chamber in the broad fork of 
a branch. Other brood-chambers are found on the sides of branches, 
as shown in Borg's figure 5 (plate 10). 

The brood-chamber is typically that of Heteropora, a low, more or 
less flat swelling, through the roof of which the peristomes of the 
autozoids penetrate and are slightly elevated above it. On the removal 
of the calcified membrane, or roof of the chamber, a considerable cavity 
is exposed, traversed by the autozoid tubules and showing the remains 
of the partially absorbed kenozoids. I have not been able to find on any 
of our specimens the large ooeciostome figured by Borg and cannot de- 
termine the location and form of the ooeciopore. 

This species differs from H. pelliculata Waters, 1879, which it some- 
what resembles, by the elevation of the peristomes over the whole surface 
and by the complete closure of the kenozoids over the whole colony up 
to the level of the margins of the cancellated tops of the branches. The 
same characters, as well as the smaller size, distinguish it from H. 
pacifica Borg. As the color of our preserved specimens is white, this may 
be the lemon-colored form recorded by Robertson (1910:259) from 
"Channel Rock, Puget Sound." 

Our specimens are from Bentinck Islands, British Columbia, without 
further data, loaned by Dr. W. A. Clemens of the University of British 
Columbia, ten colonies of various forms and sizes. Also 2 fragments, 
with ovicells, from Clayoquot Wharf, British Columbia, E. F. Ricketts, 
collector. 

Hancock Station 1490-42, off Cape Arago lighthouse, Oregon, 43° 
20'26''N, 124°22'24"W, at very low tide, 5 fragments. 

Genus BORGIOLA Strand, 1933 

Canuella Borg, 1933:331, (preoccupied by Scott, 1893). 

"Zoarium erect, arborescent, branching sparsely; its surface strongly 
rugose, showing numerous irregularly shaped elevated areas and between 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 697 

them well-marked depressions; autozoids frequently forming small 
clusters; their apertures oblique, prolonged at one side into an erect 
pointed process; kenozoids much more numerous than the autozoids, 
thick-walled, mostly with open apertures," Borg, 1933:331. Genotype 
Canuella rugosa Borg, 1933 :332. 

The genus is certainly close to Heteropora and possibly may be found 
to intergrade. However any means of separating the members of this 
family, and especially if it is found to apply to the very numerous fossil 
species, is welcome. 

The generic description requires modification to include an encrusting 
species, without erect branches, B. pustulosa new species, which is de- 
scribed below. 

Borgiola rugosa (Borg), 1933 
Plate 76, fig. 11 

Canuella rugosa Borg, 1933:331. 

The zoarium of our single specimen measures 25 mm in height and 
20 to 25 mm in width, rising from a broad encrusting base 20 by 10 mm 
across, with numerous irregular branches which are sometimes bifurcate ; 
and there is a single anastomosis. Several small subcolonies arise from 
the lateral extension of the base. The main stem is about 5 mm in 
diameter, the branches becoming progressively smaller until the terminal 
ones measure 2 mm or less. Color pure white. 

The autozoids, or functional zooecia, tend to occur in small groups 
or clusters, sometimes in radiating lines, though single autozoids are 
also common. The peristomes usually rise slightly above the surface, 
with the rim higher on one side and often extending into a sharp point ; 
or the rim may be evenly rounded. The zooecial tubes are long, those in 
the rounded branches have their origin in the center and curve outward 
as in Heteropora; those which form the basal expansion arise on the 
lamina and curve upward. There is a moderately broad basal lamina 
surrounding the basal expansion. The apertures of the autozoids are 
about 0.13 to 0.15 mm in diameter. The kenozoids are much more 
numerous than the autozoids, always noticeably smaller but varying in 
size, never rising above the surface and seldom entirely closed. The 
brood-chamber has not been observed. 

A striking feature of the zoarium is the peculiar type of rugosity 
produced by the irregular elevation of areas with increased numbers of 
autozoids, while between these are smooth areas of lower level in which 
there are fewer autozoids. 



698 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

I presume this to be the same species as that from Japan described by 
Borg, since the zoarial and zooecial characters appear to agree closely, 
but in the absence of brood-chambers the identification is necessarily 
tentative. 

The species has hitherto been known only from Sagami Bay, Japan. 

Hancock Station, 310-35, off Bindloe Island, Galapagos, 0°18'20"N, 
90°3riO"W, at 15 fms, rocky bottom. 

Borgiola pustulosa new species 
Plate 73, figs. 5, 6, 7, 8, and 9 

Zoarium encrusting on rocks and shells, with no evidence of erect 
branches, the surface with numerous low rounded or elliptical elevations 
which are rather evenly spaced, the elevated areas about as wide as 
the lower areas between them. The largest colony (type) measures about 
70 mm long by 50 mm wide by 10 mm thick, but is broken and was 
evidently considerably larger. Another fragment is 50 mm in width, and 
a third fragment which is on a shell, is 25 mm long by 10 mm wide and 
appears to have been about twice that width. The color is white to 
yellowish red. 

On the lower, general, surface the apertures of the autozoids are 
often quite regularly spaced and surrounded by a single row of keno- 
zoids about half as large as the autozoids ; on the pustules the autozoids 
are irregularly disposed and the kenozoids more numerous and irregular 
in size, and occasionally there are small areas consisting entirely of 
kenozoids. Over most of the surface the tubes of the autozoids project 
very slightly above the level of the surrounding kenozoids, the rim round 
or nearly so ; but around the borders of the pustules they are noticeably 
higher and produced on one side into a pointed process, "giving the 
aperture a distinctly oblique appearance," as Borg describes them in 
C. rugosa (1933:335). On older areas both autozoids and kenozoids 
are frequently closed, slightly below the level of the rim. The apertures 
of the autozoids are about 0.18 mm in diameter but vary considerably. 
The kenozoids vary excessively, from 0.03 to 0.12 in diameter, but 
average about 0.08 mm, and they also vary in form. 

The autozoids arise on a basal lamina which extends rather narrowly 
around the margin of the zoarium, at first prone but curving upward 
at once into an erect position. 

The brood-chambers are spacious cavities, as much as 2.50 mm in 
width and 0.50 mm in depth, resembling those of Heteropora with the 
kenozoid walls absorbed and closed on the floor of the cavity, and most 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 699 

of the autozoid tubes continued through the chamber and on above it. 
In the broken walls of old thick zoaria the chambers appear at different 
levels, some just beneath the surface, but others buried deeply by the 
regeneration of the zoarium above them. There is little or no surface 
evidence of the position of the brood-chambers, and the pustules appear 
to have no relation to them, as shown by dissection. Borg could not be 
certain as to the brood-chamber in his material of B. rugosa, but that 
of his figure 2 (plate 3) appears to be similar to those of pustulosa. I 
have observed no differentiation of ooeciostomes. 

Borgiola is certainly close to Heteropora in most of its characters, 
but the roughened nature of the zoarial surface, the grouping of the 
autozoids and heterozoids and the pointed processes of the taller zooecial 
tubes separate it. 

The occurrence of this species is of unusual interest, since no member 
of the Family Heteroporidae has hitherto been recorded from the Arctic 
Ocean. 

Type, U. S. Nat. Mus. no. 11051. 

Type locality, Point Barrow, Alaska, Arctic Research Laboratory, 
453 feet, encrusting a stone, G. E. MacGinitie, collector. Also on a stone 
from 295 feet and on a shell at 60 feet. 

Division 5. Rectangulata Waters, 1887 

(Calyptrostega Borg, 1926) 

The Lichenopores 

"They fool me to the top of my bent." Shakespeare. 

"Primary zoid adherent to the substratum, never separated by any 
joint from the pro-ancestrula ; zoarium wart-like, its basal wall adnate, 
simple; frontal wall double consisting of gymnocyst and cryptocyst; 
between zoids special coelomic cavities (alveoli) limited by calcareous 
extrazoidal walls; no vestibular sphincter; brood-chamber zoarial, a 
coelomic space corresponding to numerous alveoli, outside the fertile 
zoid • polypide degenerating first after having been functional for some 
time." (Borg, 1944:211). 

The Lichenopores have always been a "thorn in the flesh" to those 
who have attempted to work with them. Defrance established the genus 
Lichenopora in 1823. Before this time the species were usually re- 
ferred to Madrepora (a coral), or to Tubulipora Lamarck. Since then 
a large number of generic names have been proposed ; d'Orbigny was 
especially lavish in this respect, separating out ten "genera" on trivial 



700 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

zoarial characters. Some of these have proved to belong in other families 
and others placed in synonymy until only two genera, Lichenopora 
Def ranee, 1823, and Disporella Gray, 1848, have survived the research 
of later authors. Most recent authors have used only Lichenopora, but 
Borg, 1944:234-5 and 249, has given what appear to be good reasons 
for the retention of Disporella. Borg even goes so far as to propose a 
new family, Disporellidae, to include only this genus, but this appears 
to me to be unwarranted on the basis of the characters, 

A few citations will indicate the difficulty others have had : "One 
cannot help feeling despair when trying to determine the Lichenoporae." 
(Waters, 1889:282) ; "The determination of the species of Lichenopora 
is admittedly very difficult." (Harmer, 1915:160) ; "The determination 
of the species, even the recent ones, presents much difficulty." (Canu 
and Bassler, 1920:812) ; "The discrimination of the species . . . has 
scarcely but begun." (Borg, 1944:213). 

Family Lichenoporidae Smitt, 1866 

Zoaria rounded or ovate, occasionally otherwise modified in outline 
by the nature of the substratum ; more or less convex, sometimes dome- 
shaped ; attached the full width of the basal lamina, or the basal lamina 
free and turned upward at the edge, or, when on small stems, they may 
be attached by a short central stipe of variable width. The central part 
of the zoarium, varying in size with the species, is occupied by cancelli 
(alveoli, Borg), and outside of this area the functional zooecia are 
arranged in radiating series or more or less in quincunx. The "peri- 
stomes" are usually much higher next to the central area and decrease 
in height regularly to the margin ; usually the basal lamina extends in 
a thin rim around the outer edge. Zoarial budding occurs in some 
species, either vertically or near the edge, and sometimes very complex 
zoaria may be formed in this manner. The ovicells are brood-chambers 
of considerable size, occupying the central area, branching out in lobes 
between the rays, or located entirely between the rays ; covered by a 
thin calcified layer with minute pores, which may be obscured by the 
development of secondary cancelli above it. 

As stated above, Borg has indicated two families on the following 
basis : 

Lichenoporidae, alveoli soon "roofed in" by a porous calcified layer, 
with secondary alveoli above them ; the one central brood-chamber, 
which may have lobes extending between the rays; zoarial budding 
vertical. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 701 

Disporellidae, alveoli not roofed in but partially closed "iris-like" 
with a round hole at the center; brood-chambers between the rays; 
zoarial budding lateral. Z?-?) 

This separation seems to work very well with the few species/ treated 
by Borg, but, unfortunately for taxonomic purposes, the three characters 
used by him to distinguish Lichenopora and Disporella do not appear 
to be constant throughout the series. Thus in Lichenopora huskiana, 
novae-zelandiae and intricata, which have irregular cancelli closed by a 
membrane, the budding is lateral with the sub-colonies beyond the margin 
of the primary disc. The location of the ovicells, in the central area or 
interradial, shows frequent variations. Also the closure of the cancelli, 
by a thin porous membrane or by an iris-like diaphragm, is not constant, 
as both types may occur on the same zoarium. In L. mtricata Busk, the 
interradial cancelli and those of the central area of infertile discs have 
the rounded cancelli, while the fertile discs present the irregular cancelli 
above the ovicells; in several of the elongated discs of this species both 
kinds of cancelli are present, the irregular ones covering the ovicell at 
one end and the rounded cancelli at the opposite infertile end. 

Apparently the only character that seems to hold absolutely is the 
presence of irregular, thin-walled secondary cancelli covering the ovicells 
in Lichenopora. 

Genus LICHENOPORA DeFrance, 1823 

Brood-chambers, one or more, occupy the central area and may extend 
somewhat into the interradial areas; in older zoaria secondary small 
brood-chambers may appear between the rays toward the margin. Can- 
celli (alveoli) of the primary layer thin-walled, irregular in form and 
size, and closed by a thin, perforated, calcified layer; secondary cancelli 
above these may be similar to these or may be thicker-walled with large 
rounded apertures. Marginal zoarial budding sometimes occurs but 
vertical budding is the rule. The distribution of the functional zoids 
varies much among the species, in short or longer connate or non-connate 
rays or in irregular quincunx. Genotype, L. turbinata Def ranee, 1823: 
257. 

Key to the Species of Lichenopora 

1. Radiating rows of tubules biserial buskiana 

Rays uniserial or more or less in quincunx 2 



702 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

2. Ooeciostome hooded over the top and opening on the side; 

tubules distinct, high on proximal border and with one to 

several sharp points canaliciilata 

Ooeciostome wide open 3 

3. Zooecial tubules not connate and, except near the center, they 

are usually scattered or in quincunx verrucaria 

Zooecial tubules in series only, connate to the tips 4 

4. Cancelli distinctly larger than the zooecial apertures; radii in 

regular series ; pin-head spicules abundant . . novae-zelandiae 
Cancelli smaller; radii regular only close to the central area; 
very complex and intricate colonies formed by lateral bud- 
ding; pin-head spicules almost wanting intricata 

Lichenopora canaliculata (Busk), 1876 
Plate 76, figs. 3 and 4 

Discoporella canaliculata Busk, 1876:118; 1879:199. 
Lichenopora grignonensis, Ridley, 1881:57. 
Lichenopora fimbriata, Borg, 1926:184. 
Lichenopora canaliculata, Borg, 1944:235. 

Busk's description is very brief, "Zoarium circular, bordered, slightly 
convex; tubes very irregularly uniserial, with a raised canalicular fillet 
on one side, interspaces cancellous." Borg, 1944:235, had very rich 
material from the Swedish Antarctic Expedition and has given an ex- 
tended discussion of the species. 

The zoaria are circular, somewhat elevated at the center and sloping 
regularly to the basal lamina which is broad and thin; encrusting on 
shells and attached to stems. The basal lamina is traversed nearly to 
the edge by the bases of young zooecia. The central area of the zoarium 
is comparatively small, a little depressed in the young, but later filled 
in by the flat-topped brood-chamber. The zooecial tubes are irregularly 
distributed or in short radiating lines and never connate; they are 
moderately high, usually much elevated on the proximal (central) side 
and low on the side toward the border, thus forming a channel above 
the aperture; the proximal side with one to several "fillets" or longi- 
tudinal ribs which end in points; aperture rounded and 0.10 to 1.12 
mm in diameter. The cancelli are large and irregular in form, often 
twice as large as the apertures of the zooecia, the walls thin and the 
whole giving a reticulated appearance. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 703 

The brood-chamber covers the central area, with a thin calcareous 
layer which is very minutely perforated ; a coarsely reticulated layer of 
secondary cancelli later may cover it. The ooeciostome appears to be 
unique in this genus; it has the short erect cylindrical base but the 
orifice is on the side, with a peculiar "helmet" or hood-shaped cover 
which arches widely over the top, closing the orifice entirely from above ; 
also just inside from the rim of the hood there is a transverse row of 
minute pores, as described by Ridley under L. grignonensis (=L. ca- 
naliculata) . 

This species has been recorded only for Antarctic and far southern 
waters ; Kerguelen Island ; Strait of Magellan ; Kap Adare, Victoria 
Land; and New South Wales. It is therefore of special interest to 
discover it in the Arctic region. The nature of the tubules and especially 
the form of the ooeciostome with its row of perforations seem sufficient 
for positive identification. 

Point Barrow, Alaska, Arctic Research Laboratory, 110 to 522 feet, 
several colonies, only one of which is mature, G. E. MacGinitie, col- 
lector. 

Lichenopora verrucaria (Fabricius), 1780 
Plate 74, fig. 3 

Madrepore verrucaria Fabricius, 1780:430. 
Discoporella verrucaria, Busk, 1875:31. 
Lichenopora verrucaria, Hincks, 1880:478; 1884:207. 
Lichenopora verrucaria, Robertson, 1900:329; 1910:263. 
Lichenopora verrucaria, O'Donoghue, 1923:15; 1926:28. 
Lichenopora verrucaria, Canu and Bassler, 1923:205. 

Zoaria small, rarely more than 3 mm in width; encrusting on algae, 
stems, stones and shells ; high near the center and rounding off gradually 
to the margin where there is a narrow basal lamina. In younger colonies 
there is a depressed cancellous central area, but in the sexually mature 
this area is filled in with one or more brood-chambers. The zooecia are 
irregularly arranged, often in short radiating series, especially near the 
central area, but never connate ; moderately elevated, carinated on the 
side toward the center, the keel rising into a point. The cancelli vary 
greatly in size, sometimes larger than the zooecial apertures but often 
not half as large. 

Usually only one brood chamber fills the central area, but as many 
as three have been observed, in which case they fuse externally so that 
the number of ooeciostomes is the only obvious clue to the number of 



704 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

chambers; they do not extend between the zooecial rays farther than 
the central ones. The ooecial cover is a thin inflated calcareous plate 
with numerous minute pores ; secondarily the ooecial roof may be cov- 
ered by a layer of cancelli which form a very irregular reticulum. The 
ooeciostome is more or less excentric in position, a short erect cylinder 
with a flaring lip which varies from nearly round to elliptical, the ooecio- 
pore 0.10 to 0.12 mm in diameter. 

A common northern and arctic species, extending south to Cape Cod 
on the Atlantic coast and to California on the Pacific coast; abundant 
in the Arctic seas. 

Hancock Station, 1416-41, San Miguel Island, southern California, 
34°02'45'"N, the most southern record. Hein Bank, Puget Sound, J. L. 
Mohr, collector; British Columbia (Hincks and O'Donoghue) ; south- 
ern Alaska (U. S. Crab Investigation) ; Bering Sea; and abundant at 
Point Barrow, Alaska, shallow water to 85 fms, G. E. MacGinitie, 
collector. 

Lichenopora buskiana Canu and Bassler, 1928 
Plate 74, figs. 1 and 2 

Lichenopora buskiana Canu and Bassler, 1928:164. 
Not Unicavea Calif ornica d'Orbigny, 1853:972. 
Lichenopora Californica, Conrad, 1855:441. 
Lichenopora californica, Gabb and Horn, 1862:176. 
Discoporella californica. Busk, 1875:32. 
Lichenopora californica, Canu and Bassler, 1923:203. 
Lichenopora buskiana, Borg, 1944:219 and 224. 

The misidentification of this species with d'Orbigny's Unicavea cali- 
fornica has led to much misunderstanding; Unicavea was described as 
having only uniserial radii, "Toutes les lignees n'ont qu'une seule ligne 
de cellules" (d'Orbigny, 1853:970). 

The zoaria are attached to algae especially, sometimes to shells, worm 
tubes, etc. The central frontal area is comparatively small and rounded, 
with large cancellae which are slightly larger than the zooecial apertures. 
The radiating rows of zooecial tubes are connate and biserial, except 
that they may begin with a single tube which sometimes is not connate, 
and also that rarely there may be three series near the outer ends of the 
radii. Near the center the tubules are unusually high and nearly erect. 
There are usually 8 to 12 primary radii, with shorter rays originating 
between these toward the margin. As a rule the rays are very regular 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 705 

in form, size and arrangement, and separated by 1 or 2 rows of cancellae, 
often with "pin-head" spicules. The apertures of the tubules are some- 
what irregular in form and about 0.06 to 0.08 mm in diameter. 

The brood-chamber (ovicell) in smaller zoaria occupies only the 
central area, with the ooeciostome a little excentric or frequently nearer 
the border of the area. The ooeciopore is about as large as the zooecial 
apertures ; the ooeciostome is erect with a short cylindrical stalk which 
flares, trumpet-shaped, slightly ovate, 0.13 to 0.16 mm by 0.16 to 0.20 
mm in breadth at the tip. The roof of the brood-chamber is thin and 
perforated by numerous small pores, and above this are secondary can- 
cellae of irregular size and form. Strong irregular raised lines often 
give the area a coarsely reticulated appearance. 

The zoaria are often complex, with daughter colonies budding off 
from the sides, and these are frequently very irregular, both in shape 
and in the arrangement of the uniserial rays. In the daughter colonies 
the brood-chambers are irregularly situated, often small and situated 
between the series of tubules, sometimes near the margin. Even in 
larger simple colonies there may be small secondary brood-chambers near 
the margin. 

The biserial rays with high peristomes, the large and irregular can- 
celli which frequently bear "pin-head" spicules within their apertures, 
and the closure of older cancelli by a thin calcified porous membrane, 
readily distinguish this species from any other of the Pacific coast, even 
in the absence of an ooeciostome. 

It is a common species along the shore and about the islands of 
southern California, extending southward to Lower California and the 
Gulf of California ; common also in the Pleistocene deposits of the same 
area; but there are only two dredging records, which indicates that it 
is definitely a shallow water species. 

Hancock Stations: 1378-41, Catalina Island, 2 to 3 fms; 1071-40, 
San Felipe Bay, Gulf of California at 2^^ fms. 

Lichenopora novae-zelandiae (Busk), 1875 
Plate 74, fig. 4 

Discoporella novae-zelandiae Busk, 1875:32. 

Lichenopora radiata, Robertson, 1910:262. 

Lichenopora novae-zelandiae, Harmer, 1915:155 (references), 

Zoarium encrusting on shells and stems; on small stems there is a 
very short stipe, and the margin of the basal lamina is turned upward, 
saucer-shaped > zoarial budding rarely occurs at the margin. The tubules 



706 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

are in radiating uniserial rows, high next to the central area (often 1.0 
mm or more in height), and sloping gradually to the edge, which is 
surrounded by a moderately broad basal lamina. Shorter rays appear 
between the main ones toward the margin. The tubules are connate to 
the tips, and slightly compressed, the apertures about 0.08 mm in di- 
ameter, the tips prolonged into points on the central side and often also 
on the outer side. The cancelli are extremely variable in size and form, 
producing an irregular network; the largest are more than twice the 
width of the zooecial apertures and the smallest are even less than the 
apertural width. There is no evidence of closure of the cancelli by an 
"iris" diaphragm, but instead they often become closed by a thin calcified 
membrane with numerous small pores like that which covers the ovicell, 
and this is true for some of the cancelli near the outer border beyond 
the brood-chambers. 'Tinhead" spicules are present within the apertures 
of the cancelli, often in nearly every one but sometimes more rare; 
usually they are present slightly above the closing membrane, and it may 
be that these cancelli are regenerated. Between the rays there are one 
or two rows of cancelli. In old and more heavily calcified specimens the 
walls of the cancelli are thicker but not closed by an iris-like diaphragm. 

The brood-chambers occupy the central area but often extend for a 
considerable distance into the interradial spaces; the roof consists of a 
thin calcified and perforated membrane and soon becomes covered with 
a secondary layer of cancelli. The ooeciostome is excentric in position, 
the tube short, the orifice round and about as large as that of a zooecial 
tube, the lip round or elliptical and slightly flared. 

Harmer, 1915:155, includes L. holdsworthi under L. novae-zelandiae ; 
Waters, 1918:36, reverses this and includes novae-zelandiae under 
holdsworthi, though the former has page priority in publication. From 
Busk's figures of these species, 1875, plate 30, figs. 2 and 4, there appear 
differences in the height of the peristomes, the mode of closure of the 
cancelli, and especially in the size of the central area, sufficient to war- 
rant their separation. L. holdsworthi has the appearance of a Discopo- 
rella. The L. holdsworthi of Canu and Bassler, 1929, plate 88, fig. 11, 
has short biserial rays and probably should go elsewhere. 

The species was described from New Zealand and later recorded from 
Australia, Ceylon and Japan. While it has not been listed from the 
American Pacific, our specimens conform so closely to the descriptions 
and illustrations of Busk and Harmer that they appear to belong to this 
species. Also I believe that the L. radiata of Robertson from southern 
California belongs here, and possibly that of O'Donoghue (without 
description) from British Columbia. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 707 

Hancock Stations: 468-35, Charles Island, Galapagos; 1399-41, Santa 
Catalina Island; 1242, Anacapa Island; 1002, San Clemente Island; 
Palos Verdes, near San Pedro, all from southern California; Acapulco 
Harbor, Hubbs Sta. 46-244, west coast of Mexico; and Colombia (with- 
out further data) ; shore to 77 fms. 

Lichenopora intricata (Busk), 1856 

Plate 76, figs. 5, 6, 7, 8, and 9 

Defrancia intricata Busk, 1856:179, 

Apparently this species has never been referred to since Busk described 
it. In December, 1946, Dr. E. Y. Dawson, while collecting algae at 
Mazatlan, Mexico, the type locality of D. intricata, recovered several 
specimens on algae. Again, in 1949, the "Velero" dredged more than 
100 specimens at Magdalena Bay, on the west coast of Lower California. 

These specimens conform to Busk's meager description: "Disc very 
irregular in form, rows of cells radiating irregularly; orifices of cells 
and interstitial pores of equal size. The small irregular patches appear 
to be constituted by the confluence of several sets of costae, with their 
corresponding interstices, each set radiating from a depressed central 
point." 

The form of the encrusting complex zoaria varies to such an extent 
as to baffle description; adnate on algae, worm tubes, corallines, other 
bryozoans, etc., the largest colonies 3 cm or more in length, the margins 
of the zoaria sometimes extending free. The subcolonies are very numer- 
ous, more than 70 having been counted on one large zoarium, and vary 
in form from nearly round to very elongate-elliptical. The radii 
("costae," Busk) are high, closely set, and rather regularly arranged 
about the low central area ; in general they are uniserial, but often they 
are biserial next to the central area and rarely biserial for the whole 
length of the radii ; separated by one or two rows of cancelli. The outer 
ends of the radii are often extended with short tubules into meandering 
series which break up into short, separate series or sometimes form small 
clumps. The subcolonies often arise in the midst of this intricate me- 
andering series, or they may be closely associated, with the low outer 
ends of their radii in contact. 

The central area is flat and low, even when ovicells are present. When 
an ovicell is present it is covered by a thin lamina and above this the 
secondary cancelli are large, thin-walled and irregular in form, in true 
Lichenopora fashion. The cancelli between the radii and in the central 
area, in the absence of ovicells, are rounded and partially closed, sug- 



708 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

gesting Disporella. In some elongate central areas I have observed an 
ovicell at one end covered by the irregular cancelli, while the other 
end of the area, free from the ovicell, shows the rounded, partially 
closed cancelli. This throws some doubt on the complete validity of 
Disporella, as infertile subcolonies would undoubtedly be referred to 
that genus. On the complex zoaria the fertile discs are easily seen 
because of their irregular secondary cancelli, and I have not been able 
to find any evidence of ovicells in discs with the uniformly rounded 
cancelli. 

The ovicells occupy all or a part of the central area and can often 
be seen through the large irregular cancelli ; occasionally two ovicells 
are present in the same area. The ooeciostome is a short, thin-walled, 
erect tube, situated near the border of the central area. 

Collected by Dr. E. Y. Dawson at Mazatlan, Mexico (the type 
locality), about 23° 11' N. Lat., shore collection, 4 zoaria, 1 on a shell 
fragment, the others on algae ; the ones on algae are much thinner than 
those on solid substrata. 

Hancock Station 1714-49, two miles east of Entrada Point, Mag- 
dalena Bay, west coast of Lower California, 24°32'30"N, 112°01'45" 
W, at 17 fms, more than 100 complex zoaria, in a single dredge haul. 

Genus DISPORELLA Gray, 1848 

Brood-chambers, one or more, occupying interradial areas and some- 
times extending over parts of the central area; cancelli thick-walled, 
partially closed by an "iris-like" growth of the rim toward the center 
but leaving always a small round aperture, never closed by a perforated 
flat calcified membrane; lateral zoarial budding is common. As in 
Lichenopora the functional zoids may be in radiating series, uniserial, 
biserial or multiserial and connate or non-connate, or they may be more 
or less in quincunx. Genotype, Discopora hispida Fleming, 1828:530. 

Key to Species of Disporella 

1. Radii uniserial or the tubules in quincunx 2 

Radii with 2 or more (2 to 4) series of tubules, sometimes 

arranged in short clumps 5 

2. Tubules not connate, except sometimes at the base only ... 3 
Tubules closely connate to their tips, rays longer 4 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 709 

3. Peristomes slightly expanded at the tips and bearing a number 

(3 to 5) of long thin spines fimbriata 

Peristomes not expanded at the tips, sometimes prolonged into 

a single process, but never fimbriated hispida 

4. Pores of the central cancelli larger than the tubule apertures ; 

pin-head spicules very abundant californica 

Pores of cancelli small, the v^^alls more heavily calcified ; pin- 
head spicules rare or wanting ovoidea 

5. Radii usually prominent in the form of short fascicles 2 to 4 

tubules in width 6 

Radii more elongate and less prominent, with 2 to 4 rows of 

tubules, zoaria often very complex 8 

6. Zoarium high, cylindrical, with a terminal crown of high 

marginal radii, the encrusting base larger than the erect 

stem; central, vertical budding astraea 

Zoarium low, without an erect stem 7 

7. Radial fascicles small and low, cancelli of central area nearly 

closed by a funnel-shaped diaphragm octoradiata 

Fascicles larger, with more tubules, and higher; cancelli large 

and nearly wide open alaskensis 

8. Zoarium highly complex, composed of numerous lateral sub- 

colonies which are separated by rows of cancelli; radii 

moderately high, 2 to 4 rows of tubules separata 

Zoarium simple; radii usually forming a low ridge of 2 to 4 
series of tubules; central area ovoid and moderately large; 
sub-colonies superposed vertically stellata pacifica 

Disporella fimbriata (Busk), 1875 
Plate 75, figs. 2 and 3 

Discoporella fimbriata Busk, 1875 :32. 
Lichenopora fimbriata. Busk, 1886:26. 
Disporella spinulosa Jullien, 1888:83. 
Lichenopora fimbriata. Waters, 1904:96; 1905:250. 
Lichenopora fimbriata, O'Donoghue, 1923:15. 
Disporella fimbriata, Borg, 1944:229. 

Busk's original description is as follows: "Zoarium almost conical; 
cells very indistinctly serial, distant; interstitial pores almost obsolete; 
mouth expanded, peristome fimbriated." 



710 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

When the brood-chamber fills the central area the zoarium is "nearly- 
conical," as shown in Busk's illustration (1875, PL 27, figs. 1 and 2), 
but in the absence of the chamber the area is depressed and slightly 
concave. The "cells" or zoids are often in short radial series of 3 or 4, 
but frequently are irregularly quincunical. The "interstitial pores" or 
cancellae are much less numerous than in other species and usually more 
widely separated ; when young they are as large as the apertures but 
later become partly closed, with a small central pore. The peristomes 
project strongly and are somewhat flared ("mouth expanded") and 
fimbriated with 2 to 5 marginal spines. The basal lamina is very broad 
and turned upward at the edge, as shown in Busk's figure 2. 

The brood-chambers, 1 to 3 or 4 in number, are prominent, usually 
coalesced to more or less fill the central area, but in one of our specimens 
the 3 chambers are distinct; there are numerous pores in the ooecial 
cover; the ooeciostomes situated more or less between the inner ends of 
the rays, the aperture about the size of those of the zoids, the tube short 
and very slightly flaring but without a distinct lip. 

The zoaria are all small, the largest slightly over 4 mm in diameter, 
the peristome and ooeciostome about 0.10 mm. 

Busk described the species from the southern tip of South America, 
Chonos Archipelago, Tierra del Fuego, Cape Horn and Chiloe, and 
later added Tristan da Cunha. The Disporella spinulosa of Jullien was 
dredged between the Falkland Islands and the Strait of Magellan. It 
has also been recorded from Australia, Tasmania, New Zealand, the 
Azores and Cape Verde Islands; O'Donoghue has recorded it from 
Round Island, British Columbia. If these identifications are all correct, 
the species has a very wide distribution. 

Hancock collections: not dredged, but taken in low tide collecting 
by the writer at Palos Verdes near Los Angeles; by Miss A. E. Blagg 
at Pescadero Point outside of Monterey Bay; and recovered from a 
sunken buoy brought up from 45 fms, off Rocky Point, near Los Angeles, 
all from southern California. 

Disporella hispida (Fleming), 1828 
Plate 75, fig. 1 

Discopora hispida Fleming, 1828:530. 
Discoporella hispida. Busk, 1875:30. 
Lichenopora hispida, Hincks, 1880:473; 1884:207. 
Lichenopora hispida, O'Donoghue, 1923 :15 ; 1926:28. 
Lichenopora hispida, Canu and Bassler, 1923:203. 
Lichenopora hispida, Osburn, 1923 :5D; 1933:18. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 711 

The zoarium is usually rounded, attached more or less over the whole 
dorsal surface but sometimes only by a very short stipe, surrounded by 
a moderate bordering basal lamina which is sometimes turned slightly 
upward ; the central part of the colony in young stages is a little de- 
pressed and with rounded cancelli which become partially closed. Adult 
colonies, with brood-chambers, are usually evenly rounded over the top. 
The tubules vary much in their arrangement, sometimes occurring in 
radiating uniserial rows in which, however, the tubes are not connate, 
or at least are free at their tips; for the most part they are irregularly 
quincuncial, and they are separated by rounded cancelli about as large 
as the apertures of the zoids, about 0.08 mm in diameter. The peristomes 
are a little elevated, rising on the central side into a pointed cusp which 
is sometimes double or tricuspidate. 

The ovicells, or brood-chambers, are located at the edges of the central 
area and extend outward between the zooecial tubes ; occasionally, when 
more than one is present (I have noted as many as 4) their expanded 
inner ends may cover the central area. The ooeciostome is situated at 
the edge of the central area or farther out between the tubules, short, 
cylindrical, with a round aperture which is somewhat larger than that 
of the tubules, about 0.10 mm in diameter. The chamber at first is cov- 
ered by a thin, minutely perforated calcified layer, but later this may 
secondarily be covered with a cancellous layer. 

It is a well known northern and arctic species, extending on the 
Pacific coast south to Lower California. 

Hancock Stations: 1260-41, off San Eugenio Point, Lower California, 
27°49'50"N, 115°06'05'"W, the southernmost record; oflf Santa Cata- 
lina, Santa Barbara and San Miguel Islands, and Albatross Sta. 2938, 
all from southern California; from near shore to 34 fms. Also a speci- 
men labelled "Bering Sea," with no other data. 

Disporella calif ornica (d'Orbigny), 1853 /--O 

Plate 74, figs. 7, 8, and 9 

Unicavea Californica d'Orbigny, 1853:972. 

Not Lichenopora californica, Gabb and Horn, 1862:176. 

Not Discoporella calif ornica. Busk, 1875:32. 

Not Lichenopora calif ornica. Waters, 1889:282. 

Not Lichenopora californica, Robertson, 1910:261. 

Lichenopora californica, Borg, 1944:219. 

D'Orbigny's description, without illustration, reads: "Espece tres- 
convexe en dessus, ayant le centre excave, et pourvue de pores inter- 
mediares enormes. Madelaine, Basse-Californie." 



712 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

The californica of d'Orbigny was placed by him in the genus Uni- 
caveUj which indicates that his species has uniserial rays. On the other 
hand the californica of Busk, Gabb and Horn, Waters, and Robertson 
is definitely stated to have biserial or triserial rays and has been rede- 
scribed as Lichenopora buskiana by Canu and Bassler, 1928:164. The 
description of the zoarial form by d'Orbigny might apply to numerous 
species, but his final statement of the large size of the cancelli is more 
definite and is an exact statement of their nature. In older colonies the 
cancelli become partially closed by an "iris-Hke" thickening of the in- 
ternal wall, but the outlines of the large pores are evident in the raised 
separating ridges. Moreover, the species is common in the area where 
d'Orbigny obtained his material, Lower California, and we are fortunate 
to have ten specimens from, three stations in Santa Maria Bay and 
Magdalena (Madelaine) Bay, that is, in the type locality of californica. 
It appears very probable, therefore, that after a century d'Orbigny's 
species has been resurrected. 

The zoaria are round, low dome-shaped with the central area flat or 
somewhat depressed in the young. The colonies are all small, not over 
4 mm in width, the central area one-fourth to one-third as wide as the 
zoarium ; the radiating rows of tubules are all definitely uniserial, about 
10 primary rows with shorter ones between them toward the margin. 
The peristomes are only moderately elevated, slightly higher toward the 
central area, connate to their tips which usually are truncate but some- 
times are extended into short points on their distal borders; the aper- 
tures are slightly elongated in the direction of the rays, about 0.10 mm 
long by 0.08 mm wide. The cancelli of the central area are noticeably 
larger than the tubules, the apertures round and as much as 0.13 mm 
in diameter, partially closed by the characteristic "iris" diaphragm ; the 
pin-head spicules are abundant. Between the rays there are two rows 
of cancelli, occasionally only one, which are somewhat smaller than at 
the center. 

The brood-chambers are interradial or extending somewhat into the 
central area, the roof a thin calcified membrane with minute pores, 
later covered by secondary cancelli of the usual type. The ooeciostome is 
short, round, thin-walled and a little larger than the zooecial apertures. 

There is a peculiar type of zoarial budding which I have not seen 
described and which I have observed in only one other species, D. 
alaskensis new species, described in this report. The sub-colonies arise 
on the frontal side toward the margin but do not extend beyond it and 
in the three colonies at hand they are exactly similar in origin. When I 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 713 

first observed one of these I thought it might be a monstrosity or per- 
haps due to the attachment of an ancestrula, but the discovery of three 
similar triple colonies and a very young bud on another proves it to be 
a normal process. The sub-colonies are short stipitate with their borders 
and most of the dorsal side entirely free. Thus they have some resem- 
blance to d'Orbigny's "genus" Tecticavea, except that the sub-colonies 
arise near the margin and are not superposed on the central area. In 
each case the first sub-colony bears another similar to it but smaller. 
They present the same characters as the primary one, with uniserial 
radii, large central cancelli, moderately low connate peristomes and 
inter-radial brood-chambers. 

Hancock Stations: 279-34, Santa Maria Bay, Lower California, 
24°44'45"N, 112°15'20"W, and 1714-49 and 2180, Magdalena Bay, 
the type locality of californica d'Orbigny, 10 to 18 fms. Also at 1242, 
Anacapa Island, and 1662-48, Santa Cruz Island, southern Cahfornia; 
1889-49, Cortez Bank at the United States-Mexican boundary; 275, 
Raza Island, 675-37, Carmen Island, and 1044-40, Tiburon Island, 
Gulf of California; and 468-35, Port Parker, Costa Rica. Depth 5 to 
77 fms. Also 3 colonies from Tobago Island, Panama, each consisting 
of several sub-colonies, Helen Hoyt, collector. 

Disporella ovoidea new species 

Plate 75, figs. 4 and 5 

Lichenopora radiata, Canu and Bassler, 1928:163; 1930:56. 
Lichenopora radiata, Osburn, 1940:334; 1947:6. 

Zoarium more or less ovate, in older stages becoming low dome- 
shaped ; the central area large, distinctly elongate, ovoid to elliptical, 
much depressed in the young but thick and elevated nearly to the tips 
of the zooecial tubes in older colonies ; 3 to 5 mm in the longest dimen- 
sion. The zoids are in very definite uniserial rays, the longest ray noted 
having 7 zoids. The tubes are moderately short and are connate to their 
tips, which are without spinous projections or notches; the apertures 
elongated in the direction of the rays, averaging 0.07 mm wide by 0.10 
mm long, those at the outer ends of the rays usually larger than those 
near the central area. The cancelli are large, about twice the size of the 
zooecial apertures, but very soon become partially closed by an iris-like 
diaphragm so that their apertures are funnel-shaped and surrounded by 
hexagonal separating ridges. 'Tin-head" spicules are sometimes present. 



714 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

I have not been able to determine the nature of the primary brood- 
chambers near the central area, but the secondary chambers near the 
border are covered in the usual manner by a calcified porous membrane ; 
here they lie between the rays, in some cases extending on both sides of 
a short secondary ray. They are soon covered by secondary cancelli. 
The ooeciostome is hardly distinguishable from the cancelli in height 
and size, but the orifice is w^ide open, rounded and its wall thin. 

I must agree with Borg (1944:223) that the L. radiata of Canu and 
Bassler (1928:163 and plate 29, figs. 1-2) from north of Cuba, and 
those of Osburn (1940:334) from Porto Rico, cannot be identified with 
Discoporella radiata of Waters (1879:276) from the Bay of Naples, 
nor with the Melobesia radiata of Audouin (1826:235). Waters states: 
"In most specimens the cancelli appear open; but in well-preserved ones 
a delicate calcareous cover is found covering the aperture: and this is 
perforated with about 2-10 holes," which is clearly shown in his plate 
24, fig. 11a. The figures of Melobesia radiata Audouin show a round 
zoarium with a small round central area ; a central brood-chamber cov- 
ered by a calcareous porous membrane and with lobes extending between 
radii; the radii high, elongate and uniserial, the tubes connate to the 
tips and ending in sharp points. Apparently there is no other species 
recorded from the Mediterranean or Red Seas with which Waters could 
have confused his D. radiata, and we must conclude that it is Audouin's 
M. radiata and is a Lichenopora in the strict sense. 

On the other hand, the L. radiata of Canu and Bassler and of Osburn, 
from the West Indies, has an ovate or rounded zoarium with a large 
ovate central area ; the cancelli thick-walled and without a covering 
calcified membrane ; the brood-chambers not centrally located ; the uni- 
serial connate radii much less elevated. These West Indian specimens 
appear to conform in every particular with Disporella ovoidea, as de- 
scribed above, and it is probable that Canu and Bassler's reference to 
L. radiata from the Galapagos Islands is also to the same species, since 
Dr. Bassler informs me (in litt.) that it has "a large, slightly elongate 
central area, with the cancelli and rows of tubules as in the Cuban one." 
How many other references to radiata are untenable it is impossible to 
say, as it has often been recorded without description or figures, but it 
seems safe to state that it has not been found on the Pacific coast of the 
Americas. 

Our material consists of 4 colonies, 2 from the Galapagos Islands, 1 
from Colombia and 1 from southern California, a wide distribution to 
be sure, but they all agree in the elongate form of the central area, the 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 715 

short uniserial connate rays, the size and form of the zooecial apertures, 
and the size, form and nature of the closure of the cancelli. It is very 
different from any other Eastern Pacific species, but it may represent 
some of the too numerous lichenoporid species that have been inade- 
quately described from all around the world. 

Type, AHF no. 126. 

Type locality, Hancock Station 432, Tagus Cove, Albemarle Island, 

Galapagos, 80 to 100 fms, two colonies, with ovicells. Also 1662-48, 

Santa Cruz Island, 33°55'45"N, 119°32'30"W, southern California, 

23 fms ; and one colony from Colombia without further data. 

Disporella alaskensis new species 

Plate 75, figs. 7 and 8 

The zoarium is round, 3 mm in diameter, high at the central area, 
the broad cancellated thin border turned up all around, shaped like a 
miniature Mexican straw hat ; attached over most of the dorsal surface. 
The radii are multiserial (2 to 4), consisting of elevated ovoid clumps 
which are regularly arranged about the central area. The outer ends 
of the radii descend sharply to the thin bordering lamella. In our two 
specimens, the smaller has 4 radii with 2 developing between these at 
the edges, the larger has 8 rays with several smaller incomplete ones. 
The tubules are completely connate to their tips, which are not extended 
into points, the apertures rounded or slightly hexagonal and about 0.10 
mm in diameter. 

The central area is moderate in size, short-ovate in form, with large 
rounded cancelli (0.13 mm) and the cancelH of the bordering area are 
of the same size and form (occasional smaller ones are present on the 
central area and between the radii) ; there are 2 to 4 rows of cancelli 
between the radii. Small pinhead spicules are present. There is very 
little closure of the cancelli of the central area, just enough to suggest 
an "iris-like" diaphragm, and the bordering cancelli are wide open. 

A small sub-colony is present on the front, situated at the outer end 
of one of the rays and well within from the border; this has the same 
form, with edge strongly turned up and the tubules and cancelli similar 
to those of the primary colony. 

The ovicell is interradial, extending somewhat into the central area 
and covered by a thin membrane with minute pores. Unfortunately the 
ooeciostome is broken away. 

Type, U. S. Nat. Mus. no. 11052. 



716 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Type locality, Stepovak Island, Alaska, Alaska Crab Investigation, 
Sta. 84-40, 15 fms. Another colony, the older one, is from Cleveland 
Passage, Alaska, 10 fms, W. Williams, collector. 

The older colony differs from the type specimen only in the larger 
number and greater prominence of the radii and in the absence of an 
ovicell. 

Disporella stellata var. paclfica, new variety 
Plate 76, fig. 10 

Defrancia stellata Reuss, 1847:37. 

Defrancia stellata, Canu and Bassler, 1930:57. 

Defrancia Bronn, 1825, is considered synonymous with Apsendesia La- 

mouroux, 1821, by Bassler, 1935:48. 

Canu and Bassler, 1930:57 and Plate 14, figs. 7-12, described a 
specimen of this Miocene form as Defrancia stellata, from the Gala- 
pagos Islands. As they remark, "It is quite remarkable to rediscover in 
the recent seas this European fossil." However, the measurements agree 
with those of the fossils and the specimen photographed (fig. 9) corre- 
sponds in a remarkable way to the figures of the fossil specimens shown 
beside it. It is possible that a species may have continued to live from 
Miocene time and be distributed half way round the world, but the 
chances are very much against it. Since we know nothing of the ovicells 
of stellata, it seems better to give the recent form at least a varietal 
name, pending the discovery of the ovicells of stellata. 

From the Hancock dredgings at the Galapagos Islands 12 specimens 
have been recovered from 4 different stations, similar to that discussed 
by Canu and Bassler, but bearing ovicells which are definitely those of 
a Disporella. 

The zoaria are attached to corallines; discoid in form, thick, with a 
narrowly extending basal lamina- the central area large, nearly flat, 
round or ovate in form, and the radii on the slope of the zoarium ; the 
colonies are of moderate size, from 2 to 4 mm in diameter. The radii are 
multiserial with 2 to 4 (usually 3) series of tubules which are closely 
connate to their tips, and which form elevated ridges separated by 2 to 
4 rows of cancelli. The apertures measure 0.08 mm in diameter and the 
cancelli 0.08 to 0.10 mm, depending on the amount of closure. 

Vertical budding appears to be a constant character, as even the 
smallest colonies have at least one sub-colony superimposed and arising 
near the center of the frontal area ; as many as 3 sub-colonies are present 
in one specimen, vertically arranged. In one specimen a second bud is 
present at the edge of the central area, indicating the beginning of a 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 717 

branched colony. In another case what appears to be lateral budding 
involves 3 colonies (or sub-colonies) ; these might have been produced 
by the fusion of separate colonies, but if so there is no definite line of 
demarcation. 

The ovicells or brood-chambers, shown at or near the surface in two 
of our specimens, are either at the edge of the central area and extending 
between the rays or are farther out and entirely interradial or both; 
they show the calcified bottom layer, which covers the submerged can- 
celli, and the minutely perforated roofing layer. The roof of the ovicell 
is again closed by secondary cancelli of the usual type. 

The ovicells appear to place this form definitely in Disporella, and 
the character of the cancelli with thick walls (though they are but little 
closed) also suggests this disposition. At the same time, the normal verti- 
cal arrangement of the sub-colonies indicates Lichenopora but, as has 
been shown above, this character does not appear to have positive generic 
importance. 

Recorded by Canu and Bassler at Albatross Station D. 2815, Gala- 
pagos Islands. 

Type, AHF no. 127. 

Type locality, Hancock Station 143-34, Wenman Island, Galapagos, 
1°23'10''N, 91°48'45''W, 100-150 fms. 

Also at Hancock Stations: Galapagos Islands, 155-34, Albemarle 
Island; 453, Gardner Island, and 454, Hood Island; 30 to over 100 fms. 

Disporella separata new species 
Plate 74, figs. 5 and 6 

Zoarium a very complex colony of the kind known as Radiopora by 
d'Orbigny, Busk, etc. It consists of about 30 sub-colonies rather reg- 
ularly arranged over a rounded area about 15 to 20 mm, attached loosely 
and spreading over the surface of a small dead barnacle and the shell 
to which the barnacle is attached ; most of the basal lamina is free. The 
sub-colonies are all well separated from each other by a few rows of 
cancelli and are quite regular in size and form ; the discs are short-ovate, 
about 2.5 by 2 mm in diameter, with the radii varying in number from 
8 to 12. The rays consist of small ovate clusters of peristomes, biserial 
or triserial, which often become uniserial at the outer end ; not infre- 
quently uniserial rays are present, and sometimes these may become 
biserial at the outer ends; while the triserial cluster appears to be the 
dominant form, all of these variations may be found on a single sub- 
colony and on any part of the complex zoarium. The peristomes are 



718 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

moderately high near the center and become gradually shorter outward, 
connate to their tips, which form a single acuminate spine at the point 
of junction; the apertures about 0.10 by 0.08 mm. 

The central area is concave in younger stages to nearly flat in older 
sub-colonies, elliptical in outline ; the cancelli about as large as the aper- 
tures of the tubules, partially closed by an "iris-like" diaphragm with a 
large central pore. The interradial and intercolonial cancelli do not 
differ from those of the central area, except that they vary more in size 
and the amount of closure. 

The ovicells are interradial and covered by a layer of secondary can- 
celli, and the ooeciostome is short, thin-walled, round, without a flaring 
border, barely elevated above the level of the cancelli, and measures 
0.08 mm in diameter. 

Young marginal sub-colonies develop near the border along with the 
proliferation of the lamina after 3 or 4 rows of cancelli are formed. 
There are several such incomplete discs at the edge of the zoarium, with 
the first few radii outlined on the side toward the center of the zoarium. 

This species belongs to the "Radiopora" group in which the sub- 
colonies are distinct (the discs not confluent) and their discs similar to 
that of the primary colony (see Waters, 1918, plate 4, figs. 1-4), but 
appears to be different from any of the recent "Radiopora" species de- 
scribed, Discopora meandrina Peach, Radiopora irregularis J. Y. John- 
son, Discoporella pristis MacGillivray, and Lichenopora bullata and 
L. magnifica MacGillivray. 

Type, AHF no. 128. 

Type locality, Hancock Station 1889-49, Cortez Bank, west of the 
United States-Mexican boundary, 32°27'05'"N, 119°08'04"W, at 15 
to 20 fms. 

? Disporella octoradiata (Waters), 1904 
Plate 75, fig. 6 

Lichenopora octoradiata Waters, 1904:97. 
Disporella (?) octoradiata, Borg, 1944:257. 

Waters' description is as follows: "The zoarium is very solid and 
much raised, with the base narrower than the disk. There are a number 
of biserial rays, formed by a few zooecia, and in a well developed colony 
there are 8 main rays, with indications of the commencement of another 
series. The rays do not extend to the border of the zoarium, nor are 
the zooecia around the border of the disk elevated, while in the center 
of the zoarium the openings are round and vary in size." As far as it 
goes this is as fairly complete a description of our two young specimens 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 719 

form around the central area, and vary in size from 2 to 6 tubules, con- 
siderably elevated above the central area which is somewhat concave. 
The zooecial apartures measure 0.10 mm in diameter. The larger can- 
celli are of about the same size, but most of those in the central area 
are partially closed, with a smaller rounded central pore; there are 
usually two rows of cancelli between the radii. The zoids around the 
border are not at all elevated and in most cases are indistinguishable 
from the cancelli. There is a narrow basal lamina. 

The zoaria are evidently young, as there are no ovicells, and there is 
a question whether the species is the same as L. octoradiata Waters. The 
nature of the closure of the cancelli appears to relate it to Disporella 
rather than to Lichenopora. 

Waters described the species from 71°09' S. Lat., 89° 15' W. Long., 
and Borg recorded it questionably from 63°57'S, 61°50'W; both of 
these records are from the area between South America and Antarctica. 

Hancock Station 481, Cartago Bay, Albemarle Island, Galapagos, at 
12 fms, two colonies. 

Disporella astraea, new species 
Plate 76, figs. 1 and 2 

Zoarium encrusting with a broad base and rising by vertical budding 
into a short cylindrical stalk which, with the radiating fascicles, gives 
the appearance of a minute Astraeid coral. The central area is flat with 
numerous thick-walled and rather wide open round cancelli, which extend 
between the rays in two or three rows. The flat top is surrounded by a 
ring of 10 high, short fascicles. The fascicles are groups composed of 
4 to about 10 zooecial tubes which are all closely connate, their apertures 
about 0.07 mm in diameter; the apertures of the cancelli are about the 
same size, occasionally larger. The encrusting base is 2 mm in width ; 
the primary zoarium arising from it 1.30 mm wide and about 0.60 mm 
high; the secondary zoarium or vertical bud is 1.10 mm in diameter and 
about 0.80 mm high. 

There is no evidence of an ovicell, and therefore the disposition of the 
species in Disporella is questionable and based merely on zoarial char- 
acters. 

Type, AHF no. 129. 

Type locality, Hancock Station 451, off Post Office Bay, Charles 
Island, Galapagos, at 100 fms, one colony. Another somewhat smaller 
colony at Station 461, off Tagus Cove, Albemarle Island, Galapagos, 
at 80 fms ; this specimen has 9 slightly smaller and higher fascicles, but 
otherwise is similar. 



720 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

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NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 721 

Busk, G. 

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722 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

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726 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Order ECTOPROGTA 
Suborder GtENOSTOMATA 

By Raymond C. Osburn, Ph.D., D.Sc. and John D. Soule, Ph.D. 

This taxonomic report on the Pacific Coast Ctenostomata was pre- 
pared by Dr. Soule under the immediate direction of the senior author. 
The work was done in connection with a study of postlarval develop- 
ment and histogenesis and the bearing of the results on the classification 
of this group. The data on which the taxonomic changes are based will 
be published elsewhere and bear the full approval of the senior author. 
The new species which have appeared during the progress of the work 
are all to be credited to the careful work of Dr. Soule. 

R. C. O. 

Sub-Order GtENOSTOMATA Busk, 1852 

The chitinous zoaria may be incrusting, erect, stolonate or burrowing. 
The zooecial aperture is essentially simple, being closed by the inversion 
of the tentacle sheath on retraction of the polypide. In some genera spe- 
cialized apertures are present, including those that are bilabiate, pro- 
duced or even operculate. The operculum present in one genus of 
burrowing ctenostomes is analogous to the opercula of the cheilostomes. 
No avicularia or true external ovicells are present, although specialized 
gonozoids do occur. Kenozooecia, modified as stolons, are present in the 
stolonate groups, or as spines in the carnose forms. 

Division 1. Garnosa Gray, 1841 

Ctenostomata that have in common a comparatively heavy non- 
calcareous cuticle, giving the zoaria a fleshy or leathery appearance. The 
colonies included within this group are usually incrusting, but they may 
rise in thin flabellate or palmate fronds, sac-like expansions, or they may 
be cylindrical or pedunculate structures. 

Key to the Families of the Division Carnosa 

1. Zoaria primarily incrusting 2 

Zoaria erect, clavate, with kenozooecial peduncle . . Clavoporidae 

2. Zooecia with aperture closed by simple folds . . . Alcyonidiidae 
Zooecia with modified apertures 3 

3. Aperture bilabiate, zooecia with kenozooecial spines . Flustrellidae 
Aperture raised, quadrangular, zooecia with 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 727 

Family Alcyonidiidae Johnston, 1849 

Zoaria incrusting or erect in sacculate or cylindrical expansions. Aper- 
ture closed by simple folds formed by the invagination of the tentacle 
sheath when retracted, producing a puckered or drawn appearance. 

Genus ALGYONIDIUM Lamouroux, 1812 

Zoaria incrusting, coriaceous or gelatinous in appearance, forming a 
soft cover over the substrata, or arising into lobed sac-like, or cylindrical 
expansions. Zooecia closely united, not stolonate. The aperture may be 
in the center of raised papillae, or the entire ventral surface of the 
zooecia may present a smooth surface, a slight puckering at the distal 
end indicative of the aperture. Genotype: Alcyonium gelatinosum Lin- 
naeus, 1767. 

Key to the Species of Alcyonidium 

1. Zoaria primarily incrusting, spreading irregularly 2 

Zoaria primarily erect, or disc-shaped, limited 4 

2. Zoaria flat, incrusting, zooecia irregularly hexagonal . . polyoum 
Zoaria flat, incrusting, zooecia with raised apertures .... 3 

3. Zoaria argillaceous, zooecia with fine papillate border . parasiticum 
Zoaria clear, zooecial aperture mammillate . . . mammillatum 

4. Zoaria disc-shaped, flattened disctforme 

Zoaria primarily erect 5 

5. Zoaria sacculate, expanded, lobed pedunculatum 

Zoaria elongate, cylindrical enteromorpha 

Alcyonidium polyoum (Hassall), 1841 
Plate 77, fig. 1 

Sarcochitum polyoum Hassall, 1841 :484. 
Alcyonidium mytili, Robertson, 1900:329. 
Alcyonidimn polyoum, Robertson, 1900:330. 
Alcyonidium mytili, O'Donoghue, 1923:191; 1926:54. 
Alcyonidium columbianum O'Donoghue, 1926:56. 

The zoaria of the specimens in the collection show a great deal of 
color variation, ranging from transparent to brown or gray. In size the 
colonies ranged from 1 to 6 cm in breadth depending upon the size and 
type of subtrate. These zoaria are found incrusting rocks, mollusk 
shells, algal holdfasts and sometimes on the larger Crustacea. 



728 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

The zooecia are irregularly hexagonal, but zooecia that are pentagonal, 
quadrangular and some that are nearly square are not uncommon. This 
wide variation in shape may account for the differences found in the 
measurements that have been previously cited in the literature. 

The zooecial walls are usually distinct. The apertural openings in 
some of the zoaria are found on small raised papillae, distally located on 
the ventral wall, while in other zoaria there are no papillae, the ventral 
surface being smooth. In the latter case, the openings are either easily 
discerned, or are very obscure. One recent author (Silen, 1942:9-11) 
considers A. polyoum one of the species with a smooth ventral surface. 
Other authors (Hincks, 1880:501 ; Osburn, 1933:61 ; 1944:16; Marcus, 
1941:68) have all noted the presence of a raised oral papilla. It is 
possible that the presence or absence of the oral papillae may be due to 
the degree of retraction of the tentacle sheath. If so, a given living 
zoarium could exhibit no oral papillae at one time, and have them at 
another. 

The tentacle number poses another problem in this species. The vex- 
ing question is, does this species have a fixed number of tentacles, a 
variable number of tentacles, or are there two or possibly three species 
similar in external appearance being lumped together as A. polyoum? 
The reported tentacle number varies from 12 (Harmer, 1915:38) to 
20 (Silen, 1942:11). The original description (Hassall, 1841:484, 
485) reports the tentacle number as 20. If then the number to be con- 
sidered as correct is 20, what is to be the disposition of those with 16 
tentacles (Marcus, 1941:68; Rogick, 1949:47), unless A. polyoum is 
considered as having a wide variation in tentacle number. In order to 
determine the number of tentacles in the specimens from the eastern 
Pacific found in the Hancock collection, comprising at this time 9 sta- 
tions (5 from Alaska, 4 from northern California), a sample of each 
of the best preserved specimens with the external characteristics of 
A. polyoum was sectioned. None of the specimens had 20 tentacles. Two 
Alaskan specimens, one with raised papillae, and one without, had 17 
tentacles. One Californian specimen with definite oral papillae had 17 
tentacles. Three others, all from Californian waters, without oral pa- 
pillae, had 15, rarely 16 tentacles. Until such time as additional material 
can be obtained of both Pacific and Atlantic origin, the only safe con- 
clusion is that A. polyoum does have a variable number of tentacles. 

Akyonidium polyoum is widely distributed in the colder waters of 
both the Atlantic and the Pacific. In the eastern Pacific it has been 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 729 

previously reported by Robertson, 1900, and O'Donoghue, 1923, 1926, 
in the waters off Alaska, British Columbia, and Puget Sound. 

The specimens in the Hancock collection are from off Point Barrow, 
Alaska, Arctic Research Laboratory, G. E. MacGinitie collector; 
Lenard Harbor, Alaska, Canoe Bay, Alaska, Tomales Bay, California, 
AHF stations 1607-48 and 1656-48 in depths ranging from intertidal 
to 40 fathoms. (8 stations.) 

Alcyonidium parasiticum (Fleming), 1828 
Plate 77, fig. 2 

Alcyonium parasiticum Fleming, 1828:518. 
Alcyonidium parasiticum, O'Donoghue, 1923:191. 

The collection has one large zoarium, thin, incrusting upon an eroded 
mollusk shell. The individual zooecia may be distinguished with some 
difficulty, due to the deposit of sand and mud which covers most of the 
zoarium. The zooecia are small, irregular in morphology, the variation 
ranging from nearly square zooecia to those that are elongated to nearly 
diamond-shaped. All of the zooecia that could be examined possessed 
raised oral papillae on the ventral surface, and minute border papillae. 
The argillaceous cover upon the cuticle prevented sectioning of a portion 
of the specimen. 

This species is well distributed throughout the colder Atlantic waters 
and has been reported by O'Donoghue from the Pacific northwest. 

The specimen in the Hancock collection came from Tomales Bay, 
California, at a depth of 5 fathoms, collector R. C. Osburn. 

Alcyonidium mammillatum Alder, 1857 
Plate 77, fig. 4 

Alcyonidium mammillatum Alder, 1857:154. 

Alcyonidium mamillatum, O'Donoghue, 1923:191; 1926:54. 

The zoaria form dark brown, thin, rough, irregular incrustations 
upon mollusk shells. The zooecial walls are well defined, except in the 
portions of the zoaria that are covered by foreign matter. The zooecia 
vary in shape from an elongated irregular oval to rectangular. Distally 
the zooecial apertures are raised upon short cylindrical, transversely 
wrinkled projections. 

The literature reveals that this species is moderately well known from 
the cold waters of the Atlantic. On the Pacific coast of North America, 
O'Donoghue has reported it from the vicinity of Vancouver Island, 1923, 
1926. 



730 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Hancock Station 1642-48, off Point Vicente, southern California; 
also Friday Harbor, Puget Sound, J. L. Mohr, collector; Cold Bay, 
Alaska, U. S. Alaska Crab Investigation; and off Newport, southern 
California, which well may be the southern extension for this species. 
The known depth range is 15 to 70 fms. 

Alcyonidium pedunculatum Robertson, 1902 

Plate 77, fig. 3 

Alcyonidium pedunculatum Robertson, 1902:106. 
Alcyonidium pedunculatum, O'Donoghue, 1926:55. 

The zoaria of this unusual species are erect, arising from a short 
"peduncle" into wide flat saccate expansions, the largest of those in the 
Hancock collection, from Puget Sound, Washington, measuring 6.5 cm 
high and 4,5 cm wide. The largest from Alaska measures 11 cm long 
and 3 cm in width. The "peduncles" are wrinkled, rough, coriaceous 
in appearance, short, stout, cylindrical, and contain a loose reticular 
connective tissue. The zooecia are not modified as they are in the true 
peduncle of Clavopora, i.e., for bending and swaying the colony. The 
expanded portion of the zoarium is sac-like, filled with loose connective 
tissue, and may have several finger-like projections, or it may be a single 
foliaceous lobe. These lobes are smooth, light brown in color. The 
zooecial outlines are well marked, an irregular hexagonal shape. Sec- 
tioning disclosed the tentacle number to be 17. 

Miss Robertson's specimens were from the Pribilof Islands, Alaska. 
O'Donoghue (1926) reported the species from the Vancouver Island 
region. 

The specimens in the Hancock collection are from Alaska, Arctic Re- 
search Laboratory, G. E. MacGinitie, collector, and Puget Sound, 
Washington, J. L. Mohr, collector. There are 10 stations ranging in 
depth from 20 to 35 fathoms. 

Alcyonidium disciforme (Smitt), 1871 
Plate 77, figs. 5 and 6 

Alcyonidium mammillatum var. disciforme Smitt, 1871:1122, 1123. 
Alcyonidium disciforme, Osburn, 1936:540. 

The mature zoaria have a very distinctive, characteristic morphology. 
Resembling a common wide rubber washer or large coin with a circular 
hole punched from its center, these zoaria form circular, slightly convex 
discs, which apparently rest upon soft sandy substrata. Young colonies 
lack the central hole. Minute, fine root-like extensions from the basal 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 731 

side help to anchor the colony in place. These kenozooecial filaments 
are most easily found near the periphery of the zoaria. The zoaria in 
the collection measured between 2.6 and 3.0 cm in diameter. The zooecia 
are small, hexagonal, usually bearing the apertures raised on papillae 
which occupy nearly all of the ventral surface. The tentacle number is 
16, determined from sections. 

Described by Smitt from Scandinavian waters, and recorded by Os- 
burn from Captain R. A. Bartlett's dredgings in Wakeham Bay, Un- 
gava, Canada. 

Hancock collection specimens are from Point Barrow, Alaska, Arctic 
Research Laboratory, 13 fms, collected by G. E. MacGinitie. The 
species evidently has a circumpolar distribution. 

Alcyonidium enteromorpha Soule, 1951 
Plate 77, figs. 7 and 8 

Alcyonidium enteromorpha Soule, 1951:367. 

The zoaria are elongate without lateral branching, bearing a super- 
ficial resemblance to the intestinal tract of a small mammal. Of several 
zoaria in the collection the longest measured 61 cm in length and from 
4 to 6 mm in width. Coiled in several loose folds the zoaria are attached 
to the substrate without a differentiated "peduncle." The cuticle is 
firm, mottled light brown to tan in color, and only moderately thick. 
The zoaria are cylindrical and filled with a loose reticular connective 
tissue. Within this meshwork of connective tissue may be found numer- 
ous brown bodies, the product of degenerated zoids that have entered 
the central cavity when the thin dorsal zooecial walls were ruptured. 
From the ventral surface the zooecia are well defined, most easily found 
in the portions of the zoaria where the cuticle is thin. On the greater 
part of the zoaria, the lateral zooecial walls can be only faintly discerned, 
and while not totally obscured, they are rather difficult to trace. The 
ventral zooecial walls are smooth, with no oral papillae present. As 
noted before, the dorsal zooecial walls are thin, almost to the point of 
transparency. In shape the zooecia are varied, ranging from rectangular 
to irregularly hexagonal, those containing mature polypides measuring 
between 0.23 and 0.40 mm in length, and from 0.11 to 0.25 mm in 
width. The tentacle number obtained from serial sections is 17. It 
differs from A. pedunculatmn Robertson, by virtue of its cylindrical 
form, its extreme zoarial length, and its complete lack of a "peduncle. 

All of the specimens in the Hancock collection are from Alaska, off 
Point Barrow, Arctic Research Laboratory, collector G. E. MacGinitie. 
Collected at depths ranging from 80 to 123 fathoms. 



732 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Family Flustrellidae Hincks, 1880 

PiM^ Zoaria incrusting or rising in flabellate extensions. The aperture is 

( » bilabiate, closed by two lip-like flaps that are supported by chitinous 

jy rims. The analogy has been drawn by earlier authors, commenting on 

'^ the resemblance of the aperture of the Flustrellidae to the opening of 

an old fashioned clasp purse. Chitinous spines are present. 

^K5W/t^ Genus FLUSTRELLA Gray, 1848 

Zoaria incrusting, or rising in flattened fan-shaped projections. The 
zoaria are hispid, with many flexible chitinous spines, which vary in 
morphology and frequency with the species. The spines originate from 
kenozooecia. The aperture is bilabiate as described above. Genotype: 
Flustra hispida Fabricius, 1780. 

Flustrella cornicLilata (Smitt), 1871 
Plate 77, fig. 9 

Jlcyonidium corniculatum Smitt, 1871:1123. 
AlcyonidiuTn cervicornis Robertson, 1900-330. 
Alcyonidium spinifera O'Donoghue, 1923:192. 
Alcyonidium cervicorne, O'Donoghue, 1926:56. 
Flustrella corniculata, O'Donoghue, 1925:15. 

The zoaria are found in various modes of growth, depending upon 
the types of substrata. The shape varies from small cylindrical clavate 
colonies to large foliaceous flattened expansions. The color may range 
from pale tan to dark brown. Macroscopically, the zoaria have a coarse 
"fuzzy" appearance due to the presence of numerous chitinous spines. 
These spines arise from modified zooecia scattered abundantly among 
the functional zooecia. Most commonly the spines have four prongs. 
However, there are also spines bearing six prongs, and some with but 
one. The zooecia range in form from an elongated ovoid to hexagonal, 
usually with distinct lateral walls. The aperture is a narrow transverse 
slit. Occasionally specimens are found with the apertures slightly raised, 
at the summits of low papillae. The tentacle number, determined from 
sections, is 18. 

This species, described from cold European waters, has appeared in 
the Pacific literature under several different names. Robertson found it 
in the Alaskan collection of the Harriman Expedition, and O'Donoghue 
described it from the Vancouver Island region and Puget Sound. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 733 

Specimens in the Hancock collection are from off Point Barrow, 
Alaska, Arctic Research Laboratory, collector, G. E. MacGinitie; British 
Columbia; and Dillon Beach, Tomales Bay, California. Depth range, 
from intertidal to 36 fathoms. 

Flustrella gigantea Silen, 1947 
Plate 78, fig. 1 

Flustrella gigantea Silen, 1947:134. 

The zoaria are incrusting or arise into erect, flattened lobate, bilaminar 
expansions measuring 3 to 4.5 cm in height, and 0.5 to 1.0 cm in width. 
Macroscopically all the dark brown zoaria have a hirsute appearance 
due to the presence of branching chitinous spines. The zooecia are 
arranged in alternating series, varying in form from an irregular rec- 
tangle to an uneven hexagon j in length they range from 0.97 to 1.25 
mm, and in width from 0.70 to 0.83 mm. In younger portions of the 
zoaria the zooecia are distinct, but in the older areas the lateral zooecial 
walls are obscured by the pigmented cuticle. Each zooecium has a distal 
raised oral papilla with the bilabiate aperture at its summit. The hollow 
spines, arising from kenozooecia, are variable in morphology, and have 
a location pattern that is only moderately uniform. Distally, about the 
raised oral papilla on each zooecium, are 2 to 4 of the multibranched 
spines. The number of terminal prongs may vary from 9 to 21, the most 
frequent range being 11 to 14. Some spines, as well as having the normal 
numerous prongs, are modified so as to have one large grossly extended, 
thorn-like spike, giving the spine an over-all length of 1.38 to 2.05 mm. 
This spectacular form of the spine is scattered at random in generous 
quantity over the zoaria, from the growing tip to the most mature por- 
tions of the zoaria. Sections revealed the tentacle number to be 26. 

The specimens described by Silen were from the Bering Sea. The 
material in the Hancock collection is also from Arctic waters, off Point 
Barrow, Alaska, G. E. MacGinitie collector. Depth, 36 fathoms. 

Family Pherusellidae Soule, new family 

Zoaria incrusting or arising into flattened flabellate, bilaminar exten- 
sions. Aperture square or quadrangular, raised upon a stout tubular 
process. Prominent compound communication pores (multiporous sep- 
tulae), supported by heavy chitinous rings, connect adjacent zooecia, 
piercing the distal as well as the lateral walls. No spines present. Prior 
to this time the genus Pherusella has been placed under the family 



734 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

FlustrelHdae, but the morphological differences in the aperture, the 
presence of the prominent communication pores, and the lack of keno- 
zooecial spines warrant the separation of this genus into a distinct family. 

Genus PHERUSELLA Soule, 1951 

Zoaria coriaceous, incrusting, or arising from incrustations in branch- 
ing flabellate, flattened projections. The distal ends of the zooecia rise 
into prominent tubular processes, which bear the aperture. When the 
polypide is retracted, the apertures appear square to transversely quad- 
rangular in shape. The lateral walls and the distal walls are pierced by 
prominent multiporous septulae provided with heavily chitinized rims, 
apparently a unique character in the Ctenostomata. Genotype: Flustra 
tubulosa (Solander), 1786. The genus Pherusa Lamouroux, 1816, is 
preoccupied by Pherusa Oken, 1807. The name Pherusa had also been 
proposed by Leach, 1814, and Rafinesque, 1815. ,^ /. 

Pherusella brevituba Soule, 1951 
Plate 78, fig. 2 

Pherusella brevituba Soule, 1951:368. 

The chitinous zoaria are a light brown in color, leathery in appear- 
ance, and form prominent incrustations upon the holdfasts and blades 
of algae. When the zoaria are strictly incrusting, they are unilaminar, 
or they may form erect fan-like "fronds" that are bilaminar, back to 
back, where the zoarial growth exceeds the limits of the algae thalli. 

The zooecia are elongate with considerable variation in shape, from 
imperfectly rectangular to hexagonal, averaging about 0.80 mm in length 
and 0.40 mm in width. Normally the individual zooecia are distinct, 
clearly defined by the lateral walls. The zooecial walls are perforated 
by well marked compound interzooecial communication pores having an 
average diameter of 0.02 mm. The rims of the communication pores 
are strengthened by heavy chitinous rings. Within this ring are four 
minute perforations piercing a thin chitinous diaphragm. 

The distal portion of each zooecium is raised to form a short but 
prominent tubular process bearing the aperture. The upper extremity 
of this tubular process is square to transversely quadrangular in shape. 
The tentacles number 23. 

This species has been taken off Portuguese Bend, California ; collected 
in the intertidal zone at Punta Baja, Rosario, Lower California, by 
E. Y. Dawson ; and found on the holdfast of algae washed ashore near 
the Santa Barbara-San Luis Obispo county line, southern California. 
The range in depth is from intertidal to 8 fathoms. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 735 

Family GlaVOpOridae Soule, new family 

Zoaria erect, arising from a basal plate. Each zoarium is differentiated 
into two anatomically distinct portions, a capitulum composed of auto- 
zoids supported by an annulated peduncle composed of muscular keno- 
zoids. Zoaria may be solitary or in groups but are never compound. 
The aperture of each zooecium is similar to that found in the family 
Alcyonidiidae, and is usually located at the center of a small papillate 
process. The presence of two anatomically distinct regions within a 
zoarium, a situation not found elsewhere in the carnose families, justifies 
the proposal of a new family. 

Genus CLAVOPORA Busk, 1874 

Zoaria usually small, erect, coriaceous, clavate, arising from basal 
discs. As noted above, each zoarium has an annulated peduncle of 
muscular kenozooecia capable of bending and flexing the erect portion 
of the colony in any direction, and a capitulum of functional autozooecia 
capable of feeding and reproduction. The kenozooecia of the annulated 
peduncle are arranged in a series of rings, the central portion of the 
peduncle being a hollow fluid-filled tube. This tube forms a communica- 
tion between each feeding autozoid of the hollow capitulum and the 
muscular kenozooecia. Fluid containing dissolved nutriments and cel- 
lular elements may pass into the kenozooecia by means of minute simple 
pores (septulae) located in the internal zoid walls. The musculature of 
the kenozooecia consists of modified parietal muscles that run parallel 
to the long axis of the peduncle. Contraction of the muscles on one side, 
with reciprocal relaxation of the musculature of the opposite side, will 
bend the entire erect portion of the colony. In the capitulum, at the 
apex of the peduncle, the autozoids are densely packed. On the outer 
wall of the capitulum the cuticle is comparatively thick and leathery. 
The lateral walls and the internal walls of the zooecia, submerged within 
the body of the capitulum, are, in contrast, thin, lightly chitinized, and 
delicate in appearance. Genotype: Clavopora hystricis Busk, 1874. 

Glavopora occidentalis (Fewkes), 1889 
Plate 78, fig. 3 

Ascorhiza occidentalis Fewkes, 1889:1. 
Ascorhiza occidentalis, Robertson, 1902:106. 
Ascorhiza occidentalism O'Donoghue, 1923:192. 
Clavopora occidentalis, O'Donoghue, 1926:57. 



736 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

The zoaria are stalked, arising directly from small irregularly cylindri- 
cal adherent basal discs. The basal discs are firmly attached to the sub- 
strata; rocks, mollusk shells, or, as in the case of some of those in the 
Hancock collection, attached to colonies of the cheilostome bryozoan 
Discoporella umbellata (Defrance), 1823. The zoaria are a pale brown 
to light tan in color. The zoarial length is variable, ranging from 8 to 
5.8 cm. The zoaria may be solitary, or secondary zoaria may grow 
attached to the pedunculate portion of an older zoarium, where they 
have developed from settled larvae. 

Anatomically, a zoarium may be divided into two distinct sections, a 
peduncle composed of muscular kenozooecia, and a capitulum at the 
apex of the peduncle, composed of functioning autozooecia. The peduncle 
is cylindrical, stout, and strongly annulated in the older zoaria. Accord- 
ing to the figure in Fewkes' original description, the stalk is extremely 
slender. This is not the case with specimens in the Hancock collection, 
the peduncular portions of the mature zoaria having a diameter ranging 
from 0.50 to 0.75 mm. The capitulum, ranging in length from 2 mm 
to 3.5 cm, is an expanded ovoid structure, bulb-like in appearance with 
a coriaceous cuticle. It is composed of the functional autozooecia, closely 
united, somewhat indistinct, with the aperture located within the center 
of a low papillate process. The tentacle number, determined by means 
of serial sections, is 18. 

The specimens reported by Miss Robertson (1902) were dredged off 
Santa Catalina Island, California, while those recorded by O'Donoghue 
came from the vicinity of Vancouver Island. The specimens in the 
Hancock collection are from Hancock station 924-39, Socorro Island, 
Mexico; Guadalupe Island, Mexico, collector C. L. Hubbs; and Dillon 
Beach, California; in depths ranging from 17 to 46 fathoms. 

Division 2. Paludicellea Allman, 1856 

Zooecia connected by stolon-like tubular extensions that may or may 
not possess internodes separated by septulae. A zooecium may form a 
daughter zooecium by means of a bud produced near its distal extremity. 

Family Nolellidae Harmer, 1915 

"The Family Nolellidae is characterized by the great development of 
the peristomial part of the zooecium. This region is typically much 
elongated and its ectocyst frequently includes muddy particles. The 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 737 

adnate portion of the zooecium is represented by a delicate stolon-like 
tube and by the base of the peristome into which it usually passes 
abruptly, although it more rarely dilates gradually as it approaches this 
part. The branching is of the cruciform type. Gizzard absent." Harmer, 
1915:52. 

Genus NOLELLA Gosse, 1855 

Zooecia cylindrical, elongate, with considerable variation in size within 
the same zoarium. The proximal ends of the zooecia are prolonged, 
narrowed to form connecting tubular extensions. The cuticle may, on 
occasion, be covered by a very fine argillaceous coat. Genotype : Nolella 
stipata Gosse, 1855. 

Nolella stipata Gosse, 1855 
Plate 78, fig, 5 

Nolella stipata Gosse, 1855:35-36. 
Farrella gigantea Busk, 1856:93. 
Farrella dilatata, Hincks, 1860:279. 
Cylindroecium giganteum, Hincks, 1884:208. 
Cylindroecium papuense Busk, 1886:38. 
Cylindroecium giganteum, O'Donoghue, 1926:60. 

Zoaria with stolonal portion adhering to varied substrata ranging from 
hydroids and algae to eroded mollusk shells and cheilostomatous bryo- 
zoans. The zooecia are chitinous, erect, cylindrical. The cuticle is cov- 
ered with an extremely fine layer of silt, which does not, however, totally 
obscure the view of the polypide in alcoholic or wet-mount preparations. 
The zooecia are extremely variable in length, with mature specimens 
ranging in length from 0.90 to 3.80 mm, and in width from 0.17 to 
0.25 mm. The proximal portion of the zoid, the basal area, is expanded 
(dilated) forming a junction point for 2, 4, or even 6 of the stolons. 
The only stolon that is not set off from the basal dilation by a distinct 
diaphragm is the one from which the zoid arises. The degree of basal 
dilation seems to be correlated with the type of substratum. The speci- 
mens in the collection that are adherent to a soft substrate, such as the 
algae or the hydroids, have a much less prominent dilation than those 
adhering to a hard mollusk shell, where the proximal dilation is very 
great. (See Hincks, 1880:537, pi. 77, figs. 1 & 2, and pi. 79, figs. 1-3). 

This species is liberally represented in the cooler waters of the Atlantic 
on both the European side and the North American. On the Pacific 



738 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

coast of North America it has previously been reported by Hincks from 
the Queen Charlotte Islands, and by O'Donoghue from the vicinity of 
Vancouver Island, 

In the Hancock collection specimens are from Puget Sound, Wash- 
ington, Gulf of California, and the west coast of Lower California. 
Hancock stations, 650-37, San Francisco Island, Gulf of California, 
and 1714-49, Magdalena Bay, Lower California. Depth range from 
17 to 47 fathoms. 

Genus ANGUINELLA van Beneden, 1845 

Zoaria erect, branching irregularly. Zooecia cylindrical, arising from 
a small adnate proximal base. Zooecia bud directly from other zooecia. 
Genotype: Anguinella palmata van Beneden, 1844. 

Anguinella palmata van Beneden, 1845 
Plate 78, fig. 4 

Anguinella palmata van Beneden, 1845:34. 
Anguinella palmata, Osburn, 1912:253. 

Zoaria palmate, chitinous, opaque, brown in color, consisting of erect 
single stalks with zooecia branching irregularly to all sides. The zoarial 
length of the specimens in the Hancock collection ranges from 2.0 to 
3.1 cm. The zooecia are cylindrical, elongate, rounded distally, the 
aperture terminal. The cuticle is characteristically covered with a fine 
coat of silt, rendering examination of the polypide difficult even under 
optimum conditions of fixation and preservation. The zooecia bud directly 
from the sides of older mature zooecia. The polypides of the zooecia in 
the older basal and axial portions of the zoaria are suppressed, and these 
zooecia serve as support for the younger lateral and distal zooecia that 
are functional. 

No difference could be detected in the morphology of the Pacific speci- 
mens when compared with the Atlantic specimens collected at Beaufort, 
North Carolina, by R. C. Osburn, or those collected at New River, 
North Carolina, by A. S. Pearse. This is believed to be the first record 
of this genus and species from the Pacific Coast of North America. 
According to Hincks, 1880:540, it is moderately abundant in the waters 
about the British Isles and off the coast of Belgium and France. 

Hancock Stations: 277-34, Isabel Island, Mexico; 447-35, Panama 
City, Panama; 847-38, off Zorritos Light, Peru; 1449-42, Newport 
Harbor, and 2020-51, Seal Beach, southern California. Depth range, 
intertidal zone to 25 fathoms. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 739 

Division 3. Vesicularina Johnston, 1847 

The ctenostomes included within the limits of this grouping char- 
acteristically have relatively heavy, thickened, usually branching, septate 
stolons. The zooecia bud directly from the stolon. Polypide usually 
provided with a gizzard, or as Harmer, 1915:60, stated, "Gizzard 
present in most of the genera, perhaps in all," 

Family Vesiculariidae Johnston, 1838 

Zoaria erect or creeping, consisting of two types of zooecia, the keno- 
zooecia constituting the stolons, and the autozoids the feeding individ- 
uals. From within each internode of a stolon arise several zooecia, the 
arrangement being characteristic within the genera. 

Genus VESIGULARIA J. V. Thompson, 1830 

Zoaria erect, the main stolon or stolons supported on the substrate 
by a number of kenozooecial rhizoid-like runners. Zooecia ovoid to 
elongate cylindrical, distinct, arranged within an internode in a single 
series. Zooecia are contracted at the base, and the polypide is provided 
with a prominent gizzard. Genotype: Sertularia spinosa Linnaeus, 1758. 

Vesicularia fasciculata Soule new species 
Plate 78, fig. 6 

Diagnosis : Zoaria erect, unbranched, arising from a base supported 
by tubular, root-like, kenozooecial fibers. The main axis of the zoarium 
is composed of a series of 6 to 8 stout parallel or entwined stolons ad- 
herent to each other so as to form an elongated bundle. Zooecia elongate, 
cylindrical, arising from the stolons in a linear series, containing poly- 
pides each bearing 12 short tentacles and a prominent gizzard. 

Description: Of the three zoaria representing this species in the 
Hancock collection, the longest measured 2.80 cm in height, prior to 
the removal of portions for sectioning and for whole-mounts, while the 
shortest measured barely 0.6 cm. The remaining zoarium was in a very 
poor state of preservation. 

The zoaria arise in a single, non-branching axis of growth, from a 
base supported by kenozooecia in the form of tubular radiculate fibers. 
The main axis mentioned above consists of a series of 6 to 8 or more 
robust stolons adherent to and twisted about each other to form an 



740 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

elongate sheaf. Examination of a cross section of a stolonal sheaf from 
an older portion of the colony revealed stolons of uniform diameter, 
while cross sections made close to the growing tip of the colony disclosed 
one larger principal stolon surrounded by 4 to 6 secondary stolons of 
narrower diameter. An individual stolon does not as a rule traverse the 
entire length of a zoarium. One stolon will give rise to a second at a 
point located just below (proximally) a septum terminating an inter- 
node. The newly arisen stolon gains mature diameter at once and pro- 
ceeds distally paralleling its "parent" and the other stolons of the 
zoarium. The zooecia arise from the stolons, originating in a linear 
series within an internode in variable numbers. They are deciduous, the 
stolons characteristically marked with the scars of departed zooecia. 
The zooecia are constricted slightly at the point of fusion with the 
stolon. Morphologically they are elongate, cylindrical, ranging in length 
from 0.94 to 1.10 mm, and in width from 0.24 to 0.28 mm. The poly- 
pide contains a prominent gizzard. The tentacles are short, and 12 in 
number. 

Vesicularia fasciculata differs in two major aspects from the other 
species in the genus, having an unbranched zoarium, as compared to the 
branched zoaria of V. spinosa Linnaeus, V. papuensis Busk, and V. 
harmeri Silen, and it has 12 tentacles as compared to 8 tentacles in V. 
spinosa and V. papuensis. 

Holotype: U. S. N. M. no. 11053; Paratype, AHF no. 134. 

Repository: The United States National Museum, Washington, D. C. 

Paratype: The Allan Hancock Foundation, The University of South- 
ern California, Los Angeles, California. 

Type locality: Off Point Barrow, Alaska, 18 February 1950, depth 
162 feet, collector, G. E. MacGinitie. Also Point Barrow, Alaska, 
August 1, 1949, depth 321 feet, July 1, 1950, depth 118 feet, collector, 
G. E. MacGinitie. 

Genus AMATHIA Lamouroux, 1812 

Zoaria erect, stolons robust, stiff. Zooecia in biserial arrangement, 
forming a spiral within an internode. Polypide provided with a gizzard. 
Genotype: Sertularia lendigera Linnaeus, 1758. 

Amathia convoluta Lamouroux, 1816 
Plate 78, fig. 7 

Amathia convoluta Lamouroux, 1816:160. 
Amathia convoluta, Harmer, 1915:64. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 741 

The zoaria of this well known species are large, erect, prominent, 
light brown in color. The zooecia are arranged biserially, paired, form- 
ing a loose spiral that encircles the stolon within the limits of an inter- 
node. An internode is limited to one series of zooecia. The zooecia are 
completely connate along their entire length when the tentacles are 
completely retracted. The zooecia are of uniform length, ranging from 
0.71 to 0.74 mm. In width, the range is from 0.08 to 0.09 mm. The 
polypide has a gizzard. 

This species appears to be widely distributed, having been previously 
reported from European waters of the Atlantic and in North and South 
America from Chesapeake Bay to Santos Bay, Brazil. In the Pacific 
there have been several reports from the Australian region. This is the 
first report of its occurrence in the waters of the eastern Pacific. 

Hancock Stations: 133-34, Socorro Island, west of Mexico; 253-34, 
and 257-34, Port Culebra, Costa Rica; 265-34, Petatlan Bay, Mexico, 
and 486-35, Tenacatita Bay, Mexico. Depth, 5 to 20 fms. 

Amathia vidovici (Heller), 1867 
Plate 79, fig. 2 

Valkeria Vidovici Heller, 1867:128-129. 
Amathia vidovici, Osburn, 1940:340. 

Zoaria erect, tall, with elongate internodes. The zooecia are small, 
biserial, forming a spiral in the distal portion of the internode, leaving 
for the most part the proximal portion of the internode bare. Zooecia 
connate only at their point of origin and attachment to the stolon. Their 
length ranges from 0.32 to 0.41 mm. 

This species has not appeared in the literature as frequently as some 
of the other species of the genus Amathia. It was originally reported 
from the Adriatic Sea by Heller. On the Atlantic coast of North 
America it has been reported by Osburn and by Hutchins. Osburn also 
reported it from Puerto Rico. 

The specimens in the Hancock collection are from about 20 stations, 
ranging geographically from Santa Rosa Island, southern California 
(in the northern Channel Islands), to Ecuador and the Galapagos 
Islands. 

Amathia distans Busk, 1886 
Plate 79, fig. 1 

Amathia distans Busk, 1886:33. 
Amathia distans, O'Donoghue, 1925:16. 
Amathia distans, Osburn, 1940:339. 



742 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

The zoaria are comparatively small, low, straggling, with a moderately 
regular dichotomous mode of branching. The zooecia are found in bi- 
serial spirals that may, but usually do not, fill an internode, most fre- 
quently occupying only the distal portion. The zooecia are short, ranging 
in length from 0.35 to 0.46 mm, closely connate, except at the tips. This 
species differs from A. convoluta, whose zooecia are also connate, in its 
smaller size, its reptant habit, and in having the proximal half of the 
internode usually devoid of zooecia. 

A. distans has been reported previously from the South Atlantic by 
Busk, 1886:33; from Australian waters by MacGillivray, 1889:30; 
Java, Harmer, 1915:68; Puerto Rico by Osburn, 1940:339; and Puget 
Sound, O'Donoghue, 1925. 

The specimens in the Hancock collection (20 stations) range geo- 
graphically from Santa Rosa Island, southern California, to the Gulf 
of California. 

Genus ZOOBOTRYON Ehrenberg, 1831 

Zoaria loosely spreading, flaccid, not creeping, branching in an irregu- 
lar fashion. Zooecia ovoid, narrowed at the point of origin and attach- 
ment to the stolon. Polypide with a prominent gizzard. Genotype: 
Hydra verticillata delle Chiaje, 1828. 

Zoobotryon verticillatum (delle Chiaje), 1828 
Plate 79, fig. 3 

Hydra verticillata delle Chiaje, 1828:203. 

Zoobotryon pellucidus Ehrenberg, 1831: no pagination. 

Zoobotryon pellucidum, Osburn, 1940:341. 

The zoaria are flaccid, lavishly branching into tangled masses. The 
stolons are transparent, very flexible, only lightly chitinized, ranging in 
diameter from 0.40 to 0.70 mm. At intervals both the stolon and the 
zooecia may be partially obscured due to a deposit of silt. The zooecia 
are usually found arranged bilaterally along the stolons, but not in- 
frequently they occur in scattered clumps. The zooecia range in length 
from 0.36 to 0.48 mm, and in width from 0.12 to 0.17 mm; elongated- 
ovoid, rather narrow at the point of origin and attachment to the stolon, 
tapering to a bluntly square tip at the distal apertural orifice. The poly- 
pide is provided with a prominent gizzard. While this is the first direct 
description of this species from the Pacific coast of North America, Miss 
Alice Robertson, 1921:63, mentioned in her paper on the Bryozoa of 
the Bay of Bengal that she had seen this species in San Diego, California, 
and had received specimens from Hawaii. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 743 

According to Osburn, 1940:342, this species is circumtropical. It 
has been recovered from the warm waters of the Mediterranean, from 
Bermuda, Florida, Puerto Rico, Gulf of Mexico, and Brazil. 

Specimens in the Hancock collection are from San Diego, California, 
no further data given. 

Genus BOWERBANKIA Farre, 1837 

"Zooecia arising irregularly from an erect or creeping axis, commonly 
in definite groups. Tentacles 8-10. Gizzard present." Harmer, 1915:70. 
Genotype: Sertularia imbricata Adams, 1800. 

Bowerbankia imbricata (Adams), 1800 
Plate 79, fig. 4 

Sertularia imbricata Adams, 1800:11. 
Bowerbankia imbricata, Robertson, 1900:331. 
Bowerbankia imbricata, O'Donoghue, 1925:93. 

The zoaria form irregular tangled masses, with reptant stolons having 
a diameter ranging from 0.06 to 0.09 mm. The stolons are divided into 
internodes of variable length, separated by a diaphragm perforated by 
a single pore. The zooecia are elongate-tubular, straight or slightly 
curved, and have a square distal extremity. The proximal zooecial por- 
tion may be extended to form a short caudate process of one or two 
prongs. The zooecia are constricted at the point of origin on the stolon. 
The zooecial length of the eastern Pacific specimens ranges from 0.92 
to 1.15 mm. A gizzard is present. The tentacle number is 10, as de- 
termined from serial sections. 

This species appears to be well distributed in the cooler European 
waters. In the eastern Pacific, it has been previously reported from 
Alaska and Puget Sound. 

In the Hancock collection, the specimens of this species are from 
British Columbia, E. F. Ricketts, collector, no bathymetric data avail- 
able. 

Bowerbankia gracilis Leidy, 1855 
Plate 79, fig. 5 

Bowerbankia gracilis Leidy, 1855:142. 

Bowerbankia gracilis, O'Donoghue, 1923:192; 1925:93. 

Bowerbankia gracilis, Osburn, 1940:341. 



744 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

The zoaria consist of tangled gray masses of stolons and zooecia, 
repent, not erect, with extremely irregular branching. The zooecia are 
tubular, narrow, tapering slightly at both the distal and the proximal 
ends. The distal extremity is square in most cases. The polypide is pro- 
vided with a prominent gizzard, measuring between 0.08 and 0.09 mm 
in diameter. The zooecial length ranges from 1.02 to 1.52 mm. None 
of the zoaria had specimens of mature zoids with a measurement of less 
than 1.0 mm. As a rule, the specimens with a caudate appendage 
proximally were the longest. The zooecia are attached to a creeping 
stolon with or without a lateral extension. The zooecia may occur 
single, in pairs, or in dense clusters. The stolonal diameter is variable, 
ranging from 0.03 to 0.05 mm. The stolons have internodes of variable 
length, separated by diaphragms which are perforated by a single pore. 

In the eastern Pacific specimens in the collection, it was found that 
both caudate and non-caudate individuals occur within the same zoaria, 
with the non-caudate form predominant. No zoaria were found in which 
the caudate individuals occurred solely. 

Bowerbankia gracilis is a "cosmopolitan species," having been previ- 
ously reported from Greenland to Puerto Rico to Brazil. 

Specimens in the Hancock collection are from Puget Sound, Wash- 
ington; Dillon Beach, Tomales Bay, California, R. C. Osburn collector; 
Los Angeles Harbor; and the Gulf of California. All collections were 
made in the intertidal range. Hancock station, 510-36, Espiritu Santo 
Island, Gulf of California. 

Bowerbankia gracilis aggregata O'Donoghue, 1926 

Plate 79, fig. 6 

Bowerbankia gracilis var. aggregata O'Donoghue, 1926:58-60. 

The zoaria form dense tangled masses which completely obscure the 
substrata. The stolons, as in B. gracilis Leidy, have internodes of vari- 
able length, limited by diaphragms perforated by a single pore. The 
zooecia are very greatly elongated, ranging in length from 1.77 to 2.25 
mm. The tentacle number is 8. 

This variety was described by O'Donoghue from the Vancouver 
Island region. 

The specimens in the Hancock collection are from Point Barrow, 
Alaska, Arctic Research Laboratory, G. E. MacGinitie, collector; Puget 
Sound, Washington, J. L. Mohr, collector ; Dillon Beach, Tomales Bay, 
California, R. C. Osburn collector ; and Los Angeles harbor, California. 
The depths range from intertidal to 9 fathoms. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 745 

Division 4. Stolonifera Ehlers, 1876 

Zoaria with delicate creeping stolons, with occasional points of ex- 
pansion where a diaphragm occurs and either stolonal branches or zooecia 
may arise. A gizzard may or may not be present. 

Family Valkeriidae Hincks, 1877 

"Zooecia contracted below, deciduous, destitute of a membranous 
area." Hincks, 1880:551. 

Genus VALKERIA Fleming, 1823 

Zoaria repent, with creeping stolons. Zooecia ovoid to cylindrical, 
originating at the distal end of a short internode close to the diaphragm. 
No gizzard present. Genotype: Ser tularin uva Linnaeus, 1767. 

Valkeria tuberosa Heller, 1867 
Plate 79, fig. 7 

Valkeria tuberosa Heller, 1867:129. 
Valkeria tuberosa, Harmer, 1915:76. 

Zoarium stolonate, internodes of variable length, ranging from 0.52 
to 0.94 mm in length. At the internodes the stolon is expanded slightly, 
with lateral branches arising immediately distal to the diaphragm. Here 
the zooecia arise. The zooecia are small, ranging from 0.43 to 0.55 mm 
in length, and have a narrow wrinkled base 0.03 to 0.04 mm in width. 
Tentacles are 8 in number. Polypide lacking a gizzard. 

Previously reported from the Adriatic Sea, Red Sea, and Borneo. It 
has not been previously recorded from the eastern Pacific. 

The specimens in the Hancock collection are from Lower California, 
C. L. Hubbs, collector, no bathymetric data given. 

Genus AEVERRILLIA Marcus, 1941 

Zoaria creeping, minute. Stolons with short lateral peduncles to which 
the zooecia are attached. Polypide with a prominent gizzard. Geno- 
type: Biiskia setigera Hincks, 1887. 

Aeverrillia setigera (Hincks), 1887 
Plate 79, fig. 8 



Buskia setigera Hincks, 1887:127. 
Buskia setigera, Osburn, 1940:343. 
Aeverrillia setigera, Marcus, 1941 :74. 



746 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL, 14 

The zoaria are adherent to the substrate, minute, delicate, rather 
difficult to see without the aid of a lens. The zoaria consist of primary 
and secondary stolons, usually at right angles to each other, with the 
secondary stolons originating in pairs, one stolon on either side of the 
primary stolon, adhering closely to the substrate. The primary stolons 
are septate, divided into internodes. Septa are also found at the junction 
of the secondary or lateral stolons. The internodes are of variable length. 
The diameter of the stolons ranges from 0.02 to 0.05 mm. The zooecia 
arise in pairs from kenozooecia placed at each side of either a primary 
or secondary stolon. In a number of instances in the eastern Pacific 
material, the substrate was Amathia convoluta and Amathia vidovici, 
and thus did not permit paired zooecia to arise consistently. The zooecia 
range in length from 0.57 to 0.62 mm, and in width from 0.16 to 0.20 
mm. The basal portion of the zooecia is rounded, somewhat swollen, 
and usually bears 2 spine-like processes. Distally, the zooecia taper, 
and each bears upon the oral extremity 4 spine-bearing protuberances 
that encircle the aperture. A long setigerous collar may or may not 
project from the aperture. The polypide contains a prominent gizzard. 
The tentacles number 8. 

Aeverrillia setigera, previously unreported from the eastern Pacific, is 
a semitropical species, having been reported from the warmer waters of 
the southwest Pacific (Ceylon, New Guinea, Gulf of Bengal, China 
Sea), from the waters off Puerto Rico, from the Suez Canal, from 
Brazil, and as far north as Long Island Sound, Connecticut, and New 
Bedford and Woods Hole, Massachusetts, on the Atlantic Coast of 
North America. 

Hancock Stations: 133-34, Socorro Island, west of Mexico; 445-35, 
Panama City, Panama ; and 847-38, southwest of Zorritos Light, Peru. 
Depth, intertidal to 35 fms. 

Family Buskiidae Hincks, 1880 

"Zooecia contracted below, not continuous with the creeping stolon, 
with an aperture on the ventral surface." Hincks, 1880:531. In the 
light of present knowledge of this family, the above diagnosis must be 
modified : Zooecia contracted proximally, arising directly from the 
stolon, aperture terminal. 

Genus BUSKIA Alder, 1857 

Zoaria repent or erect, stolonate. Zooecia arising directly from the 
stolon. Polypide with a prominent gizzard. Genotype: Buskia nitens 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 747 

Buskia nitens Alder, 1857 
Plate 80, fig. 1 

Buskia nitens Alder, 1857:156. 
Buskia nitens, Hincks, 1884:208. 
Cylindroecium repens O'Donoghue, 1923:192. 
Buskia nitensj O'Donoghue, 1926:60. 

Zoaria minute, repent, inconspicuous. Stolons thin, thread-like, sub- 
divided by septa into internodes. Zooecia very small, ranging in length 
from 0.31 to 0.50 mm. The zooecia arise directly from the stolons and 
in most cases, but not invariably, the proximal one-third of a zoid is 
adherent to the stolon and the substrate, w^ith the distal two-thirds free. 
In some cases the entire zoid arises directly away from the stolon and 
is free in its entirety. Proximally, some zooecia exhibit one or two pairs 
of short thorn-like protuberances. Distally, some zooecia show a short 
setigerous collar projecting from the aperture. 

This species is evidently well distributed in both warm and cool marine 
waters, but because of its minute size is easily overlooked. It has been 
reported from England, Brazil, Puerto Rico, and British Columbia. 

Hancock Stations: 277-34, Isabel Island, Mexico, and 1407-42, Coos 
County, Oregon, intertidal to 25 fms. 

Buskia seriata Soule, new species 
Plate 80, fig. 2 

Diagnosis: Zoaria erect, branching. Stolons robust, septate, bearing 
clusters of short stocky zooecia arranged in a paired linear series, alter- 
nate, and arising directly from the stolon. Zooecia wrinkled distally 
and may exhibit a setigerous collar protruding from the aperture. The 
polypide contains a prominent gizzard. The tentacle number is 8. 

Description: The zoaria are large, branching, erect, macroscopically 
bearing a superficial resemblance to specimens of the genus Amathia. 
The stolons are robust, septate, with internodes of variable length, rang- 
ing from 0.90 to 1.30 mm, and in width from 0.07 to 0.09 mm. The 
zooecia, which are arranged in clusters, arise from the stolon in an 
irregular alternate, paired linear series. These zooecial clusters may 
contain from 5 to 14 short, stout zooecia, with 11 occurring most fre- 
quently. On the younger stolonal branches only 1 or 2 developing 
zooecia may be in evidence. Invariably, there is but a single zooecial 
cluster to each stolonal internode. The zooecia arise directly from the 
stolon. The proximal portion of the zooecium is constricted, but the 
body proper rarely adheres to the stolon or substrate for any distance, 



748 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

although individuals of this type do occur. The zooecia are small, rang- 
ing in length from 0.35 to 0.45 mm, and in width from 0.11 to 0.13 
mm proximally, and 0.09 to 0.11 mm distally. All of the zooecia have 
a broad rounded proximal portion, w^here 1 or 2 small, pointed, spine- 
like protuberances may appear. The zooecia gradually taper distally, 
where, shortly beyond the point midway between the two extremities, 
they become wrinkled transversely. Some of the zooecia exhibit a 
setigerous collar protruding from the aperture. The polypide is pro- 
vided with a large and prominent gizzard. The tentacle number is 8, 
as determined from examination of serial sections. 

Although erect, Buskia seriate has comparatively short zooecia, differ- 
ing from B. socialis which, according to Marcus, has zooecia measuring 
0.75 mm in length. Being erect, it is easily distinguished from the 
reptant B. nitens with its minute creeping zooecia. 

Holotype: AHFno. 133. 

Repository: Allan Hancock Foundation, The University of Southern 
California, Lx)s Angeles, California. 

Type locality: Galapagos Islands, N. Seymour Island, January 16, 
1931, tidepools. 

Additional distribution: Hancock station 1111-40, February 14, 1940, 
San Lorenzo Channel, Gulf of California, west coast, 24°21'55'"N, 
110°15'15''W, depth 6-13 fathoms, bottom sandy, shells. 

Family Triticellidae G. O. Sars, 1874 

"Stolon delicate without free branches, zooecia erect with a long 
slender base-like pedicel, with a flattened membranous frontal area and 
without spines at the distal end around the oral aperture." Osburn, 
1944:26. 

Genus TRITIGELLA Dalyell, 1848 

Zoaria with creeping stolons. Zooecia pedicellate, erect, attached to 
the stolon by means of a movable joint. Zooecia are elongate, ovoid, 
with a membranous frontal area. No gizzard. Genotype: Triticella 
ftava Dalyell, 1848. 

Triticella pedicellata (Alder), 1857 

Plate 80, fig. 4 

Farrella pedicillata Alder, 1857:158. 

Triticella pedicellata, O'Donoghue, 1923:193, 1926:61. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 749 

The zoaria are stolonate, creeping. The zooecia may be clustered, 
arising from short lateral internodes of the stolon. The pedicel is slender, 
measuring between 0.03 and 0.04 mm in diameter near the base, becom- 
ing slightly enlarged toward the zooecia proper. At the point of junction 
with the zooecia, the pedicel becomes transversely wrinkled. The zooecia 
are elongate, elliptical, ranging in length from 0.80 to 1.25 mm, and in 
width from 0.16 to 0.23 mm. A flattened frontal area extends the full 
length of the zooecia proper. The polypide does not have a gizzard. 
The tentacles number 12. 

There is some variation in the length of the pedicels, but they are 
usually about twice the length of the zooecia. The longest measured 
2.40 mm, which when combined with its zooecial measurement of 1.19 
mm, gave a total height of 3.59 mm. 

This species has been previously reported in the cool waters of Eng- 
land and northern Europe. In the eastern Pacific it has been previously 
reported from the Vancouver Island region. 

The specimens in the Hancock collection are from Canoe Bay, Alaska, 
and Union, Washington. The depth of the Alaskan specimens is un- 
known; those from Washington were collected at 10 fathoms. 

Triticella elongata (Osburn), 1912 
Plate 80, fig. 5 

Hippuraria elongata Osburn, 1912:256. 
Triticella elongata^ Osburn, 1944:26. 

Zoaria living in the gill chambers of the pea crab, Scleroplax granulate 
Rathbun. The adnate stolons give rise to erect zooecia, which are usually 
paired in clusters. The zooecia arise from short internodes, rather than 
directly from the stolons. The zooecia range in length from 0.90 to 
1.80 mm, including the pedicel. The length of the zoids proper ranges 
between 0.50 and 0.90 mm. In width, the zooecia range from 0.18 to 
0.24 mm. The polypide has 16 to 18 tentacles. 

Osburn, 1944:26, reports this species from Chesapeake Bay, and its 
geographical distribution on the Atlantic coast of North America from 
Vineyard Sound, Massachusetts to Beaufort, North Carolina. It has 
not been previously reported from the eastern Pacific. 

Specimens in the Hancock collection are from Elkhorn Slough, 
California, collector R. I. Smith. No depth data available. Found on 
Scleroplax granulata Rathbun. 



750 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Genus FARRELLA Ehrenberg, 1838 

Zoaria with reptant stolons. Zooecia arising within the internodes 
along the entire length of the stolon. No gizzard. Genotype : Lagenella 
repens Farre, 1837. 

Farrella elongata (P. J. van Beneden), 1845 
Plate 80, fig. 3 

Laguncula elongata van Beneden, 1845a :26. 
Triticella tegeticula O'Donoghue, 1923:193. 

The zoaria are comprised of creeping stolons, which may in the older 
colonies form a dense mat-like network upon the substrate. In the young 
zoaria the zooecia are seen to arise from the creeping stolons within the 
internodes, budding forth laterally and vertically without apparent order. 
In the older colonies, this lack of arrangement packs the pedunculate 
zooecia closely together. The zooecia are robust, elongate, ovoid to sub- 
cylindrical in form, and are situated at the end of a long peduncle that 
may attain a length of from 0.50 to 0.80 mm. The peduncle is trans- 
versely wrinkled and gradually widens into the zooecia proper without 
a definite joint. The overall length of the zooecia ranges from 1.16 to 
1.35 mm, while the width varies from 0.32 to 0.39 mm. The diameter 
of the primary stolon is about 0.03 mm. The polypide lacks a gizzard. 
The tentacle number of 16 was determined from serial sections. 

A striking feature of this species is the morphology of the zooecial 
aperture. The aperture deviates from the typical rounded or squared 
form of the stolonate ctenostomes in that it is bilabiate. Close examina- 
tion of the apertural area will reveal a pair of lip-like structures, each 
reinforced by a thin but definite chitinous rim. These "lips" are found 
only in the zooecia that have reached maturity. Farre, 1837:403, in his 
work on Lagenella repens (Farre), 1837, a very closely related species, 
considered the labiate structure to be opercula. 

Marcus, 1926:50, using Farrella repens (which according to Farre, 
van Beneden, and Hincks, has only 12 tentacles) and experimentally 
causing unfavorable conditions, produced the "Form" elongata and at 
the same time reduced the tentacle number. Marcus overlooked the 
fact that van Beneden reported 16 tentacles for Farrella elongata, the 
same number that was found in the Pacific specimens. 

Triticella tegeticula O'Donoghue, 1923, is here suggested as a pos- 
sible synonym of F. elongata, because of its habit of growth as well as 
the morphology of the zooecia. Although O'Donoghue failed to mention 



NO. 3 OSBURN: EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 751 

in his description the presence of a bilabiate aperture, the figure of his 
specimen strongly suggests the bilabiate type of structure. 

Farrella elongate appears to be well represented in the cooler European 
waters in the vicinity of England and the Adriatic Sea. 

Hancock Station, 1489-42, Coos County, Oregon; also taken at 
Tomales Bay, California, by R. J. Menzies. Intertidal. 

Division 5. Terebriporina Soule, new division 

Ctenostomes with stolonate zoaria that are characteristically imbedded 
within the calcareous shells of living or dead mollusks, brachiopods, or 
barnacles, their presence marked by the apertural openings of the zoids 
appearing at the surface of the shell. The stolons are thin, thread-like, 
septate. 

The three families that are placed under the Terebriporina cannot 
be readily differentiated by the pattern of the tracings that appear upon 
the surface of the shell in which the zoaria are immersed. The only 
means of postive identification of the families and the genera is examina- 
tion of zoaria that have been removed from shells by decalcification. 
The identification of species involves not only the study of zoid anatomy, 
but serial sections of the autozoids to determine definitely the tentacle 
number. The family Penetrantiidae can be anatomically identified by 
its zoaria with primary and secondary stolons, its typical gonozoids, and 
the operculated autozoids. Terebriporidae, also with primary and sec- 
ondary stolons, lacks the operculated autozoids, while Immergentiidae, 
having autozoids with typical ctenostomatous apertures, has zoaria devoid 
of true gonozoids, the colonies being composed of a series of zoids joined 
by stolon-like tubules that are direct extensions of the zoids. 

Family Terebriporidae d'Orbigny, 1847 

Zoaria burrowing, stolonate. The zooecia are connected to the pri- 
mary or main stolons by means of short secondary stolons emitted from 
near the distal zooecial extremity. 

Genus TEREBRIPORA d'Orbigny, 1847 

Zoaria stolonate, consisting of primary stolons joined to the zoids by 
secondary stolons, with the point of union being nearly midway between 
the distal and proximal extremities, but always closer to the distal end. 
Polypide provided with a gizzard. Genotype: Terebripora rmnosa 
d'Orbigny, 1847. 



752 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Terebripora comma Soule, 1950 
Plate 80, fig. 6 

Terebripora comma Soule, 1950:380. 

The zoaria have successive zoids alternately placed to the right and 
left of the primary stolon at the end of a short secondary stolon. The 
short lateral stolon has a septum at the junction point where the stolon 
meets the zoid. The secondary stolons enter the zoids about midway 
between the distal and proximal extremities of the zoids, but always 
nearer to the distal end. Two types of zoids are in evidence, the auto- 
zoids (feeding individuals) and zoids modified for reproduction that 
may be termed gonozoids for convenience. Anatomically, the autozoids 
are typical of the usual ctenostomate type. The polypide bears a promi- 
nent globular gizzard. In length the autozoids range from 0.32 to 0.35 
mm, and in width from 0.06 to 0.08 mm. The tentacles are short, and 
are 8 in number. The autozoids are elongate, with the distal aperture 
bluntly square, and the proximal portion terminating in a tapering 
rounded point. No brown bodies were seen. The reproductive zooecia 
or gonozoids have a prominent, large, oval embryo measuring about 
0.06 mm in diameter at maturity. In length, the gonozoids range from 
0.29 to 0.33 mm, and in width from 0.07 to 0.08 mm. 

Hancock Station, 1937-50, Anacapa Island, southern California. Also 
off Newport, southern California. Depth, 18 to 43 fms. 

Family Immergentiidae Silen, 1946 

Zoaria with only primary stolons, that are not stolons in the strict 
sense, being prolongations of the zoids. These stolons arise directly from 
the distal tips of the preceding zoids, and connect one zoid with another 
in a series. 

Genus IMMERGENTIA Silen, 1946 

Zoaria imbedded in the shells of both living and dead mollusks. The 
stolonal connections between zoids are slender, thread-like, originating 
at the distal ends of the zooecia. The zoids are small in size, elongate, 
narrow. The proximal end may be bluntly rounded or tapered to a 
narrow point. The distal tip bears a centrally placed square shaped 
aperture. No zoid specifically modified for reproduction is known to 
occur. Genotype: Immergentia calif ornica Silen, 1946. 



NO. 3 OSBURN: EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 753 

Immergentia californica Silen, 1946 

Plate 80, fig. 7 

Immergentia californica Silen, 1946:6. 
Immergentia californica, Soule, 1950:364. 

The specimens of /. californica in the Hancock collection are identical 
in all important respects with the paratype material generously donated 
by Dr. Lars Silen. The zoaria have the zoids arranged in straight rows, 
with lateral rows branching to the sides at rather irregular intervals. 
In length the zoids range between 0.32 and 0.34 mm, and in width the 
range is from 0.08 to 0.09 mm. The tentacle number is 10 as determined 
by serial sections. 

This species was originally described by Silen from material collected 
at Pacific Grove, California. 

Specimens in the Hancock collection are from San Pedro and Portu- 
guese Bend, southern California. All intertidal. 

Family Pcnetrantiidae Silen, 1946 

Zoaria with septate stolons. Zoids joined to the main stolons by 
means of short lateral stolons entering the zoids near the distal ex- 
tremity. Zoids have a double cuticle and are provided with an operculum. 
The polypide has a gizzard. Reproductive zoids, the gonozoids, have 
rudimentary polypides and bear large ovoid embryo chambers. 

Genus PENETRANTIA Silen, 1946 

Zoaria are imbedded in the shells of both living and dead mollusks 
and cirripeds. The zoids are connected by thin septate stolons. From 
a primary stolon a short thin lateral branch enters the zoid laterally at 
the distal end. A zoid modified for reproduction, the gonozoid, is present. 
The zoids are operculated. The polypide is provided with a gizzard. 
Genotype: Penetrantia densa Silen, 1946. 

Penetrantia densa Silen, 1946 
Plate 80, fig. 8 

Penetrantia densa Silen, 1946:2. 
Penetrantia densa, Soule, 1950:360. 

The zoaria characteristically have the zooecial openings crowded 
closely together. These openings in the molluscan shells vary in shape 
from circular to strongly oval. The primary stolons are serrated upon 



754 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

their upper surfaces. From a primary branch short lateral branches 
extend to the zoids, entering them at the distal end. The autozoids 
(feeding individuals) are usually straight, or only slightly curved. In 
length they range from 0.47 to 0.55 mm, and in width they vary from 
0.09 to 0.11 mm. Infrequently, an autozoid with a sharply pointed 
rather than a bluntly rounded proximal end will be found. The ten- 
tacles are 12 in number, determined from serial sections. The repro- 
ductive zoid, the gonozoid, is usually as long as but may be slightly 
shorter than the autozoid. Its proximal portion, extending below the 
embryo chamber, is long, thin and slightly curved in the direction of the 
embryo chamber. The embryo chamber is globular, giving the gonozoid 
a "pot-bellied" appearance. The tentacle number of the gonozoid is 8, 
but only in immature gonozoids will they be found, where the polypide 
has not completely degenerated. 

The specimens described in the original report by Silen were collected 
from South Africa and the Cape of Good Hope, and there was one 
"doubtful" specimen from Panama. 

The specimens in the Hancock collection are from numerous localities 
from San Pedro to La Jolla, southern California, all intertidal. 

Penetrantia concharum Silen, 1946 
Plate 80, fig. 9 

Penetrantia concharum Silen, 1946:5. 
Penetrantia concharum, Soule, 1950:360. 

The zoids of the colonies are well spaced, without the crowding noted 
in P. densa. The openings in the shell are well defined, reniform in 
shape. The autozoids are straight, slender, with the proximal extremity 
tapering to a point. The autozoids range in length from 0.46 to 0.54 
mm, and in width from 0.08 to 0.10 mm. The tentacles number 10, as 
determined from sections. The gonozoids are comparatively rare. The 
proximal extremity of the gonozoids is straight and visibly thicker than 
the gonozoid of P. densa. 

Penetrantia concharum was found by Silen to occur in numerous 
localities in Sweden and Norway. 

The specimens in the Hancock collection are from several localities 
ranging from San Pedro to La Jolla in southern California and south- 
ward to Rosarito, Lower California, Mexico (Hancock station 1597- 
47). All are intertidal. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 755 

Penetrantia sileni Soule, 1950 
Plate 80, fig. 10 

Penetrantia sileni Soule, 1950:361. 

The openings in the scaphopod shell made by this species vary con- 
siderably in shape, from a simple circular to a highly exaggerated reni- 
form appearance. The stolon is thin, not serrated on its upper surface, 
and is in general circular in cross section. The zoids are placed close to 
each other but are not crowded. The autozoid ranges in length from 
0.35 to 0.36 mm, and in width from 0.07 to 0.08 mm. The autozoids 
may be slightly curved, and they may have a pointed rather than a 
rounded proximal extremity. The tentacle number, as determined from 
serial sections, is 11. The mature gonozoid has a very characteristic 
morphology. It is little more than one-half the length of the autozoid, 
ranging from 0.19 to 0.20 mm in length. The narrowed proximal portion 
barely reaches below the swollen embryo chamber. 

Specimens in the Hancock collection are from off the San Benito 
Islands, Mexico, Hancock station 1010-39, 28°12'05''N, 115°33'45"W. 
The depth range is from 71 to 86 fathoms. It is as yet known only 
from the eastern Pacific. 



756 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

REFERENCES 
1. Papers dealing only with Pacific Coast Bryozoa 

Fewkes, J, W. 

1889. A Preliminary notice of a Stalked Bryozoon (Ascorhiza occidentalis). 
Ann. and Mag. Nat. Hist. ser. 6, vol. 3, pp. 1-6, pi. 1. 

HiNCKS, T. 

1884. Report on the Polyzoa of the Queen Charlotte Islands. Ann. and Mag. 
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O'DoNOGHUE, C. H. and Elsie O'Donoghue 

1923. A Preliminary List of Bryozoa (Polyzoa) from the Vancouver Island 
Region. Contr. Canad. Biol. Fish, new ser., vol. 1, pp. 143-201 (1-59), 
pis. 1-4. 

1925. Notes on certain Bryozoa in the collection of the University of Wash- 
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pp. 15-23. 

1925a. List of Bryozoa from the vicinity of Puget Sound. Wash. [State] Univ. 
Puget Sound Biol. Sta. Pubs. vol. 5, pp. 91-108. 

1926. A Second List of the Bryozoa (Polyzoa) from the Vancouver Island 
Region. Contr. Canad. Biol. Fish, new ser., vol. 3, pp. 49-131 (1-85), 
pis. 1-5. 

Robertson, Alice 

1900. Papers from the Harriman Alaska Expedition. VI. The Bryozoa. Proc. 

Wash. Acad. Sci. vol. 2, pp. 315-340, pis. 19-21. 
1902. Some observations on Ascorhiza occidentalis Fewkes, and related 
Alcyonidia. Proc. Calif. Acad. Sci. ser. 3, Zool. vol. 3, pp. 99-108, pi. 14. 

Soule, J. D. 

1950. Penetrantiidae and Immergentiidae from the Pacific (Bryozoa: Cten- 
ostomata). Trans. Amer. Micros. Soc. vol. 69, pp. 359-367, pis. 1-2. 

1950a. A new species of Terebripora from the Pacific (Bryozoa: Ctenosto- 
mata). Jour. Wash. Acad. Sci. vol. 40, pp. 378-381, figs. 1-3. 

1951. Two new species of incrusting ctenostomatous Bryozoa from the Pa- 
cific. Jour. Wash. Acad. Sci. vol. 41, pp. 367-370, figs. 1-4. 

2. General References 

Adams, J. 

1800. Descriptions of some Marine Animals found on the Coast of Wales. 
Trans. Linn. Soc. London, vol. 5, pp. 7-13, pi. 2. 

Alder, J. 

1857. A catalogue of the Zoophytes of Northumberland and Durham. Trans. 
Tyneside Nat. Field Club. vol. 3, pp. 93-162, pis. 3-9. 

BeNEDEN, J. P. VAN 

1845. Recherches sur I'anatomie, la physiologie et le developpement des 
Bryozoaires qui habitent la cote d'Ostende. Nouv. Mem. Brussels Acad. 
Roy. de Belg. vol. 18, pp. 1-44, pis. 1-5. 

1845a. Recherches sur I'organisation des Laguncula. Nouv. Mem. Brussels 
Acad. Roy. de Belg. vol. 18, pp. 1-29, pis. 1-3. 

Busk, G. 

1852. An account of the Polyzoa, and Sertularian Zoophytes, collected . . . 

on the coasts of Australia and the Louisiade Archipelago. In Mac- 

Gillivray, J. Narrative of the Voyage of H. M. S. Rattlesnake . . . 

during . . . 1846-50. Appendix. London, vol. 1 (4), pp. 343-388. 
1856. Zoophytology. Quart. Jour. Micros. Sci. new ser., vol. 4, pp. 93-96, 

pis. 5-6. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 757 

1874. On Clavopora hystricis — a new Polyzoon belonging to the family 
Halcyonelleae. Quart. Jour. Micros. Sci. new sen, vol, 14, pp. 261-262, 
pi. 9. 

1886. Report on the Polyzoa collected by H. M. S. Challenger during the 
years 1873-1876. Part II. — The Cyclostomata, Ctenostomata and Pedi- 
cellina. In Report on the Scientific Results of the Voyage of H. M. S. 
Challenger during the years 1873-76. Zoology, vol. 17 (3), pp. i-viii, 
1-47, pis. 1-10. 

DELLE ChIAJE, S. 

1828. Memorie sulla storia e notomia degli Animali senza vertebre del 
Regno di Napoli. vol. 3. 

Ehrenberg, C. G. 

1831. Symbolae Physicae . . . Pars Zoologica. Anim. evert, exclus. insect 
Berolini. pis. 1-10 with descr. letterpress. 

Farre, a. 

1837. Observations on the Minute Structure of some of the higher forms of 
Polypi, with views of a more natural arrangement of the Class. Phil. 
Trans. Roy. Soc. London, vol. 127 (2), pp. 387-426, pis. 20-27. 

Fleming, J. 

1828. Zoophyta. In his A History of British Animals. Edinburgh, London, 
pp. 505-554. 

GossE, P. H. 

1855. Notes on some new or little known Marine Animals. (Fascis II.) 
Ann. and Mag. Nat. Hist, sen 2, vol. 16, pp. 27-36, pis. 3-4. 

Harmer, S. F. 

1915. The Polyzoa of the Siboga Expedition. Part 1, Entoprocta, Ctenosto- 
mata and Cyclostomata. In Siboga-Expeditie. Leyden. Mon. 28a, pp. 
1-180, pis. 1-12. 

Hassall, a. H. 

1841. Description of two new genera of Irish Zooph)^es. Ann. and Mag. 
Nat. Hist. vol. 7, pp. 483-486. 

Heller, C. 

1867. Die Bryozoen des adriatischen Meeres. Verhandl. Zool.-Bot. Gesell. 
Wien. vol. 17, pp. 77-136, pis. 1-6. 

HiNCKS, T. 

1860. Descrintions of new Polyzoa from Ireland. Quart. Joun Micros. Sci. 

vol. 8,*pp. 275-285, pis. 30-31. 
1880. A History of the British Marine Polyzoa. London. 2v. 

1887. The Polyzoa of the Adriatic: a supplement to Prof. Heller's "Die 
Bryozoen des adriatischen Meeres." Ann. and Mag. Nat. Hist, sen 5, 
vol. 19, pp. 302-316, pi. 9. 

1887a. On the Polyzoa and Hydroida of the Mergui Archipelago. Joun Linn. 
Soc. London. Zool. vol. 21, pp. 121-134, pi. 12, figs. 1-13. 

Johnston, G. 

1838. A History of the British Zoophytes. London, Edinburgh. 341p., 44 pis. 
1847. Idem, i'^d. 2). London. 2v. 

Lamouroux, J. V. F. 

1816. Histoire des polypiers coralligenes flexibles, vulgairement nommes 
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758 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Leidy, J. 

1855. Contributions towards a knowledge of the Marine Invertebrate Fauna 
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Marcus, E. 

1926. Beobachtungen und Versuche an lebenden Meeresbryozoen. Zool. 
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Letr. vol. 22 (Zool. 5), pp. 3-208, pis. 1-18. 

OSBURN, R. C. 

1912. The Bryozoa of the Woods Hole region. Bui. U. S. Bur. Fisheries, vol. 

30, pp. 205-266, pis. 18-31. 
1933. Bryozoa of the Mount Desert Region. Repr. from the Biological Survey 

of the Mount Desert Region. Philadelphia. 67p., 15 pis. 
1936. Bryozoa collected in the American Arctic by Captain R. A. Bartlett. 

Jour. Wash. Acad. Sci. vol. 26, pp. 538-543, 1 fig. 
1940. Bryozoa of Porto Rico with a Resume of the West Indian Bryozoan 

Fauna. In Scientific Survey of Porto Rico and the Virgin Islands. 

New York. vol. 16 (3), pp. 321-486, pis. 1-9. 
1944. A survey of the Bryozoa of Chesapeake Bay. Chesapeake Biol. Lab. 

Contr. no. 63, pp. 1-55, pis. 1-5, text figs. 1-28. 

Robertson, Alice 

1921. Report on a Collection of Bryozoa from the Bay of Bengal and other 
Eastern Seas. Rec. Indian Mus. vol. 22, pp. 33-65. 

ROGICK, M. D. and H. Croasdale 

1949. Studies on Marine Bryozoa III, Bryozoa associated with Algae. Biol. 
Bui. vol. 96, pp. 32-69, figs. 1-10. 

SiLEN, L. 

1942. Carnosa and Stolonifera (Bryozoa) collected by Prof. Dr. Sixten 
Bock's expedition to Japan and the Bonin Islands, 1914. Arkiv for 
Zool. vol. 34A (8), pp. 1-33, text figs. 1-24. 

1946. On two new Groups of Bryozoa living in Shells of Molluscs. Arkiv 
for Zool. vol. 38B (1), pp. 1-7, text figs. 1-12. 

1947. On the spines of Flustrella (Bryozoa). Zool. Bidr. Uppsala, vol, 25, 
pp. 134-140, pi. 1, text figs. 1-7. 

Smitt, F. a. 

1871. Kritisk forteckning ofver Skandinaviens Hafs-bryozoer. Ofversigt af 
Svenska Vetensk. Akad. Forhandl. vol. 28, pp. 1115-1134, pis. 20-21. 

Thompson, J. V. 

1830. On Polyzoa, a new animal discovered as an inhabitant of some Zoo- 
phites — with a description of the newly instituted genera of Pedi- 
cellaria and Vesicularia, and their Species. In his Zoological Researches 
IV. Memoir 5, pp. 89-102, pis. 1-3. 

ZiRPOLO, G. 

1924. Zoobotryon pellucidum Ehrbg. = Z. verticillatum (delle Chiaje). 
Bol. Soc. Nat. Napoli. vol. 36 (Commun. verb.), pp. 6-7. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 759 

Phylum Entoprogta 

By Raymond C. Osburn, Ph.D., D.Sc. 

Phylum ENTOPROGTA Nitsche, 1869 

Subphylum Entoprocta, various authors. 
Phylum Entoprocta, Hatschek, 1888. 
Phylum Calyssozoa Clark, 1921 :19 and 23. 
Phylum Kamptozoa Cori, 1929:5. 
Phylum Entoprocta, Hyman, 1951:521. 

Until recent years this group has generally been considered a sub- 
phylum or subclass of the Bryozoa, although Hatschek as early as 1888 
(Text-book of Zoology) separated the Phylum Entoprocta. In 1921 
Clark recognized the differences and proposed the Phylum Calyssozoa. 
Again Cori in 1929, though he was familiar with the name Clark had 
suggested, thought it necessary to rename the group as Phylum Kamp- 
tozoa. These writers considered the Entoprocta to be much simpler than 
the Ectoprocta and separated them widely. Marcus (1939:208-288) 
raised objections to this wide separation and gave some very cogent rea- 
sons for retaining the group as a subphylum of the Bryozoa. Recently 
Dr. Libbie Hyman, in the third volume of "The Invertebrates" (1951 : 
521-554) again separates the group as a phylum. She assigns it to a 
place much lower in the scale and retains the name Entoprocta for the 
very good reason that it is unnecessary to invent a new name when a 
perfectly good one already exists. Hyman's discussion is a very satisfac- 
tory analysis of the knowledge of the Entoprocta, and it seems unneces- 
sary to argue the matter further at present. 

Whether or not the entoprocts are closely allied to the ectoprocts and 
wherever they may eventually be placed in the taxonomic scale, it hap- 
pens that the only taxonomists who have paid much attention to them 
are the bryozoologists, and for this reason the species which are known 
from the Pacific coast are appended to the Bryozoa. The list is small 
as the species are not numerous, and the littoral species are rarely found 
among the dredgings. 

The Entoprocta are stalked, with naked heads or calyces (polypides), 
the tentacles rolled inward instead of being withdrawn into a zooecium, 
and the anal opening is within the ring of tentacles instead of outside 
as it is in the Ectoprocta. 

The family Loxosomatidae is unique in that the individuals live singly 
and do not form colonies, and they live as epizoites on other animals, 
usually on sponges, worms, other Bryozoans, etc. The only other family, 
Pedicellinidae, is colonial and is represented among our material by 4 



760 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Family Loxosomatidae Hincks, 1880 

The individuals are not colonial but live singly, attached to some other 
animal by a muscular sucking disc at the base of the stalk. They are 
unchitinized, flexible and capable of bending in any direction, and some 
of them, at least, are capable of moving about and re-attaching them- 
selves. They produce buds from the sides of the calyx, but these sever 
their connections when their growth is complete and live singly there- 
after. They are all small, some of them microscopic in size. There are 
two genera, Loxosoma Keferstein and Loxocalyx Mortensen, depending 
on whether the foot-gland disappears after attachment or remains func- 
tional. 

Genus LOXOSOMA Keferstein, 1863 

Loxosoma davenporti Nickerson, 1898 

Loxosoma Davenporiii Nickerson, 1898:220. 
Loxosoma Davenporti Nickerson, 1899:368. 
Loxosoma davenporti Nickerson, 1901:351. 
Loxosoma davenporti, Osburn, 1912:212. 

This species has been noted only once on the Pacific coast, by 
O'Donoghue at low tide in Hammond Bay Lagoon, British Columbia. 
It is a commensal in worm tubes. 

The entire animal is about 2 mm long, somewhat vase-shaped, the 
pedicel cylindrical and about as long as the calyx into which it merges 
gradually; foot-gland wanting in the adult; lophophore with 18 to 30 
tentacles, the body somewhat narrowed below the lophophore ; usually 
with a pair of flask-shaped glandular organs on the ventral side of the 
body near the lower end of the stomach. The species was originally ob- 
tained by Nickerson and later by Osburn in the Woods Hole region, 
Massachusetts. 

? Loxosoma sp. 

A small species which was epizoic on an annelid worm at Point Bar- 
row, Alaska, and on account of the preservation is unidentifiable even 
to the genus. The calyx expands gradually from the pedicel upward; 
width of calyx at the upper end 0.18 to 0.22 mm, height 0.33 to 0.40 
mm, length of pedicel about 0.40 mm. Apparently there is no foot gland, 
and the tentacles cannot be counted. 



NO. 3 OSB URN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 761 

Genus LOXOGALYX Mortensen, 1911 

In this genus the foot-gland is evident and functional throughout life. 
Genotype Loxosoma raja Schmidt, 1876. 

Loxocalyx sp. 

Three individuals attached to the parapodia of an annelid vi^orm, 
Gattyana cirrosa, from Puget Sound. The calyx expands abruptly, and 
its base is rounded; w^idth of cal}^ 0.26 mm, height 0.33 mm, length 
of pedicel 0.40 mm. The foot-gland is present, but the tentacle number 
cannot be estimated. 

Family Pedicellinidae Johnston, 1847 

Colonial, the individuals erect from a creeping segmented stolon, the 
pedicels and stolon more or less chitinized. In the genera Myosoma and 
Pedicellina the pedicel is muscular and flexible, w^ithout a special muscu- 
lar enlargement at the base, w^hile in Barentsia and Coriella the pedicel 
is more chitinized and bears an enlarged, barrel-shaped muscular en- 
largement at its base. In Barentsia daughter individuals are often pro- 
duced by budding from joints of the pedicel. 

Key to Genera of Pedicellinidae 

1. Pedicels not heavily chitinized, muscular and flexible, rising 

directly from the stolon vv^ithout a specialized muscular base . 2 
Pedicels usually stiff and inflexible, with an enlarged cylin- 
drical, somew^hat barrel-shaped base 3 

2. The lophophore (tentacle crov^^n) is diagonally placed on the 

ventral side; the pedicel thick and with strong diagonal 

muscles Myosoma 

The lophophore is terminal, the pedicel narrower and without 

diagonal muscles Pedicellina 

3. Individuals always arising from short stolon internodes ; erect 

branches formed by the fusion of stolons ; pedicels never 

jointed Coriella 

Individuals arising from short stolon internodes, or from the 
sides of the pedicels; erect branches sometimes formed by 
enlarged pedicels; the stolons do not fuse to form erect 
branches Barentsia 



762 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Genus MYOSOMA Robertson, 1900 

"Zoarium with stolon composed partly of successive polypide-bearing 
segments and partly of alternate non-polypide-bearing segments; both 
stalk and calyx muscular, the muscle fibers continuous from one into 
the other; polypide oblique." (Robertson, 1900:324). Genotype, M. 
spinosa Robertson, 1900:324. 

The pedicel is unusually thick, flexible, and has a conspicuous set of 
diagonal muscles in addition to longitudinal ones. The stolon is entirely 
adnate. 

Myosoma spinosa Robertson, 1900 
Plate 82, fig. 1 

Myosoma spinosa Robertson, 1900 :324. 

The creeping stolon gives rise to branches vi^hich sometimes unite side 
by side but more frequently ramify and cross each other, forming a 
rather close mat; all of the internodes are comparatively short, ranging 
from 0.20 to 0.70 mm, the infertile internodes 0.08 to 0.12 mm in 
diameter, the fertile ones somewhat thicker, especially near the origin of 
zooecial buds. The zooecia arise from the internodes without any special 
differentiation, and even the muscles extend down into the stolon. The 
pedicels are exceptionally large, as much as 0.26 mm in diameter, nar- 
rowing upward to about 0.13 mm below the calyx, varying greatly in 
height to as much as 2.50 mm. They are highly muscular, with the unique 
diagonal muscles in addition to the longitudinal ones. 

The calyx is moderately large, ovoid in shape, averaging about 0.65 
mm long by 0.40 mm in width, the dorsal side more curved ; the lopho- 
phore is diagonally placed on the shorter ventral side; the tentacle 
number is apparently 16. Chitinous spines, varying in number, are 
present on the dorsal side of the calyx and also on the stolon. 

Robertson listed the species from Dillon Beach, Tomales Bay, and 
from Fort Point and San Diego, California. 

Hancock collections, numerous specimens from Dillon Beach, Cali- 
fornia, the type locality. Dr. R. J. Menzies, collector. The writer has 
also taken it at Newport Bay and at La Jolla, California. It is a littoral 
species and, as far as known, occurs only on the coast of California. 

Genus PEDIGELLINA M. Sars, 1835 

The zoarium is entirely adnate, consisting of fertile and infertile 
internodes, the latter rather regularly 0.40 mm in length ; lateral stolons 
sometimes cause the zoarium to cover a considerable area. The pedicel 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 763 

is large, as much as 0.25 mm in diameter at the base and about half as 
wide as its distal end, as much as 2 mm long but usually much shorter; 
flexible and with longitudinal muscles only which do not extend into 
the calyx. The expanded caljoi is cup-shaped, with the tentacle crown 
transverse at the top. Spines present on the stalk and calyx, or wanting 
on one or both of them. Genotype, Brachionus cernuus Pallas, 1771. 

Pedicellina cernua (Pallas), 1771 
Plate 82, fig. 2 

Brachionus cernuus Pallas, 1771 :57. 
Pedicellina americana Leidy, 1855:143. 
Pedicellina nutans, Robertson, 1900:332. 
Pedicellina echinata, Robertson, 1900:344. 
Pedicellina cernua, O'Donoghue, 1926:7. 

The stolon is slender, more or less transparent, branching, consisting 
of an irregular succession of fertile and infertile internodes, the fertile 
ones shorter than the others, as a rule, and somewhat swollen. The 
pedicel is broadest at the base, about 0.25 mm in diameter but often 
smaller, diminishing in size upward to about half of the basal width; 
thin walled and flexible, with longitudinal muscles which do not enter 
the calyx. Usually there is a slight constriction between the pedicel and 
calyx. The pedicel may be 2 mm or more in length but is usually shorter. 

The calyx is cup-shaped with a well-marked gibbosity on the dorsal 
side, varying greatly in size, in our largest specimens about 0.55 mm 
long by 0.40 mm in diameter. The lophophore is terminal and transverse, 
with the tentacle number varying from 14 to 24. 

Spines are sometimes present on the stalk and also on the calyx, and 
this feature has led to the erection of several species names, P. echinata 
M. Sars, 1835:5, P. glabra Hincks, 1880:565 and P. hirsuta Jullien, 
1888:13. However there is so much variation in the presence and dis- 
tribution of the spines that these must be considered merely nominal 
varieties. In our material, which extends from British Columbia to 
southern California, most of the zooecia are without spines, while a 
few spines occur occasionally even on the same colonies with bare zooecia. 

The species is cosmopolitan. Robertson listed it as P. nutans ( ?) 
from Yakutat, Alaska, and as echinata from Tomales Bay, California, 
and O'Donoghue recorded it from several localities in British Columbia. 

Hancock collections: Five Fingers, British Columbia; Tomales Bay 
and Lime Point, California; and the writer has obtained it at Newport 
Harbor and La Jolla, southern California. It is a littoral species, usually 
found on the piles of docks or at low tide. 



764 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Genus BARENTSIA Hincks, 1880 

Pedicellinopsis Hincks, 1884. 
Ascopodaria Busk, 1886. 
Gonopodaria Ehlers, 1900. 
Arthropodaria Ehlers, 1900. 

This genus is distinguished by the presence of a large, barrel-shaped, 
muscular swelling at the base of the pedicel (a character which it also 
shares with Coriella Kluge) and by the adnate stolons which never fuse 
to form erect branches (as they do in Coriella). The pedicel is narrow 
above the muscular base and is usually well chitinized and stiff; joints 
occasionally appear in most of the species and in some of them are 
characteristic. In most of the species the pedicel appears to be more or 
less perforated, but the outer chitinous layer is complete and there are 
no pores ; the cavities are limited to the internal layer. The calyx is more 
or less ovoid or vase-shaped, the lophophore terminal and transverse, and 
at its base the calyx is separated from the chitinized pedicel by a short 
flexible portion. The muscular base permits swinging back and forth, 
and the short flexible base of the calyx also permits the head to move 
freely on the pedicel. In at least one species, B. laxa Kirkpatrick, the 
pedicel is only slightly chitinized, provided with longitudinal muscles, 
and can be bent or looped in any direction. Genotype, B. bulbosa Hincks, 
1880a :285. 

The differences in the jointing and branching of the pedicel caused 
the erection of several other generic names, but there is so much variation 
in this character that it is often not even of specific value. Some species 
{e.g. discreta Busk) very rarely are jointed, some others {ramosa Robert- 
son) are very regularly jointed and branched; sometimes the joints are 
enlarged and muscular, or the muscular enlargement may be wanting. 
These characters have some value for the determination of species but 
are scarcely valid for the separation of genera. 

Key TO Species of Barentsia 

1. Upper half of the basal internode and the whole of the short 

second internode thin-walled and flexible ; base of the lower 

internode well chitinized subrigida 

Stalks with chitinized walls, not flexible 2 

2. Stalks never branched, usually without joints 3 

Stalks more or less jointed and branched, the nodes often 

much enlarged 5 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 765 

3. Stalk short, basal bulb as long as or longer than the following 

internode, calyx large robusta 

Stalk usually much longer than the basal bulb 4 

4. Small species, seldom 2 mm in height, the internode little or 

not at all "perforated," joints simple, rare gracilis 

Taller species, 2 to 4 mm high, internode very thickly "per- 
forated," joints simple, rare discrete 

5. Zoarium very large, reaching a height of 5 centimeters, pro- 

fusely branched ; muscular bulbs of varying size, sometimes 

gigantic; high Arctic gorbunovi 

Branches few, arising from enlarged stem nodes, basal bulbs 

of one size 6 

6. Tall slender species, long internodes, nodes moderately swollen ; 

branching occasionally at the nodes, the basal bulb short 

and ovate geniculata 

Stouter species, internodes shorter, the nodes much enlarged, 
nearly every stalk bears one or more branches; the inter- 
nodes with a few "perforations" ramosa 

Barentsia gracilis (M. Sars), 1835 

Plate 82, fig. 3 

Pedicellina gracilis M. Sars, 1835:6. 
Pedicellina gracilis, Hincks, 1884:208. 
Ascopodaria gracilis, Robertson, 1900:345. 
Gonypodaria nodosa, O'Donoghue, 1923:5. 
Barentsia gracilis nodosa, O'Donoghue, 1926:7. 
Barentsia gracilis, Marcus, 1938:8. 

A small, delicate species, usually less than 1.0 mm in height. The 
stolon is creeping, usually among hydroids and other bryozoans. The 
basal bulb is of moderate size; the stalk short and lightly chitinized 
with few or no "perforations." The largest calyx in our specimens 
measures 0.25 mm high by 0.18 mm wide; the pedicel 0.95 mm high; 
the basal bulb 0.35 mm high by 0.12 mm in diameter. The pedicel 
usually bears no "perforations" but a few may be present. The joints, 
which mark the variety nodosa Lomas, 1886, are rare in our material, 
and they are only slightly enlarged. 

Cosmopolitan. Recorded by Hincks and by O'Donoghue from a num- 
ber of localities in British Columbia, and by Robertson from Lime Point 
and San Pedro, southern California. 



766 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Hancock Stations: 1274-41, Point Hueneme, 30 fms, and 1292-41, 
Santa Rosa Island, 28 fms, southern California. Also collected by the 
writer at low tide, Corona del Mar, southern California. 

Barentsia discreta (Busk), 1886 
Plate 82, fig. 8 

Ascopodaria discreta Busk, 1886:44. 
Pedicellina australis Jullien, 1888:13. 
Barentsia timida Verrill, 1900:594. 
Barentsia discreta, Osburn, 1912:214; 1940:327. 
Barentsia discreta, Harmer, 1915:29. 
Barentsia discreta, Marcus, 1937:15. 
Ascopodaria misakiensis Oka, 1890:234. 
Barentsia misakiensis. Oka, 1895:76. 

The zoarium consists of a slender creeping stolon and erect slender 
zoids which may reach a total height of 3 mm, but usually range between 
1.0 and 2 mm. The variation in height is almost entirely in the length 
of the slender stalk. The muscular basal bulb is moderately large, 
averaging about 0.50 mm high by 0.18 mm in width; the slender stalk 
from less than 1.0 mm to more than 2 mm; the calyx in our largest 
specimens measures 0.78 mm in height by 0.60 mm in width, but it is 
often much smaller. The slender stalk is straight, widening slightly and 
gradually upward, and is especially characterized by the large number 
of "perforations" of the inner layer of the stalk. The stalk is rarely 
jointed, but simple joints sometimes occur to the number of 2 or 3. 
Harmer states that the tentacle number is "about 20-24," but in smaller 
calyxes the number may be as few as 14. 

It is a widely distributed species, occurring around the world in 
tropical and temperate waters. On the Atlantic coast it occurs from 
Cape Cod to Brazil, and Jullien described his Pedicellina australis from 
Cape Horn. Oka had it from near Tokyo, Japan, and Harmer from 
the East Indies. It has not been previously known from the Pacific 
coast, except for Waters' record at Magellanes, Chili. 

Hancock Stations: 391-35, Lobos de Afuera Islands, Peru, 6°55'40"S, 
shore collection; 401-35, Mantua Bay, Ecuador, 1 fm; 1217-40, Point 
Fermin, California, shore collection, 33°42'30'"N, and 1232-41, off San 
Pedro Breakwater, 17 fms, California. Also on piles of docks at Corona 
del Mar (R. C. Osburn) and Upper Newport Bay, shallow water, 
(J. D. Soule), southern California. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 767 

The known range on the Pacific coast is from 33°42'30"N, southern 
California, to Ecuador, Peru and Cape Horn. It appears to be a species 
of shallow waters, though Busk described it from Tristan de Cunha 
Island at a depth of 100-150 fms. 

Barentsia ramosa (Robertson), 1900 
Plate 82, Figs. S and 6 

Gonypodaria ramosa Robertson, 1900:337. 
Gonypodaria ramosa, O'Donoghue, 1923:6. 
Barentsia ramosa, O'Donoghue, 1926:8. 

The stolon is creeping and adherent, becoming heavily chitinized and 
brown, 0.07 mm in width. From this arise erect, jointed, and branched 
sub-colonies, with 3 or 4 joints in series, and these usually give rise to 
1 or 2 branches which in turn may develop secondary branches. The 
internodes vary much in length, from 0.65 to 2.60 mm; in width they 
measure 0.06 mm at the base and 0.10 to 0.13 mm near the tips. The 
muscular nodes vary in length from 0.24 to 0.30 mm and in width from 
0.15 to 0.20 mm. The basal bulb is short, 0.30 to 0.35 mm long by 0.20 
to 0.26 mm wide. The calyx is short and wide, the height, excluding 
the tentacles, 0.35 to 0.50 mm and the width 0.40 to 0.55 mm. Tentacle 
number, 16-20. The total height of the tallest sub-colonies is about 5 mm. 

Conspicuous features of the internodes are the heavy chitinization of 
the wall, the brownish color, and the conspicuous "pores," which are 
larger than in B. discrete; also the sub-colonies appear to be more rigid 
than in other members of the genus. 

The species was described by Robertson from Pacific Grove (Monte- 
rey Bay), California, and recorded also from Fort Point and Land's 
End, California and from Channel Rocks, Puget Sound. O'Donoghue 
recorded it from several localities in British Columbia. 

Hancock Collections: Carmel Cove, south of Monterey Bay, Cali- 
fornia, on the stems of the hydroid Garveia. It is a littoral species, but 
O'Donoghue dredged it at 20 fms. 

Barentsia gorbunovi Kluge, 1946 

Plate 82, figs. 10-12 

Barentsia gorbunovi Kluge, 1946:153 and 157. 

This is a remarkable species growing in bushy form to a height of 5 
cm or more. There is an adherent stolon, 0.10 to 0.20 mm in width, 
from which arise very complex sub-colonies. At the bases of these there 
are gigantic muscular bulbs of the usual shape, but with a height of 1.0 



768 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

to 2 mm and a diameter of as much as 0.80 mm. In other parts of the 
sub-colony the bulbs are much smaller, from 0.40 to 0.65 mm high and 
from 0.13 to 0.26 mm in diameter. 

The erect branches should be homologous with the stalks of other 
barentsias, but they differ in bearing a series of zoids throughout their 
length without joints or septa; they appear like erect stolons, except for 
the large muscular basal bulb. These erect stems bear branches, all in 
the same plane, up to the fifth generation of zoids. The unbranched 
internodes are comparatively short, usually less than 1.0 mm and about 
0.08 mm in diameter. The calyx is similar to those of other barentsias, 
0.40 to 0.50 mm high by 0.35 to 0.40 mm in width. 

The unusual features of this species are the large size and complexity 
of the colony, the nature of the branching, the absence of nodes, and the 
variation in the size of the basal bulbs. 

Known only from Kluge's record (Drifting Ice Expedition in the 
central Arctic Ocean in the Ice-breaking Str. "G. Sedov," 1937-40). 

Point Barrow, Alaska, Arctic Research Laboratory, 246 feet, G. E. 
MacGinitie, collector, several colonies. 

J ^^^ 

^ *.l— *- Barentsia robusta new species 

Plate 82, fig. 7 

The stolon adnate, ramifying, 0.08 mm wide. The individual zooecia 
are short-stalked with large calyces, reaching a total height of about 2 
mm. The stalks are disproportionate, as the basal bulbs are longer than 
the narrow internodes. The basal muscular bulbs are unusually long 
and measure rather regularly 0.90 mm in height by 0.18 mm in diameter. 
The internode is 0.70 to 0.78 mm long; at the base it varies from 0.06 
to 0.08 mm wide and gradually enlarges to 0.11 to 0.13 mm in diameter 
at the top ; the internode wall is moderately chitinized and its inner 
layer is punctured by scattering conspicuous "pores," similar to those 
of B. discreta but larger and much fewer in number. The calyx is large, 
the height to the base of the tentacle ring as much as 0.65 mm, and its 
width varying from 0.52 to 0.65 mm, cup-shaped, widest at the top and 
the base rounded, the dorsal side curved, the ventral side much straighten 
The tentacles cannot be counted accurately but apparently they number 
about 20 to 24. 

The larger calyces all contain embryos in August. The stout appear- 
ance of the zooecia, all about the same height, the large calyces (nearly 
one third of the total height) and the very long and comparatively 
slender basal bulbs which are consistently longer than the internodes, 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 769 

distinguish the species. It was collected by the writer in 1902 and has 
remained unidentified in his collection for the past 50 years awaiting 
a proper place for publication. 

Type, AHFno. 131. 

Type locality, near Port Renfrew, Vancouver Island, at the site of 
the former University of Minnesota Biological Station ; from the rocky 
wall of a deep tide-pool, R. C. Osburn, collector. 

Barentsia geniculata Harmer 

Plate 82, fig. 4 

Barentsia geniculata Harmer, 1915:33. 

1 Ascopodaria macropus, Robertson, 1900:345. 

The creeping stolon is thick-walled and brown, 0.06 or 0.07 mm in 
width; the fertile internodes short, 0.30 to 0.40 mm long, the infertile 
ones varying greatly in length to as much as 1.30 mm. The zoids are 
tall, reaching a total length of 5.20 mm, the stalk with 1 to 4 bulbous 
muscular joints. The stalk internodes are slender, only 0.04 mm at the 
base and enlarging slightly upward, with a few "pores" ; the basal stalk 
internodes are long, from 1.50 to 1.70 mm, the later ones somewhat 
graduated in length, the shortest terminal one only 0.45 mm. The stalk 
nodes are enlarged, muscular and bulbous, the swelling about 0.20 mm 
long by 0.15 mm in width. The basal bulb is short and wide, about 
0.40 mm high by 0.30 mm in diameter, coarsely wrinkled. The calyx 
is 0.30 mm high by 0.25 mm wide. The tentacle number cannot be 
determined accurately but they are numerous, at least 20. 

One stalk is branched twice, at the first node and again at the first 
node of a branch. This feature resembles B. ramosa, but the nodes are 
much smaller and the internodes more slender, while the basal bulb is 
strikingly different. 

This is probably the Ascopodaria macropus of Robertson, for which 
she gives no description, but states that it occurs at San Pedro, southern 
California. I believe it to be the B. geniculata of Harmer, as it agrees 
in the thick wall of the internodes, the form of the joints, the small 
number of "pores" ("tubercles," Harmer), the form of the muscular 
base and the size and form of the calyx. It differs in the longer inter- 
nodes and in the rare formation of branches at the internodes. 

Harmer described the species from the East Indies (Siboga Expedi- 
tion). 

Hancock Station 1292-41, near Santa Rosa Island, southern Cali- 
fornia, 33°53'30"N, 120°W, at 28 fms, two fragments. 



770 



ALLAN HANCOCK PACIFIC EXPEDITIONS 



VOL. 14 



/^A. 



•,i^ 



f-?' 



Barentsia subrigida new species 
Plate 82, fig. 9 

A large species, reaching a height of more than 5 mm ; it is especially 
characterized by the stalk which possesses regularly two internodes; the 
basal one is very elongate, 1.30 to more than 4 mm; the basal half, more 
or less, has the wall chitinized and rigid while the upper part remains 
thin-walled, muscular and flexible; this is followed by a somewhat ex- 
panded muscular joint, and above this is a short thin-walled and wrinkled 
flexible internode, only 0.40 to 0.55 mm in length. When completely 
developed, the top of the basal internode and the base of the terminal 
internode are both enlarged to about twice the width of the stem with 
a short muscular section between them; the terminal one has a definite 
diaphragm just above the node, similar to the one at the base of the 
first internode. Very old basal internodes may become chitinized for 
the full length, but I have seen only two such internodes. In any case 
the short terminal internode is always transparent and flexible. The 
basal internode usually bears a few "perforations," scattered and gen- 
erally disappearing entirely on the upper half. 

The basal muscular bulb varies greatly in height, from 0.25 to 0.55 
mm, probably depending on the nature of contraction, and the width 
is about 0.15 mm. 

The calyx is large, averaging about 0.35 mm high to the base of the 
tentacle crown, by 0.40 mm in width ; the largest calyx measures 0.50 
mm high by 0.52 mm wide; the dorsal side is more rounded than the 
frontal side, and the base is broadly rounded and attached by a rounded 
bulb to the upper internode. The tentacle number cannot be counted 
accurately but it appears to be at least 20. 

The very regular disparity in the length of the two internodes, the 
thin-walled flexible upper ends of the basal internodes and the constant 
thin-walled, muscular, short terminal internodes, together with the large 
size, apparently mark this as a hitherto unknown species. It is possible 
/that the Ascopodaria macropus of Robertson may belong here instead 
of under B. geniculata. The only other known species with flexible inter- 
nodes is B. laxa Kirkpatrick, which has no joints in the stalk, no "per- 
/forations," and the stalk is not heavily chitinized basally. 

Type, AHF no. 132. 

Type locality, Hancock Station 1274-41, three and one-half miles 
south of Hueneme, southern California, 28°23'20''N, 115°ir52"W, 
at 55 fms, two colonies. Another specimen in the collection is labeled 
simply "California." 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 771 

Genus CORIELLA Kluge, 1946 

Stolon adnate and creeping, but giving off clusters of erect stolons 
which fuse into complex sub-colonies. The zoids arise from the fertile 
internodes of the erect stolons in large numbers on all sides of the com- 
plex branch. The zoids are simple and unbranched, with the usual 
muscular basal bulbs, and the stalks are provided with scattering "pores" 
similar to those of some species of Barentsia. 

Genotype, C. stolonata Kluge, 1946:155. 

The genus appears to be similar to Barentsia in all essential characters 
except the fusion of erect stolons to form branches. 

Goriella stolonata Kluge, 1946 
Plate 82, figs. 13 and 14 

Coriella stolonata Kluge, 1946:155 and 157. 

The complex erect branch varies greatly from 3 to 10 or 12 stolons, 
to a height of more than 1 cm in our specimens. The calyx is large, 
reaching 0.80 mm high by 0.65 mm in width ; the stalk attains a length 
of 2 mm and a diameter of 0.06 to 0.08 mm, with rather numerous 
"pores"; the basal bulb measures 0.40 to 0.45 mm in height by 0.16 to 
0.22 mm wide; it arises from a cup-shaped enlargement which is set at 
an angle on the side of a fertile stolon internode. The total height of 
the tallest zoid is 3.25 mm, but they are usually much shorter. The 
tentacle number, according to Kluge, is 22 to 24. 

The erect clustered stolons distinguish this species from all members 
of the Pedicellinidae. 

The only record is that by Kluge (Drifting Ice Expedition in the 
central Arctic Ocean in the Ice-breaking Str. "G. Sedov," 1937-40). 

Point Barrow, Alaska, Arctic Research Laboratory, 295 feet, G. E. 
MacGinitie, collector. 



772 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

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Sars, M. 

1835. Beskrivelser og iagttagelser over nogle maerkelige eller nye i havet 
ved den Bergenske kyst levende dyr af Polypernes, Acalephernes, 
Radiaternes, Annelidernes og Molluskernes classer . . . Bergen, xii, 
81p., 15 pis. 

Verrill, a. E. 

1900. Additions to the Tunicata and Molluscoidea of the Bermudas. Trans. 
Conn. Acad. Sci. vol. 10, pp. 588-594. 



774 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Addenda 

The following species in Ectoprocta-Cheilostomata have come to the 
author's attention since Parts 1 and 2 were prepared for publication. 
Also the ovicell of Pachyegis brunnea (Hincks), hitherto unknown, is 
figured (plate 81, fig. 11), and Cheilopora praelucida (Hincks), dis- 
cussed on page 465 of Part 2, is illustrated on plate 81, fig. 12. All but 
one of these are from the last collections made at Point Barrow, Alaska, 
by Prof, and Mrs. G. E. MacGinitie. Their persistent efforts for two 
seasons resulted in 104 species from a very limited area around the 
Arctic Research Laboratory, which add greatly to our knowledge of 
circumpolar distribution of the Bryozoa. 

No doubt careful collecting all along the American coasts, in special 
habitats and especially in deeper waters, will add many more species to 
the long list already recorded. 

Gheilostomata 

Anasga 

Membranipora annae, new name 

Acanthodesia serrata (Hincks), Hastings, 1930:707 (not M. membra- 

nacea, form serrata Hincks, 1882:469) 
Membranipora hastingsae Osburn, 1950:29 (preoc. by M. (Electra) 

hasting sae Marcus, 1940) 

Dr. Anna B. Hastings misidentified this species with the M. serrata 
of Hincks (1882:469, which was already preoccupied by the M. serrata 
of MacGillivray (1868:131). [See under M. serrilamella, new name. 
Part 1, p. 22] The present author renamed it hastingsae (Part 1, p. 
29), overlooking the fact that Marcus (1940) had already used this 
name for another species. Dr. Marcus (in litt. Aug. 27, 1950) kindly 
called my attention to the error: "In Danmarks Fauna, 1940, Electra 
is treated as a subgenus of Membranipora, and M. (Electra) hastingsae 
is dealt with. So the specific name is preoccupied and your species will 
have to receive a new name." 

I have taken the liberty of using Dr. Hastings' given name for this 
very attractive little species. 

Hincksina gothica new species 
Plate 81, fig. 1 

Zoarium unilaminar, encrusting on shells, stones and bryozoans, white 
to light brown. Our largest colony, rounded and about 25 mm in 



NO. 3 OSBURN: EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 775 

diameter, spreads across a frond of Porella compressa, the edges extend- 
ing beyond the side of the frond but remaining unilaminar. The zooecia 
are large, 0.65 to 1.0 mm or more in length by 0.45 to 0.60 mm in 
width, arranged quincuncially. In the infertile zooecia the opesia occu- 
pies practically all of the frontal area; the side walls are very thin and 
finely granulated. Large interzooecial avicularia are scattered over the 
zoarium, usually at the beginning of the new series of zooecia. The 
chamber is nearly as long as the normal zooecia, 0.50 to 0.65 mm, but 
narrower, about 0.40 mm in width. The mandible, 0.35 to 0.50 mm 
long by 0.20 to 0.26 mm wide, has the shape of a gothic arch, the sides 
parallel and the pointed tip strongly decurved to fit inside the recurved 
tip of the rostrum. Minute avicularia (mandible 0.06 to 0.10 mm 
long) are situated at the distal zooecial comers and often there are one 
or two additional ones, with the mandible directed forward instead of 
backward as in H. nigrans; occasionally there are similar small avicularia 
on the side walls and in these the position of the mandible is reversed, 
pointing backward. Near the center of the colony there are 5 or 6 short 
stout spines and one or two of these may be present on the zooecia over 
the whole zoarium. 

The endozooecial ovicells are similar to those of nigrans, but the cover- 
ing transverse ridges are m.uch less developed and there is always a pair 
of the small avicularia distally placed on either side of the ovicell with 
the mandible directed forward. 

The species is evidently related to H. nigrans, but the differences in 
the avicularia and the cover of the ovicell and the presence of spines 
appear sufficient to give it specific rank. H. nigrans also has large inter- 
zooecial avicularia, but they are short with a short triangular mandible. 

Type, U. S. Nat. Mus. no. 11048. 

Type locality. Point Barrow, Alaska, 453 feet. Numerous colonies 
were obtained by Prof. G. E. MacGinitie at depths ranging from 216 to 
522 feet. 

Amphiblestrum trifolium (S. Wood), 1850 
Plate 81, fig. 2 

Flustra trifolium S. Wood, 1850:20. 
Membranipora solida Packard, 1867:272. 
Membranipora Flemingii var. solida, Verrill, 1879:29. 
Me?nbranipora trifolium, Hincks, 1880:167. 
Membranipora trifolium, Whiteaves, 1901:97. 
Membranipora trifolium, Osburn, 1912:279. 
Amphiblestrum trifolium, Osburn, 1923:9; 1932:9; 1933:26. 



776 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

Zoarium encrusting on stones and shells, the ectocyst sometimes with 
brown pigment. The zooecia are arranged more or less regularly in 
quincunx, length 0.60 to 0.80 mm, width 0.35 to 0.60 mm, varying 
much in form, separated by slightly elevated thin margins which are 
decorated with coarse granules. The most striking feature is the thick, 
finely granulated cryptocyst which covers the proximal half or more of 
the frontal area and extends narrowly around the sides of the trifoliate 
opesia. Spines are entirely absent from our specimens, though small ones 
are said to occur occasionally. Avicularia are rare, somewhat elevated, 
usually located near the proximal end and directed either forward or 
backward, the mandible short-pointed. 

The ovicell is hyperstomial, prominent; in final calcification there is a 
strong arcuate transverse rib which often rises to a central point, and 
proximal to this is a semilunar smooth area ; width 0.25 to 0.30 mm. 

It is a northern and arctic species, evidently circumpolar ; in Atlantic 
waters it extends southward to the British Isles and to the Maine coast 
of North America, but it has not been recorded from the Pacific coast 
before. 

Point Barrow, Alaska, Arctic Research Laboratory, 262 to 328 feet, 
G. E. MacGinitie, collector. 

Bugula flabellata acuminata new variety. 
Plate 81, figs. 3 and 4 

The zoarium is erect with broad flabellate branches, the secondary 
branches biserial but sometimes becoming quadriserial near the tips. The 
mode of branching is like that of flabellata and the origin of the zooecia 
is also similar, with long prongs extending distally down the dorsal sides 
of the preceding zooecium. Height of colony about 20 mm. 

The zooecia are long and slender, length 0.55 to 0.75 mm, width at 
base 0.13 mm and Avidening gradually to 0.15 mm near the tip; the 
membranous area extends nearly but not quite to the proximal end, 
narrowing downward ; there are no jointed spines, but the distal corners 
are extended into short, stout processes, sometimes wanting on the inner 
corner. The avicularia, present on nearly all of the zooecia, are attached 
by a short stalk more or less above the middle of the zooecium but not 
near the extremity ; moderately large and somewhat compressed, length 
0.20 to 0.30 mm, width 0.08 to 0.10 mm, height 0.13 to 0.15 mm; the 
beak sharply decurved at the tip with a narrow rounded point. The 
mandible, 0.15 to 0.20 mm long, is unique in my experience as it sud- 
denly becomes constricted near its distal end into a long acuminate 
recurved process sharply differentiated from the basal part. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 777 

The ovicells, on nearly all of the zooecia, are set transversely across 
the distal end of the zooecia, rounded and prominent, 0.18 mm wide by 
0.14 mm long, complete, but the orifice large, the surface not striated, 
in reproduction in February. 

The complete absence of jointed spines, the form of the avicularium 
and especially the sharply acuminated mandible appear to separate this 
variety sufficiently from B. flabellata. 

Type, AHF no. 130. 

Type locality, Hancock Station 66-33, Tagus Cove, Albemarle Island, 
Galapagos, 0°16'17"S, 91°22'41"W, at 10 to 20 fms, several colonies. 

ASCOPHORA 

Pachyegis brunnea (Hincks) 
Plate 81, fig. 11 

See Part 2, p. 315. 

The ovicells can now be described as I have found them at the margin 
of a large colony. They resemble those of P. princeps (Norman), hyper- 
stomial and prominent but considerably depressed on the base of the 
succeeding zooecium. With increased calcification they become more 
submerged but lack the heavy collar which develops across the front of 
P. princeps; width and length about 0.40 mm. 

Emballotheca stylifera (Levinsen), 1886 
Plate 81, fig. 5 

Escharella stylifera Levinsen, 1886:17. 
Schizoporella condylata Nordgaard, 1906:18. 
Schizoporella (Emballotheca) stylifera, Levinsen, 1916:453. 
Schizoporella stylifera, Nordgaard, 1918:57. 

Zoarium encrusting shells and stones, unilaminar, and the shining 
ectocyst with reddish-brown pigment. The zooecia are moderate in size, 
0.60 to 0.75 mm long by 0.40 to 0.55 mm wide, elongate-hexagonal in 
form and rather regularly arranged in quincunx, separated by very thin 
raised lines. The moderately inflated frontal is a tremocyst with fewer 
pores than is usual in the genus. Beneath the rather thick ectocyst the 
frontal is finely granulated, but there are no other surface decorations, 
and spines and avicularia are both wanting. Dietellae conspicuous on 
the dorsal surface. The aperture is nearly round, 0.16 to 0.18 mm wide, 
somewhat transverse proximal to the prominent cardelles, and between 
these is a shallow arcuate sinus nearly half as wide as the aperture. 



778 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

There is a narrow vestibular arch. The primary peristome is low and 
thin and becomes covered on the sides by the encroachment of the frontal 
wall, which modifies the form of the aperture only slightly. 

The ovicell is hyperstomial, considerably depressed and is closed by 
the operculum; secondary calcification from the three adjacent distal 
zooecia produces three sutural lines on the surface, and often there is 
a pore or fenestra at the central junction of the sutures, width 0.26 to 
0.30 mm. 

The species was described from the Kara Sea and later recorded from 
northeast Greenland (Levinsen) and the North American Archipelago 
(Nordgaard). 

Point Barrow, Alaska, Arctic Research Laboratory, 217 to 522 feet, 
G. E. MacGinitie, collector, rather common. 

Hippodiplosia cancellata (Smitt), 1867 
Plate 81, fig. 6 

Escharella porifera jorma cancellata Smitt, 1867:9. 
Smittina cancellata, Nordgaard, 1906:29. 

Apparently this species has not been seen since Smitt described it. 
Nordgaard placed it under Smittina, but without having seen it. 

The zoarium is encrusting, spreading over shells, reddish-brown in 
color. The zooecia are moderately large, 0.65 to 1.0 mm long by 0.40 
to 0.65 mm wide; the front a tremocyst with large pores which enlarge 
upward until the surface is coarsely cancellous or tessellated, with only 
narrow rims separating the pores. The only exception to this is the 
smoother area proximal to the aperture which is characteristic of the 
genus and upon which the secondary layer does not encroach. The aper- 
ture is large, 0.18 to 0.20 mm wide, usually a little wider than long, 
evenly rounded back to the small cardelles ; the proximal border broadly 
arcuate, nearly straight, or often arched forward slightly above the 
operculum (this last feature gives the aperture somewhat the appearance 
of a Smittina, but it does not appear to be homologous with the lyrula 
of the Smittinidae). Smitt described the species "Avicularia desunt," 
but small avicularia are frequently present (though often wanting over 
large areas), with a rounded mandible and located close to the aperture 
in the middle on the preoral smooth area. There are no spines or other 
surface structures. 

The ovicells are large, about 0.40 mm wide, hemispherical, hyper- 
stomial but rather deeply embedded, smooth at first but soon becoming 
covered with a cancellous layer similar to that of the frontal. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 779 

The species is related to H. reticulato-punctata (Hincks), but the 
measurements are larger, the aperture shorter and wider, the preoral 
area shorter and the cancellation of the front much more strongly 
developed. 

Smitt described the species from Spitsbergen and carefully figured it 
(plate 24, figs. 40 and 41), but I have not been able to find any other 
record of its occurrence. 

Point Barrow, Alaska, Arctic Research Laboratory, 438 feet, G. E. 
MacGinitie, collector, apparently rare. 

Microporella arctica (Norman), 1903 
Plate 81, fig. 7 

Microporella arctica Norman, 1903:105. 

Porina ciliata Smitt, 1867:6 (part). 

Microporella ciliata var. arctica, Nordgaard, 1918:60. 

Zoarium encrusting stones and shells, heavily calcified, and the ectocyst 
brown in color. The zooecia are much larger than those of M. ciliata 
in all their measurements, 0.65 to 0.80 mm long by 0.40 to 0.65 mm 
wide; the aperture 0.12 mm long by 0.15 mm wide; the ovicell averaging 
0.40 mm in width. The frontal is a thick tremocyst with numerous small 
pores which often become occluded in complete calcification ; inflated 
with deep separating grooves and smooth, except occasionally there is a 
minute umbo proximal to the ascopore. The aperture (except for size) 
and the ascopore are similar to those of ciliata. Avicularia, rarely present, 
are located near the lateral margin proximal to the aperture, the man- 
dibles varying in length, the longer ones noticeably broader (more 
ligulate) than those of ciliata. Spines are usually wanting but I have 
noted 4 very minute (vestigial) ones on a few young marginal zooecia. 

The ovicell is hemispherical, delicately ribbed radiately in the young 
stage but soon becoming very thick-walled and smooth, with a small 
umbo distally situated on the top. 

Recorded previously by Smitt, Norman and Nordgaard from Spits- 
bergen and northern Norway. The differences mentioned above indicate 
that it should be separated from M. ciliata, and the following record 
shows that it is a circumpolar species. 

Point Barrow, Alaska, Arctic Research Laboratory, 328 feet, G. E. 
MacGinitie, collector. 



780 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 



g ^' Escharoides jacksoni (Waters), 1900. 

f' ^^ Plate 81, fig. 8 

Smittia Jacksonii Waters, 1900:87. 
Escharoides jacksoni, Nordgaard, 1918:55. 
Peristomella (Escharoides) jacksoni, Osburn, 1923:10. 

Zoarium encrusting, the ectocyst reddish-brown. The zooecia are large, 
0.65 to 1.0 mm long by 0.50 to 0.65 mm wide; the frontal a pleurocyst 
with numerous large areolar pores between which narrow costal ridges 
extend upon the front, usually leaving a smoother central area but some- 
times extending to the tip of the high lip of the peristome. The zooecia 
are considerably inflated and elevated toward the distal end, ovate in 
form and arranged in quincunx. The most striking feature is the form 
of the peristome which is unusually high above the proximal border of 
the aperture, where it is more or less pointed, and extends sharply down- 
ward on each side to the first spine, resembhng an inverted scoop; it 
lacks the median denticle near the tip which is found in some others of 
the genus. There are 4 stout oral spines, the proximal pair usually remain- 
ing at the corners of the ovicell. The aperture is large, 0.20 to 0.26 mm in 
either direction, rounded proximally, and broader at the distal end, 
where it is broadly arcuate or sometimes nearly transverse. There are 
no condyles and no lyrula. The avicularia are large and conspicuous, 
situated characteristically at one or both sides of the peristome and 
directed laterally, but often located more proximally on the front and 
directed more or less backward, often wanting; the rostrum is elevated, 
the mandible hooked at its tip and measuring 0.25 to 0.30 mm in length 
and 0.15 to 0.20 mm wide at the base. 

The ovicells are correspondingly large, averaging 0.40 mm in width 
and length, hyperstomial, hemispherical and conspicuous, perforated but 
with a small central imperforate area on the top. 

It is an arctic species, described from Franz Josef Land and recorded 
from Greenland and the western part of the American Archipelago. 
The following record extends its distribution much farther to the west 
and it is evidently a circumpolar species. 

Point Barrow, Alaska, Arctic Research Laboratory, 453 feet, G. E. 
MacGinitie, collector. 

Porella minuta (Norman), 1868 
Plate 81, fig. 9 

Lepralia minuta Norman, 1868:308. 
Porella ininuta, Hincks, 1880:326. 
Porella alba Nordgaard, 1906:25. 

n 7j- „.: j_ XT_„j J inio.'7i 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA — CYCLOSTOMATA 781 

Zoarium small, encrusting on stones and shells, white and glistening. 
The zooecia are small, 0.40 to 0.50 mm long by 0.25 to 0.30 mm wide; 
the front considerably inflated and delicately granulated; the areolar 
pores are few in number and conspicuous only in the younger stages. 
The zooecia are very distinct, with deep separating grooves which never 
become filled even in the oldest and most complete stages of calcification. 
The aperture is semicircular, 0.13 mm wide by 0.10 mm long, the 
proximal border straight or slightly arcuate and without any indication 
of a lyrula. The primary peristome is low and thin and there are no 
spines, even in young marginal zooecia. There is a rounded median 
avicularium with a prominent bulbous chamber which covers nearly 
half of the frontal area proximal to the aperture. In advanced calcifica- 
tion, the frontal becomes very thick, extends around the sides of the 
aperture, and sometimes forms a small umbonate process on the top of 
the avicularian chamber. 

The ovicell at first is prominent, 0.24 mm wide by 0.22 mm long, 
but soon becomes more or less immersed, thick-walled and granulated 
like the frontal. 

The species resembles P. concinna (Busk), but the smaller dimen- 
sions, the proportionally larger avicularian chamber, and the distinct 
separation of the zooecia at all stages clearly distinguish it. It is known 
from the British Islands (Norman and Hincks), and Nordgaard rede- 
scribed it as P. alba from the North American Archipelago. 

Point Barrow, Alaska, Arctic Research Laboratory, at 216 to 522 
feet, G. E. MacGinitie, collector, rather common. 

Mucronella microstoma (Norman), 1868 
Plate 81, fig. 10 

Lepralia microstoma Norman, 1864:87. 

Mucronella microstoma, Hincks, 1880:370. 

1 Mucronella microstoma, O'Donoghue, 1923:46. 

This species bears much resemblance to M. labiata, but it is smaller; 
the primary aperture 0.13 mm wide by 0.10 long, semicircular, straight 
on the proximal border with a wide lyrula which is low and without 
denticles at the corners. The secondary aperture also is much smaller 
than in labiata, wider than long, and the peristome is more or less 
tubular, rising high on the proximal border and often continued into a 
central point. The dietellae are small and numerous. There are 2 to 4 
short stout oral spines on the distal border of the aperture, the two 
median ones the largest. 



782 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

The ovicell is prominent, about 0.40 mm in either direction, the 
surface finely granulated like the frontal. 

The most characteristic feature is the spout-like peristome and the 
small secondary aperture, about 0.18 mm wide by 0.12 mm long (though 
there is considerable variation). It has been recorded positively only 
from the British Isles at depths from 80 to 205 fms. O'Donoghue listed 
it somewhat doubtfully from Northumberland Channel, British Co- 
lumbia, at 15 to 18 fms, but as he did not give a full description and 
mentioned only a few variations this record must remain in doubt until 
the species is recovered from that area. 

Point Barrow, Alaska, Arctic Research Laboratory, 48 to 80 fms, 
G. E. MacGinitie, collector, rather common. 



NO. 3 OSBURN : EASTERN PACIFIC BRYOZOA CYCLOSTOMATA 783 

REFERENCES 

Hastings, A. B. 

1930. Cheilostomatous Polyzoa from the Vicinity of the Panama Canal, col- 
lected by Dr. C. Crossland on the Cruise of the S. Y. 'St. George'. 
Proc. Zool. Soc. London. 1929, vol. 2, pp. 697-740, 17 pis. 

HiNCKS, T. 

1880. A History of the British Marine Polyzoa. London. 2v. 
1882. Report on the Polyzoa of the Queen Charlotte Islands. Ann. and Mag. 
Nat. Hist. ser. 5, vol. 10, pp. 459-471. 

Levinsen, G. M. R. 

1886. Bryozoer fra Kara-Havet. Adv. reprint from Dijmphe-Togtets zoolo- 

gisk-botaniske Udbytte . . . Udgivet . . . ved C. F. Lijtken. Kj0ben- 

havn, 1887. 
1916. Bryozoa. [Denmark] Medd. Groenl. vol. 43, pp. 432-472, pis. 19-24. 

(Danmark-Exspeditionen til Groenlands Nord0stkyst 1906-08. vol. 3 

no. 16.) 

MacGillivray, p. H. 

1868. Descriptions of some new Genera and Species of Australian Polyzoa; 
to which is added a List of Species found in Victoria. Trans, and Proc. 
Roy. Soc. Victoria, vol. 9, pp. 126-148. 

Marcus, E. 

1940. Mosdyr (Bryozoa eller Polyzoa). Danmarks Fauna, Kj0benhavn. no. 
46, pp. 1-401, 221 figs. 

NORDGAARD, O. 

1906. Bryozoa from the second "Fram" expedition, 1898-1902. Rpt. of the 

Second Norwegian Arctic Expedition in the "Fram" 1898-1902. Kris- 

tiania, 1907. vol. 2 no. 8, pp. 1-44, pis. 1-4. 
1918. Bryozoa from the Arctic regions. Tromso Mus. Aarshefter. vol. 40, 

pp. 1-99. 

Norman, A. M. 

1864. On undescribed British Hydrozoa, Actinozoa, and Polyzoa. Ann. and 

Mag. Nat. Hist. ser. 3, vol. 13, pp. 82-90. 
1868. Shetland Final Dredging Report. — Part H. On the Crustacea, Tuni- 

cata, Polyzoa, Echinodermata, Actinozoa, Hydrozoa, and Porifera. 

Class Polyzoa. Rpt. Brit. Assoc. Adv. Sci. 38th meeting, 1869, pp. 

303-312. 
1903. Notes on the Natural History of East Finmark. Ann. and Mag. Nat. 

Hist. ser. 7, vol. 12, pp. 87-128, pis. 8-9. 

O'DoNOGHUE, C. H. and Elsie O'Donoghue 

1923. A Preliminary List of Bryozoa (Polyzoa) from the Vancouver Island 
Region. Contr. Canad. Biol. Fish, new sen, vol. 1, pp. 143-201 (1-59), 
pis. 1-4. 

OSBURN, R. C. 

1912. Bryozoa from Labrador, Newfoundland, and Nova Scotia, collected 
by Dr. Owen Bryant. Proc. U. S. Natl. Mus. vol. 43, pp. 275-289. 
1923. Bryozoa. In Report of the Canadian Arctic Expedition 1913-1918. 
Ottawa, vol. 8 (D), pp. 1-13. 



784 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 

1932. Biological and Oceanographic conditions in Hudson Bay. 7. Bryozoa 
from Hudson Bay and Strait. Contr. Canad. Biol. Fish, new ser., vol. 
7, pp. 361-376 (1-16), 1 pi. 

1933. Bryozoa of the Mount Desert Region. Repr. from the Biological Sur- 
vey of the Mount Desert Region. Philadelphia. 67p., 15 pis. 

1950. Bryozoa of the Pacific Coast of America. Part 1, Cheilostomata-Anasca. 
Allan Hancock Pacific Expeditions, vol. 14, no. 1, pp. 1-269, 29 pis. 

Packard, A. S. 

1867. Observations on the Glacial Phenomena of Labrador and Maine, with 
a View of the recent invertebrate Fauna of Labrador. H. View of the 
recent invertebrate Fauna of Labrador. Mem. Boston Soc. Nat. Hist. 
vol. 1, pp. 262-303, pis. 7-8. 

Smitt, F. A. 

1867. Kritisk forteckning ofver Skandinaviens Hafsbryozoer. Ofversigt af 
Svenska Vetensk. Akad. Forhandl. vol. 24 (Bihang), pp. 1-230, pis. 

24-28. 

Verrill, a. E. 

1879. Preliminary Check-list of the Marine Invertebrata of the Atlantic 
Coast, from Cape Cod to the Gulf of St. Lawrence. New Haven, Conn, 
pp. 28-31. 

Waters, A, W. 

1900. Bryozoa from Franz-Josef Land, collected by the Jackson-Harmsworth 
Expedition, 1896-1897. Jour. Linn. Soc. London. Zool. vol. 28, pp. 
43-105, pis. 7-12. 

Whiteaves, J. F. 

1901. Catalogue of the marine invertebrata of eastern Canada. Rpt. Geol. 
Surv. Canada [sep. pubn.] Polyzoa, pp. 91-114. 

Wood, S. V. 

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Nat. Hist. vol. 13, pp. 10-21. 



PLATES 

Practically all of the illustrations for the following; plates, 65 to 82, 
are the work of Mrs. Virginia Sewell, to whom the author is much 
indebted for her careful attention to details and interested cooperation. 

The scale of enlargement is not indicated on the plates, except in the 
Ctenostomata. but exact measurements are given for each species in the 
text. 



788 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 



PLATE 65 

Fig. 1. Stomaiopora f/ranulata (Milne-Edwards), part of zoarium. 

Fig. 2. The same, ovicell with ooeciostome. 

Fig. 3. Prohoscina sif/mala new species, part of zoarium. 

Fig. 4. The same, enlarged to show details of ovicell. 

Fig. 5. Prohoscina major (Johnston), part of zoarium with ovicell. 

Fig. 6. Oncousoecia ahrupta new species, zoarium. 

Fig. 7. The same, enlarged to show details of 3 ovicells. 

Fig. 8. Oncousoecia ovoidca new species, zoarium. 

Fig. 9. The same, enlarged to show details of ovicell. 

Fig. 10. Oncousoecia canadensis Osburn, showing usual type of 

ovicell. 

Fig. 11. The same, at less magnification, \\\t\\ broader ovicell. 



XO. 3 OSBURN: EASTERN PACIFIC BRYOZOA CYCLOSTOMATA PL. 65 




790 ■ ALLAN HAXCOCK PACIFIC EXPEDITIONS VOL. 14 



PLATE 66 

F'ig. 1. Proboscina iturassala (Smitt), zoarium. 

Fig. 2. The same, portion of a lobe with oviceil and ooeciostome. 

Fig. 3. Proboscina lamcllifera Canu and Bassler, part of lobe with 

oviceil and ooeciostome. 
Fig. 4. Oncousoccia diasto [sondes (Norman), oviceil and ooecio- 
stome. 
Fig. 5. Plagioecia (iriinaldii (JuUien), showing terminal, median 

position of ooeciostome. 
Fig. 6. Plagioccia meandrina (Canu and Bassler) showing erect 

branches and lobes. 
Fig. 7. The same, part of a lobe showing position of oviceil and 

ooeciostome. 
Fig. 8. Pla{/ioecta ambiffua new species, zoarium, with simple and 

expanded ovicells; note different positions of ooeciostomes. 
Fig. 9. Plarjioeiia anacapcnsis new species, outline of zoarium and 

linear arrangement of tubules. 
Fig. 10. The same, showing oviceil and ooeciostome. 
Fig. 11. Diapcroecia claviformis new species, part of a lobe showing 

proximal position of ooeciostome. 



XO. 3 OSBURN : EASTERX PACIFIC BRVOZOA CYCLOSTOMATA PL. 66 




Id .<f '"f 




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792 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 



PLATE 67 

Fig. 1. Diaf>eroecia calif ornica (d'Orbigii\ ) , showing arrangement 
of tubules and mode of branching. 

Fig. 2. The same, ovicell and large, tall ooeciostome. 

Fig. 3. Diaperoecia florijana Osburn, part of a branch with elon- 
gate ovicell and large, flared, reflected ooeciostome. 

F'ig. 4. Diaperoecia jolmstoni (Heller), portion of a lobe \vith ovi- 
cell, and ooeciostome at the side of a peristome. 

Fig. 5. Tuhulipora concinna MacGillivray, ovicell with ooecio- 
stome. 

Fig. 6. Tuhulipora ecjregia new species, an ovicell with ooeciostome 
which has an oval and slightly flared lip. 

Fig. 7. The same, portion of a zoarium with lobate ovicells. 

Fig. 8. Placjioecia tortuosa new species, outline of an irregular 
free lobe, with ovicell and ooeciostome. 

Fig. 9. The same, outline of a zoarium, showing the beginning of 
a vertical bilaminate Inbe and the position of four ovicells. 



NO. 3 OSBURX : EASTERN- PACIFIC HRVOZOA — CYCLOSTO.MATA PL. 67 




H r m M^' ^ © 



794 



ALLAN HANCOCK PACIFIC EXPEDITIONS 



VOL. 14 



PLATE 68 



Fig. 


1 


Fig. 


2 


Fig. 


3 


Fig. 


4 


Fig. 


5 


Fig. 


6 


Fig. 


7 


F-ig. 


8 


Fig. 


9 


Fig. 


10 



Tuhulipora pacifira Robertson, with ovicell and ooeciostome. 
Tuhulipora pulclira MacGiiiivray, ovicell and ooeciostome. 
The same, base oi zoariiim with lateral attachment proc- 
esses. 

The same, portion of dorsal side with attachment processes. 
Tuhulipora admiranda new species, form of zoarium and 
distribution of ovicells. 

The same, ovicell with large erect ooeciostome. 
The same, base of zoarium with primary zooecium. 
Tuhulipora flabellaris (Fabricius), a small ovicell with 
narrow erect ooeciostome and slit-like aperture. 
Tuhulipora tuha (Gabb and Horn), showing large fas- 
cicles and ovicell with ooeciostome. 

Tuhulipora tuba var. fasciculifcra (Hincks), with small 
fascicles and ovicell with ooeciostome. 



XO. 3 OSBURN : EASTERX PACIFIC BRYOZOA CYCLOSTOMATA PL. 68 




796 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 



PLATE 69 

Fig. 1. PlaUinea elonrjata new species, with very elongate ovicell, 
and position of ooeciopore, ooeciostome broken off. 

Fig. 2. Platonea velfronis new species, with short ovicell and posi- 
tion and form of ooeciostome. 

Fig. 3. Platoiira cxpansa new species, with long fascicles, broad 
marginal lamina and position and form of ooeciostome. 

Fig. 4. Pxifliysoi'da bassleri new species, outline of zoarium show- 
ing position of two ovicells. 

Fig. 5. The same, depressed arcuate ovicell with slit-like ooecio- 
stome, the two lobes surrounding peristomes and a fascicle; 
note the very short peristomes. 

Fig. 6. The same, younger part of colony showing partially closed 
ends of erect tubes before the peristomes are developed. 

Fig. 7. Batliysoccia liastitK/sae new species, sketch of ovicell, the 
arcuate ovicell modified by several lobes. Note the position 
and form of the ooeciostnme. 

Fig. 8. FiUfascigera clariotwnsis new species, portion of zoarium 
showing mode of branching and position of ovicell. 

Fig. 9. The same, enlarged to show the details of the ovicell and 
ooeciostome. 

Fig. 10. The same, a double erect fascicle, enlarged. 

Fig. 11. Disiocylis californua new species, with broad, thin base, 
short peduncle and the broad capltulum with the large 
rounded ovicell on its dorsal side. 



NO. 3 OSRL RX : EASTERN PACIFIC HRYOZOA 



CYCLOSTO.MATA PL. 69 






6 ^'^ 












798 ALLAX HAXCOCK PACIFIC EXPEDITIONS VOL. 14 



PLATE 70 

Fig. 1. Fasriculipora pacifita new species, young colony with about 

7 branches beginning to form. 
Fig. 2. The same, still younger with the beginning of branches at 

the right and left borders, more enlarged. 
Fig. 3. The same, a very young colony with the proancestrula and 

first tubule; origin of secondary tubules on the right and 

the beginning of a branch on the left. 
Fig. +. The same, from an adult zoarium, showing a simple ovicell 

at the right and a broader one involving two peristomes at 

the left. 
Fig. 5. Diaperoecia intermedia (O'Donoghue), an erect branch 

with ovicell and central ooeciostome. The capltulum here 

is narrower than usual. 
Fig. 6. Enialopliora symmetrica new species, showing method of 

branching at right angles and the position of the ovicell 

at the left. 
Fig. 7. The same, simple ovicell with terminal ooeciostome. 
Fig. 8. Entalophora proboscideoides (Smitt), portion of zoarium 

with simple ovicelF 
Fig. 9. The same, side view of ovicell and ooeciostome. 
Fig. 10. Birntalnphora cylindrica new species, outline of zoarium; 

the left branch is beginning to branch again in the same 

plane. 
F'ig. 11. The same, showing arrangement of peristomes and the 

covering layer of closed kenozoids. 



NO. 3 OSRURN: EASTERN PACIFIC BRYOZOA CVCLOSTOM ATA PL. 70 




800 ALLAX HAXCOCK PACIFIC EXPEDITIOXS VOL. 14 



PLATE 71 

Fig. 1. Crisiciia rnrnuta (L.), part of zoariiim showing mode of 
branching and position of spine-like processes. 

F'ig. 2. Birrisia edivardsianu d'Orbigny, \vith oviceil and spine- 
like processes. 

Fig. 3. Crista occidentalis Trask, showing mode of branching and 
normal form of oviceil. 

Fig. 4. The same, distorted oviceil due to curved internode. 

Fig. 5. The same, pointed tip of terminal internode, often present. 

Fig. 6. Crista operculata Robertson, frontal view of oviceil and 
ooeciostome. 

Fig. 7. The same, sketch of side view showing the cap or "oper- 
culum" above the ooeciopore. 

Fig. 8. Crisia pugeti Robertson, frontal view of oviceil and, at 
left, a sketch of the side view with betit ooecif'stome. 

Fig. 9. Crisia elongata Milne-Edwards, frontal view with short, 
wide expansion of the oviceil. 

Fig. 10. Crisia ehurnea (L.), with short, transverse ooeciostome. 

Fig. 11. Tiihulipora flextiosa (Pourtales), a portion of a branch 
with short oviceil and ooeciostome distal to a peristome. 



NO. 3 OSRURN : EASTERN PACIFIC BRYOZOA — CYCLOSTO.M ATA PL. 71 




802 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 



PLATE 72 

Fig. 1. Crisia cribrar'ta Stimpson, mode of branching and ovicell. 

Fig. 2. Crisia scrrulata new name, ovicell and mode of branching; 
note at left the usual short inserted basis rami. 

Fig. 3. Crisia maxima Robertson, ovicell and mode of branching; 
note long free end of the tubules and the small reflected 
ooeciostome. 

Fig. 4. Filicrisia franciscana Robertson, mode of branching and 
frontal position of ooeciostome. 

Fig. 5. Filicrisia geniculata (Milne-Edwards), infertile internode, 
ovicell with dorsal position of ooeciostome, and smaller 
dimensions. 

Fig. 6. Crisulipora occidrntalis Robertson, a fertile internode show- 
ing long peristomes and ovicell spreading among peri- 
stomes, ooeciostome centrally placed near distal end. 

Fig. 7. Hornrra pinnata Canu and Bassler, sketch of frond and 
base. 

Fig. 8. The same, frontal view of branch with lateral pinnule. 

Fig. 9. The same, dorsal view of tip of branch. 

Fig. 10. Hornera pectinata Busk, sketch of frond and base. 

Fig. 11. The same, frontal view of tip of branch, showing the 
irregular, pectinate tips of the tubules. 

Fig. 12. The same, dorsal side of tip of branch. 



NO. 3 OSBLRX : EASTERX PACIFIC BRYOZOA CYCLOSTOMATA PL. 72 




804 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 



PLATE 73 

Fig. 1. Diplosolen ohrlium (Johnston), ovicell enclosing peristomes 

of normal and diminutive tubules; ooeciostome near center. 
Fig. 2. PlcKjioecia tuhiahurth'a (Canu and Bassler), ovicell with 

terminal ooeciostome. 
Fig. 3. Plagioecia sarniensis (Norman), sho\ving basal tubule and 

transverse expansion of ovicell and terminal ooeciostome; 

one peristome enclosed. 
F'ig. 4. Platjioecia patina (Lamarck), basal tubule and usual broad 

expansion of ovicell, with terminal ooeciostome. 
Fig. 5. Rorgiola pustulosa new species, surface view with "pus- 
tules," and broken edge of type specimen. 
Fig. 6. The same, at broken edge showing brood-chainber traversed 

b\- tubules. 
Fig. 7. The same, showing the occasional pointed peristomes at the 

edge of a pustule. 
Fig. 8. The same, margin of zoarium. 
Fig. 9. The same, enlargement of a pustule. 
Fig. 10. Heteropora alaskensis (Borg), terminal portion of branch 

\vith brood-chamber, the roof partly removed to show the 

cavit\ traversed by tubules; note also the high peristomes. 
Fig. 11. The same, closure and partial closure of peristomes near 

base of zoarium. 
Fig. 12. The same, mode of branching of a >()ung zoarium. 
Fig. 13. Heteropora magna O'Donoghue, a characteristic zoarium 

showing mode of branching and anastomosis; the primary 

base and two secondarv attachments. 



NO. 3 OSBURN: EASTERN PACIFIC RRYOZOA CYCLOSTO.M ATA PL. 73 



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806 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 



PLATE 74 

Fig. 1. Lklienopora huskiana Catui and Bassler, roof of ovicell 
broken away; the distribution of the radii on the disc is 
often irregular. 

Fig. 2. The same, irregular secondary cancelii above ovicell, posi- 
tion and form of ooeciostome. 

Fig. 3. Liclienopora -verrucaria (Fabricius), showing quincuncial 
arrangement of peristomes, irregular cancellated cover of 
ovicell, and position and form of ooeciostome. 

F'ig. 4. Liclienopora novae-zclandiac (Busk), irregular secondary 
cancelii covering ovicell, position and form of ooeciostome, 
and occasional extra tubules in the uniserial radii. 

Fig. 5. Disporella separata new species, small portion of zoarium 
showing form and separation of the discs. 

Fig. 6. The same, enlarged, showing the irregular nature of the 
radii and the round partially closed cancelii. 

Fig. 7. Disporella calif arnica (d'Orbigny), diagram of complex 
zoarium with two complete discs and three incomplete 
marginal ones. 

Fig. 8. The same, at margin of central area, showing ovicell with 
perforated cover (seen at left through the rounded sec- 
ondary cancelii) ; position and form of ooeciostome. 

Fig. 9. The same, enlargement of cancellar pores with pin-head 
spicules. 



NO. 3 OSBURX : EASTERX PACIFIC HRVOZOA — CYCLOSTOMATA I'l,. 74 




808 ALLAN HANCOCK. PACIFIC EXPEDITIONS VOL. 14 



PLATE 75 

Fig. I. Disporflla liispida (Fleming), showing form of zoarium, 
scattered peristomes, small cancellar pores, and two ooecio- 
stomes near the border indicating the presence of inter- 
radial ovicells. 

Fig. 2. Disporflla fimhriata (Busk), the ovicel! obscured by heavy 
calcification of the secondary canceili. 

Fig. 3. The same, a younger zoarium with three ovicells occupy- 
ing most of the central area, the ooeciopores marginal, 
ooeciostomes not developed. 

Fig. 4. Dispori'lla ovoidea new species, outline showing ovate form 
of zoarium and central area, uniserial radii and position 
of ovicel 1. 

Fig. 5. The same, much enlarged, an interradial ovicel! at left 
covered by rounded secondary canceili, and interradial 
ooeciostome ; beside this another interradial o\'icell dis- 
sected away to show the cavity. 

Fig. 6. ? Disporflla octoradiata (\^'ate^s), a \()ung zoarium with- 
out ovicell. 

Fig. 7. Disporella alaskcnsis ne\v species, side view, showing the 
height of the radii, the upturned margin and the small 
daughter zoarium. 

F'ig. 8. The same, frontal view, showing the complex nature of the 
radii, the position of the ovicell (roof broken away) and 
the position of the submarginal vertical bud of the daughter 
zoarium. 



XO. 3 OSBURX : EASTERX PACIFIC HRVOZOA— CYCLOSTOMATA 



PL. 75 











1© «.'®~-S^ft#'^^iJ'^ 












810 ALLAX HAXCOCK PACIFIC EXPEDITIOXS \'OL. 14 



PLATE 76 

Fig. 1. Disporella astraca new species, showing attachment, vertical 
budding and crown from side view. 
The same, top view of cro\vn (disc). 

Lichenopora canaliculata (Busk), view of disc with central 
ovicell and hooded ooeciostome. 
The same, enlargement of ooeciostome. 

Lichenopora intricata (Busk), portion of complex zoarium 
showing irregular distribution of peristomes between discs. 
The same, a fertile disc, with ooeciostome and irregular 
secondary cancelli covering ovicell. 

The same, infertile disc with uniformlv round small can- 
celli. 

The same, one end of a fertile disc with ooeciostome and 
irregular secondary cancelli covering ovicell. 
The same, the other end of the same disc with irregular 
cancelli over the end of the ovicell, and smaller rounded 
cancelli beyond the edge of the ovicell. 

Disporella siellata pacifica new variety, disc with large 
central area, multiserial radii and interradial position of 
ovicells. 

Borc/iola rugosa (Borg), side view of zoarium with broad 
encrusting base and erect irregular branches; a small sub- 
colony from the same base at the right. 



Fig. 


2. 


Fig- 


3. 


Fig. 


4. 


Fig. 


5. 


Fig. 


6. 


Fig. 


7. 


Fig. 


8. 


Fig. 


9. 


Fig. 


10 


Fig. 


11. 



XO. 3 OSniRX : EASTERN PACIFIC RRYOZOA CYCLOSTOM AT A I'L. 76 




812 ALLAN HAXCOCK PACIFIC EXPEDITIONS VOL. 14 



PLATE 77 

Fij^. 1. .lUyonhiium polyouin (Hassall), a portion of a zoarium 

■with a zooeciiim in detail. X 46. 
Fig. 2. .Ill yotiiiiiurn parasiticum (F'leming) , note minute border 

papillae on detailed zooeciunn. X 46. 
Fig. 3. AUyoTiidium pediinculatum Robertson, a portion of a zoari- 
um \vith one zooecium in detail. X 46. 
Fig. 4. .llcyotiidium matnmiUatum Alder, note aperture at apex of 

raised oral protuberance. X 46. 
Fig. 5. .Ilcyonidium disciforme (Smitt), a portion of a zoarium 

with a zooecium in detail. X 46. 
Fig. 6. The same, a drawing of a zoarium, ventral aspect, natural 

size. 
Fig. 7. Alcyonidium cntcromorpha Soule, a portion of a zoariun^. 

with one zooecium in detail. X 46. 
Fig. 8. The same, dorsal view of dissected zooecium, showing the 

anatomy of the polypide. X 73. 
Fig. 9. Flustrella corniculata (Smitt), a portion of a zoarium with 

a zooecium in detail. X 26. 



NO. J) OSRURX: EASTERX PACIFIC BRVOZOA CYCLOSTOM ATA PL. 77 




814 ALLAN HAXCOCK PACIFIC EXPEDITIONS VOL. 14 



PLATE 78 

Fig. 1. Flustrella (ji(/antea Silen, a portion of a zoarium ^vith a 

zooecium in detail. X 26. 
Fig. 2. Plwrusella brei'itiil/a Soule, a portion of a zoarium with a 

zooecium in detail. X 46. 
Fig. 3. CAavopora occidcnialis (F'ewkes), an entire zoarium. X 26. 
Fig. 4. Anguitiflla palmata van Beneden, a portion of a zoarium 

showing the arrangement of the zooecia. X 26. 
Fig. 5. Nolella stipata Gosse, a portion of a zoarium showing 

mode of growth, note polypide anatomy. X 26. 
Fig. 6. Vesicularia fasciculaia ne\v species, a portion of a zoarium 

showing mode of growth, one zooecium with polypide 

anatomy. X 46. 
Fig. 7. Amathia convoluta Lamouroux, a portion of a zoarium 

showing the characteristic spiral pattern of zooecial growth. 

X 46. 



NO. 3 OSRURN : EASTERN PACIFIC BRYOZOA CYCLOSTOM ATA PL. 78 




816 ALLAN HAXCOCK PACIFIC EXPEDITIONS VOL. 14 



PLATE 79 

Fig. 1. A mat Ilia distans Busk, a portion of a zoariiim showing the 
position of the zooecia. X 46. 

Fig. 2. Amatliia 'vido'vici (Heller), a portion of a zoarium show- 
ing the position of the zooecia. X 46. 

Fig. 3. y.oobotryon vertliillaluin (delle Chiaje), a portion of a 
zoarium showing the placement of the zooecia. X 26. 

Fig. 4. Boivfrbankia imliruata (Adams), a portion of a zoarium 
showing the position of the zooecia ; note anatomical de- 
tails. X 26. 

Fig. 5. Boivfthankia graaiis Leidy, a portion of a zoarium show- 
ing the position of the zooecia. X 26. 

Fig. 6. Roivcrhankia f/racilis a(j(jre(jata O'Donoghue, a portion of 
a zoarium showing the mode of zooecial growth. X 46. 

Fig. 7. Valkcna tubcrosa Heller, a portion of a zoarium with one 
zooecium in detail. X 46. 

Fig. 8. A ei'ertillia si'fiffcra (Hincks), a pair of zooecia. X 46. 



XO. 3 OSHURN: EASTERN^ PACIFIC RRVOZOA— CYCLOSTOM ATA IM.. 79 




ALLAN HAXCOCK PACIFIC EXPEDITIONS VOL. 14 



PLATE 80 

F'ig. 1. Buskia nilrns Alder, two zooecia showing mode of growth. 
X 46. 

Fig. 2. Buskia seriata new species, a portion of a zoarium showing 
mode of zoarial growth, one zooecium with polvpide anato- 
my. X 46. 

Fig. 3. Fanella elotujata (van Heneden), a portion of a zoarium, 
one zooecium showing the anatomy of polvpide. X 26. 

Fig. 4. Triticclla pedicellata (Alder), a portion of a zoarium, one 
zooecium with anatomical details. X 26. 

Fig. 5. Triticrlla elontjata (Osburn), a portion of a zoarium, one 
zooecium showing details of polypide anatomy. X 26. 

Fig. 6. Terchripora comma Soule, a portion of a zoarium removed 
from a mollusk shell, one zooecium with polvpide anatomv. 
X 46. 

Fig. 7. Immcrgentia caliinrmca Silen, a portion of a zoarium re- 
moved from a mollusk shell, one zooecium with anatomical 
detail. X 46. 

Fig. 8. Pcnctrantia densa Silen, a portion of a zoarium removed 
from a mollusk shell, one zooecium \vith polypide anatomy, 
and also a typical gonozoid. X 46. 

Fig. 9. Pcnetrantia concharum Silen, a portion of a zoarium re- 
moved from a mollusk shell, one zooecium with anatomical 
detail, and also a typical gonozoid. X 46. 

Fig. 10. Penetrantia silent Soule, a portion of a zoarium removed 
from a mollusk shell, one zooecium with pol\pide anatomy, 
and also a typical gonozoid. X 46. 



XO. 3 OSBURX : EASTERX P. 



ACIFIC RKVOZOA— CYCLOSTO.MATA PL. f 



80 




820 



ALLAN HAXCOCK PACIFIC EXPEDITIOXS 



VOL. 14 



PLATE 81 



Fig. 


1. 


Fig. 


2. 


Fig. 


3. 


Fig. 


4. 


Fig. 


5. 


Fig. 


6. 


Fig. 


7. 


Fig. 


8. 


Fig. 


9. 


Fig. 


10. 


Fig. 


11. 


Fig. 


12. 



Hincksina gothiia new species, with zooecium, ovicell and 
large and snnall avicularia. 

A mphiblcstruin trifolium (S. Wood), zooecia \vith trifoliate 
opesia and ovicell. 

Bu(/ula flahfllala acuminata new variety, part of zoarium 
\vith different sizes of avicularia. 

The same, large avicularium, partial side view, and front 
view of mandible with acuminate point. 
Emballotheca stylifera (Levinsen), zooecia with ovicell. 
Hippodiplosta cancellata (Smitt), zooecia \vith cancellate 
frontal wall, minute median suboral avicularium, and 
ovicell. 

Microporella arctica Norman, showing thick-walled frontal, 
ligulate avicularium, and ovicell. 

Esiliai aides jacksoni (Waters), zooecia with spout-like peri- 
stome, spines, avicularia and ovicell. 

Porclla minuta (Norman), zooecia with suhoral aviculari- 
um and ovicell. 

Muironclla microstoma (Norman), zooecia showinjj nar- 
row aperture and spines. 

Pacliyccjis hrunnca (Hincks), ovicell (for description of 
species see Part 2, p. 315). 

Clifilopora praelucida (Hincks), ovicell (for description 
of species see Part 2, pp. 464-65). 



NO. 3 OSBURX : EASTERN PACIFIC HRYOZOA CVC 



LOSTO.MATA PL. 81 






"'V WW/ .\ 










822 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 14 



PLATE 82 

Owing to the nature of the material all of the figures are more 
or less diagrammatic, all to the same scale except figs. 6, 10 and 13. 

Fig. 1. Myosoma spinosa Robertson, portion of zoarium, zoid, stolon 
and very joung zoid. Note diagonal position of tentacle 
ring. 

Fig. 2. PedtcfUtna ccrnua (Pallas), a fertile internode with zoid; 
the spines are irregular in distribution and often wanting. 

Fig. 3. Barentsia (jracilis |M. Sars), fertile internode and zoid; 
the stalk is often twice as long as that figured. 

Fig. 4. Barentsia f/eniculata Harmer, short, wide basal bulb, mus- 
cular joints, comparatively small calyx. 

Fig. 5. Barentsia ramosa (Robertson), details of joints and form 
of calyx. 

F'ig. 6. The same, habit sketch to show mode of branching. 

Fig. 7. Barentsia roJiusta new species, showing large calyx, tall 
basal bulb and short internode (often shorter than the 
basal bulb), and attachment of bulb to stolon. 

Fig. 8. Barentsia discreta (Busk), large calyx; very elongate inter- 
node with "pores" for its entire length. 

Fig. 9. Barentsia suhriyida new species, stalk walls thin and flex- 
ible except at the base of the lower internode ; the propor- 
tions of the two internodes are very constant. 

Fig. 10. Barentsia gorhuno'vi Kluge, habit sketch of branch, inter- 
nodes without septa and three sizes of basal bulbs. 

Fig. 11. The same, details of part of branch, with medium and 
small basal bulbs. 

Fig. 12. The same, giant basal bulb at base of large branch, drawn 
to the same scale as fig. 11. 

Fig. 13. Coriella stolonata Kluge, habit sketch of erect branch 
formed of connate stolons. 

Fig. 14. The same, details of zoid; note that the basal bulb arises 
from a cup-shaped process of the fertile internode. 



XO. 3 OSBURX: EASTERN PACIFIC RRVOZOA — CYCLOSTOMATA IM.. S2 




INDEX 
Plate illustrations are in bold face. 



abrupta, Oncousoecia, 626, 789 
Acamptostega, 614, 617 
Acanthodesia serrata, 774 
Actinopora, 627, 628 

regularis, 628 
admiranda, Tubulipora, 649, 656, 795 
Aeverrillia, 745 

setigera, 745, 817 
alaskensis, Disporella, 709, 712, 715, 

809 

Heteropora, 695, 805 
alba,Porella, 780, 781 
Alcyonidiidae, 726, 727 
Alcyonidium, 727 

cervicorne, 732 

cervicornis, 732 

columbianum, 727 

corniculatum, 732 

disciforme, 727, 730, 813 

enteromorpha, 727, 731, 813 

raammillatum, 727, 729, 813 

mammillatum var. disciforme, 730 

mytili, 727 

parasiticum, 727, 729, 813 

pedunculatum, 727, 730, 731, 813 

polyoum, 727, 728, 813 

spinifera, 732 
Alcyonium gelatinosum, 727 

parasiticum, 729 
Alecto, 619 

diastoporides, 624 

dichotoma, 619 

dilatans, 624 

granulata, 619 

major, 621 

retiformis, 623 
Amathia, 740, 747 

convoluta, 740, 742, 746, 815 

distans, 741, 817 

vidovici, 741, 746, 817 
ambigua, Plagioecia, 629, 631, 638, 791 
americana, Pedicellina, 763 
Amphiblestrum trifoHum, 775, 821 
anacapensis, Plagioecia, 630, 637, 791 
Anasca, 774 
Anguinella, 738 

palmata, 738, 815 
annae, Membranipora, 774 
Apsendesia, 716 
arctica, Microporella, 779, 821 
Arthropodaria, 764 
Articulata, 614, 674 
Ascophora, 777 
Ascopodaria, 764 

discreta, 766 



gracilis, 765 

macropus, 769, 770 

misakiensis, 766 
Ascorhiza occidentalis, 735 
astraea, Disporella, 709, 719, 811 
atlantica, Tubulipora (Idmonea), 654 
atlantica var. flexuosa, Idmidronea, 654 
atlantica var. flexuosa, Idmonea, 653, 

654 
atlantica var. flexuosa, Tubulipora, 654 
atlantica var. tenuis, Idmonea, 653, 654 
australis, Pedicellina, 766 
Barentsia, 761,764, 771 

bulbosa, 764 

discreta, 764, 765, 766, 767, 768, 823 

geniculata, 765, 769, 823 

gorbunovi, 765, 767, 823 

gracilis, 765, 823 

gracilis nodosa, 765 

laxa, 764, 770 

misakiensis, 766 

ramosa, 764, 765, 767, 769, 823 

robusta, 765, 768, 823 

subrigida, 764, 770, 823 

timida, 766 
bassleri, Bathysoecia, 658, 659, 660, 797 
Bathysoecia, 648, 658 

bassleri, 658, 659, 660, 797 

hastingsae, 659, 660, 797 
Berenicea, 627, 628, 639 

obelia, 640 

prominens, 627 

sarniensis, 632 
Bicrisia, 674, 676 

edwardsiana, 676, 801 
Bidiastopora, 627, 628 
Bientalophora, 667, 669, 670 

cylindrica, 670, 799 

(Entalophora) regularis, 671 

regularis, 671 
Borgiola, 692, 696, 699 

pustulosa, 697, 698, 699, 805 

rugosa, 697, 699, 811 
boryi, Proboscina, 620 
Bowerbankia, 743 

gracilis, 743, 744, 817 

gracilis var. aggregata, 744, 817 

imbricata, 743, 817 
Brachionus cernuus, 763 
brevituba, Pherusella, 734, 815 
brunnea, Pachyegis, 774, 777, 821 
Bugula flabellata, 776, 777 

flabellata acuminata, 776, 821 
bulbosa, Barentsia, 764 
bullata, Lichenopora, 718 



[827] 



828 



INDEX 



VOL. 14 



Buskia, 746 

nitens, 746, 747, 748, 819 

seriata, 747, 819 

setigera, 745 

socialis, 748 
buskiana, Lichenopora, 701, 704, 712, 

807 
Buskiidae, 746 
californica, Crisia, 685 

Diaperoecia, 641, 642, 643, 793 

Discocytis, 690, 797 

Discoporella, 704, 711 

Disporella, 709, 711, 807 

Idmonea, 642, 643 

Immergentia, 752, 753, 819 

Lichenopora, 704, 711 

Unicavea, 704, 711, 712 
Calvetiidae, 687 
Calyptrostega, 615, 699 
Calyssozoa, 759 
Camptostega, 614, 674 
canadensis, Discocytis, 690, 691 

Oncousoecia, 625, 789 
canaliculata, Discoporella, 702 

Lichenopora, 702, 703, 811 
Cancellata, 614, 687 
cancellata, Hippodiplosia, 778, 821 

Smittina, 778 
Canuella, 692, 696 

rugosa, 697, 698 
capitata, Diaperoecia, 668 

Diaperoecia (Entalophora), 645 

Entalophora, 668 
Carnosa, 726 
catillus, Diastopora, 637 
cellarioides, Entalophora, 667 
Ceriopora cryptopora, 692 
Cerioporina, 615, 692 
cernua, Pedicellina, 763, 823 
cernuus, Brachionus, 763 
cervicorne, Alcyonidium, 732 
cervicornis, Alcyonidium, 732 

Diastopora, 628 
Cheilopora praelucida, 774, 821 
Cheilostomata, 613, 774 
ciliata, Microporella, 779 

Porina, 779 
ciliata var. arctica, Microporella, 779 
cirrosa, Gattyana, 761 
clarionensis, Filifascigera, 672, 673, 797 
clavata, Diaperoecia (Entalophora), 

645 

Entalophora, 669 
claviformis, Diaperoecia, 641, 642, 647, 

791 
Clavopora, 730, 735 , 

hystricis, 735 

occidentalis, 735, 815 
Clavoporidae, 726, 735 
columbianum, Alcyonidium, 727 
comma, Terebripora, 752, 819 



compressa, Porella, 775 
concharum, Penetrantia, 754, 819 
concinna, Porella, 781 

Tubulipora, 649, 655,793 
condylata, Schizoporella, 777 
convoluta, Amathia, 740, 742, 746, 815 
Coriella, 761, 764, 771 

stolonata, 771, 823 
corniculata, Flustrella, 732, 813 
corniculatum, Alcyonidium, 732 
cornuta, Crisia, 675, 676 

Crisidia, 675, 676, 801 

Sertularia, 675 
cribraria, Crisia, 679, 683, 803 
Criserpia johnstoni, 645 
Crisia, 618, 619, 648, 652, 653, 655, 674, 

675,678,686,687 

californica, 685 

cornuta, 675, 676 

cribraria, 679, 683, 803 

denticulata, 679, 684,685 

eburnea, 679, 680, 682, 801 

eburnea var. cribraria, 683 

eburnea forma denticulata, 684 

edwardsiana, 676 

elongata, 678, 684, 801 

franciscana, 677 

geniculata, 676, 677 

maxima, 679, 682, 803 

occidentalis, 677, 679, 680, 683, 801 

operculata, 678, 681, 801 

pacifica, 679, 680 

pugeti, 679, 684, 801 

punctata, 685 

serrata, 679, 680 

serrulata, 679, 682, 685, 803 
Crisidia, 674, 675, 676 

cornuta, 675, 676, 801 

edwardsiana, 676 

franciscana, 677 

gracilis, 677 
Crisiidae, 614, 615, 674, 686 
Crisina serrata, 679 
Crisinidae, 687 
Crisulipora, 614, 641, 674, 675, 685, 686 

occidentalis, 685, 686, 803 
cryptopora, Ceriopora, 692 
Ctenostomata, 613, 726 
Cyclostomata, 613, 614 
cylindrica, Bientalophora, 670, 799 
Cylindroecium giganteum, 737 

papuense, 737 

repens, 747 
Cytisidae, 687, 690 
davenporti, Loxosoma, 760 
dawsoni, Tubulipora, 642, 643 
Defrancia, 716 

intricata, 707 

stellata, 716 
densa, Penetrantia, 753, 754, 819 
denticulata, Crisia, 679, 684, 685 



NO. 3 



INDEX 



829 



depressa, Diaperoecia (Stomatopora), 

645 
Diaperoecia, 627, 628, 629, 636, 638, 640, 

641, 642, 646, 648, 667, 668, 670 

californica, 641, 642, 643, 793 

capitata, 668 

claviformis, 641, 642, 647, 791 

(Entalophora) capitata, 645 

(Entalophora) clavata, 645 

(Entalophora) vancouverensis, 645 

floridana, 641, 642, 644, 793 

intermedia, 641, 642, 646, 799 

intricata, 642, 643 

johnstoni, 641, 642, 645, 647, 793 

major, 621 

meandrina, 635, 641 

radicata, 644 

rugosa, 644 

(Stomatopora) depressa, 645 

(Stomatopora) expansa, 645 

striatula, 641 

subpapyracea, 641 

(Tubulipora) labiata, 645 

(Tubulipora) striata, 645 
Diaperoeciidae, 628, 686 
Diastopora, 627, 628, 639, 694 

catillus, 637 

cervicornis, 628 

foliacea, 627 

lactea, 639 

meandrina, 635 

obelia, 640 

patina, 631 

sarniensis, 628, 632 
Diastoporidae, 618, 627, 639 
diastoporides, Alecto, 624 

Oncousoecia, 624, 625, 638, 791 

Stomatopora, 624, 625 
dichotoma, Alecto, 619 

Filifascigera, 671 
digitata, Supercytis, 691 
dilatans, Alecto, 624 
dilatata, Farrella, 737 
Diplopora, 639 
Diplosolen, 627, 628, 629, 639, 641 

obelium, 640, 805 

obelium var. arctica, 640 
disciforme, Alcyonidium, 727, 730, 813 
Discocytis, 690 

californica, 690, 797 

canadensis, 690, 691 
Discopora hispida, 708, 710 

meandrina, 718 
Discoporella, 706 

californica, 704, 711 

canaliculata, 702 

fimbriata, 709 

hispida, 710 

novae-zelandiae, 705 

pristis, 718 

radiata, 714 



umbellata, 736 

verrucaria, 703 
discreta, Ascopodaria, 766 

Barentsia, 764, 765, 766, 767, 768, 
823 
Disporella, 700, 701, 708, 716, 717, 719 

alaskensis, 709, 712, 715, 809 

astraea, 709, 719, 811 

californica, 709, 711,807 

fimbriata, 709, 809 

hispida, 709, 710, 809 

octoradiata, 709, 718, 809 

ovoidea, 709, 713,714, 809 

separata, 709, 717, 807 

spinulosa, 709, 710 

stellata var. pacifica, 709, 716, 811 
Disporellidae, 700, 701 
distans, Amathia, 741, 817 
eburnea, Crisia, 679, 680, 682, 801 

Sertularia, 678, 682 

Stomatopora, 619 
eburnea var. cribraria, Crisia, 683 
eburnea forma denticulata, Crisia, 684 
echinata, Pedicellina, 763 
Ectoprocta, 613,726 
edwardsiana, Bicrisia, 676, 801 

Crisia, 676 

Crisidia, 676 
egregia, Tubulipora, 649, 655, 793 
(Electra) hastingsae, Membranipora, 

774 
elongata, Crisia, 678, 684, 801 

Farrella, 750, 819 

Hippuraria, 749 

Laguncula, 750 

PI atone a, 664, 797 

Triticella, 749, 819 
Emballotheca stylifera, 777, 821 
(Emballotheca) stylifera, Schizoporella, 

777 
Entalophora, 617, 667, 669 

capitata, 668 

cellarioides, 667 

clavata, 669 

proboscideoides, 668, 669, 799 

raripora, 669 

regularis, 670 

symmetrica, 667, 799 

vancouverensis, 669 
(Entalophora) capitata, Diaperoecia, 

645 

clavata, Diaperoecia, 645 

regularis, Bientalophora, 671 

vancouverensis, Diaperoecia, 645 
Entalophoridae, 617, 666, 667 
enteromorpha, Alcyonidium, 727, 731, 

813 
Entoprocta, 759 
Escharella porifera forma cancellata, 

778 

stylifera, 777 



830 



INDEX 



VOL. 14 



Escharoides jacksoni, 780, 821 
(Escharoides) jacksoni, Peristomella, 

780 
eudesii, Pelagia, 690 
expansa, Diaperoecia (Stomatopora), 

645 

Platonea, 663, 797 
Farrella, 750 

dilatata, 737 

elongata, 750, 819 

gigantea, 737 

pedicellata, 748 

repens, 750 
fasciculata, Filifascigera, 672 

Proboscina, 672 

Stomatopora, 672, 673 

Vesicularia, 739, 740,815 
fasciculifera, Tubulipora, 651 
Fasciculipora, 617, 641, 648, 664 

pacifica, 665, 666, 799 

ramosa, 665, 666 
Fascigera, 665 
Fascigeridae, 665 
Filicrisia, 675, 676 

franciscana, 677, 678, 803 

geniculata, 677, 678, 803 
Filifascigera, 671, 673 

clarionensis, 672, 673, 797 

dichotoma, 671 

fasciculata, 672 

parvipora, 671 

pluripora, 671 

robusta, 671 
fimbria forma pulchra, Tubulipora, 653 
fimbriata, Discoporella, 709 

Disporella, 709, 809 

Lichenopora, 702, 709 
flabellaris, Tubipora, 657 

Tubulipora, 649, 657, 795 
flabellata, Bugula, 776, 777 
flabellata acuminata, Bugula, 776, 821 
flava, Triticella, 748 
flemingii var. solida, Membranipora, 

775 
flexuosa, Idmonea, 653 

Tubulipora, 649, 653, 801 
floridana, Diaperoecia, 641, 642, 644, 

793 
Flustra hispida,732 

trifolium, 775 

tubulosa, 734 
Flustrella, 732 

corniculata, 732, 813 

gigantea, 733,815 
Flustrellidae, 726, 732, 734 
foliacea, Diastopora, 627 
franciscana, Crisia, 677 

Crisidia, 677 

Filicrisia, 677, 678, 803 
frondiculata, Hornera, 688 
Frondipora, 641, 664, 671, 673 



Frondiporidae, 618, 665, 671 
Gattyana cirrosa, 761 
gelatinosum, Alcyonium, 727 
geniculata, Barentsia, 765, 769, 823 

Crisia, 676, 677 

Filicrisia, 677, 678, 803 
gigantea, Farrella, 737 

Flustrella, 733, 815 
giganteum, Cylindroecium, 737 
glabra, Pedicellina, 763 
Gonopodaria, 764 
Gonypodaria nodosa, 765 

ramosa, 767 
gorbunovi, Barentsia, 765, 767, 823 
gothica, Hincksina, 774, 821 
gracilis, Ascopodaria, 765 

Barentsia, 765, 823 

Bovverbankia, 743, 744, 817 

Crisidia, 677 

Pedicellina, 765 
gracilis var. aggregata, Bovverbankia, 

744,817 
gracilis nodosa, Barentsia, 765 
granulata, Alecto, 619 

Scleroplax, 749 

Stomatopora, 619, 789 
grignonensis, Lichenopora, 702, 703 
grimaldii, Mesenteripora, 634 

Plagioecia, 630, 634, 635, 791 
harmeri, Vesicularia, 740 
hastingsae, Bathysoecia, 659, 660, 797 

Membranipora, 774 

Membranipora, (Electra), 774 
Heteropora, 613, 692, 693, 695, 696, 697, 

698,699 

alaskensis, 695, 805 

magna, 693, 695, 805 

pacifica, 693, 694, 695, 696 

pacifica var. alaskensis, 668, 695, 
696 

pelliculata, 693, 694, 695, 696 
Heteroporidae, 615, 692, 699 
Heteroporina, 615, 692 
Hincksina gothica, 774, 821 

nigrans, 775 
Hippodiplosia cancellata, 778, 821 

reticulato-punctata, 779 
Hippuraria elongata, 749 
hirsuta, Pedicellina, 763 
hispida, Discopora, 708, 710 

Discoporella, 710 

Disporella, 709, 710, 809 

Flustra, 732 

Lichenopora, 710 
holdsworthi, Lichenopora, 706 
Hornera, 613, 687, 688 

frondiculata, 688 

pectinata, 688, 689, 803 

pinnata, 689, 803 
Homeridae, 614, 687, 688 
Hydra verticillata, 742 



NO. 3 



INDEX 



831 



hystricis, Clavopora, 735 

Idmidronea atlantica van flexuosa, 654 

Idmonea atlantica var. flexuosa, 653, 

654 

atlantica var. tenuis, 653, 654 

californica, 642, 643 

flexuosa, 653 

milneana, 644 

palmata, 642, 643 
(Idmonea) atlantica, Tubulipora, 654 
imbricata, Bowerbankia, 743, 817 

Sertularia, 743 
Immergentia, 752 

californica, 752, 753, 819 
Immergentiidae, 751, 752 
incrassata, Proboscina, 623, 791 

Stomatopora, 623 

Tubulipora (Proboscina), 623 
intermedia, Diaperoecia, 641, 642, 646, 

799 

Tubulipora, 646 
intricaria, Pustulopora, 641 
intricata, Defrancia, 707 

Diaperoecia, 642, 643 

Lichenopora, 701, 702, 707, 811 
irregularis, Radiopora, 718 
jacksoni, Escharoides, 780, 821 

Peristomella (Escharoides), 780 

Smittia, 780 
johnstoni, Criserpia, 645 

Diaperoecia, 641, 642, 645, 647, 793 

Stomatopora, 645 
Kamptozoa, 759 
labiata, Diaperoecia (Tubulipora), 645 

Mucronella, 781 
lactea, Diastopora, 639 

Plagioecia, 639 
Lagenella repens, 750 
Laguncula elongata, 750 
lamellifera, Proboscina, 623, 791 
laxa, Barentsia, 764, 770 
lendigera, Sertularia, 740 
Lepralia microstoma, 781 

minuta, 780 
Lichenopora, 613, 699, 700, 701, 707, 

708,714,717,719 

bullata, 718 

buskiana, 701, 704, 712,807 

californica, 704, 711 

canaliculata, 702, 703,811 

fimbriata, 702, 709 

grignonensis, 702, 703 

hispida, 710 

holdsworthi, 706 

intricata, 701, 702, 707, 811 

raagnifica, 718 

novae-zelandiae, 701, 702, 705, 807 

octoradiata, 718,719 

radiata, 705,706, 713, 714 

turbinata, 701 

verrucaria, 702, 703, 807 



Lichenoporidae, 615, 700 
liliacea, Tubulipora, 649, 661 
lobulata, Tubularia, 658, 660 

Tubulipora, 624, 658, 659, 660, 661 
Loxocalyx, 760, 761 
Loxosoma, 760 

davenporti, 760 

raja, 761 
Loxosomatidae, 759, 760 
macropus, Ascopodaria, 769, 770 
Madrepora, 699 

verrucaria, 703 
magna, Heteropora, 693, 695, 805 

Tretocycloecia, 693 
magnifica, Lichenopora, 718 
major, Alecto, 621 

Diaperoecia, 621 

Oncousoecia, 621 

Proboscina, 621, 789 

Stomatopora, 621 
mammillatum, Alcyonidium, 727, 729, 

813 
mammillatum var. disciforme, 

Alcyonidium, 730 
maxima, Crisia, 679, 682, 803 
meandrina, Diaperoecia, 635, 641 

Diastopora, 635 

Discopora, 718 

Mesenteripora, 633, 634, 635 

Plagioecia, 630, 635, 791 
Mecynoeciidae, 628 
Melobesia radiata, 714 
membranacea form serrata, 

Membranipora, 774 
Membranipora annae, 774 

(Electra) hastingsae, 774 

flemingii var. solida,775 

hastingsae, 774 

membranacea form serrata, 774 

serrata, 774 

serrilamella, 774 

solida, 775 

trifolium, 775 
Mesenteripora, 627, 628 

grimaldii, 634 

meandrina, 633, 634, 635 

michelini, 628 
michelini, Mesenteripora, 628 
Microecia, 627, 628, 632 

sarniensis, 632 

tubiabortiva, 636 
(Microecia) tubiabortiva, Plagioecia, 

636 
Microporella arctica, 779, 821 

ciliata, 779 

ciliata var. arctica, 779 
microstoma, Lepralia, 781 

Mucronella, 781, 821 
milneana, Idmonea, 644 
minuta, Lepralia, 780 

Porella, 780, 821 



832 



INDEX 



VOL. 14 



misakiensis, Ascopodaria, 766 

Barentsia, 766 
Mucronella labiata, 781 

microstoma, 781, 821 
Myosoma, 761, 762 

spinosa, 762, 823 
Myriozoum, 670 
mytili, Alcyonidium, 727 
neocomiensis, Reptotubigera, 662 
nigrans, Hincksia, 775 
nitens, Buskia, 7+6, 747, 748, 819 
nodosa, Gonypodaria, 765 
Nolella, 737 

stipata, 737, 815 
Nolellidae, 736 
novae-zelandiae, Discoporella, 705 

Lichenopora, 701, 702, 705, 806 
nutans, Pedicellina, 763 
obelia, Berenicea, 640 

Diastopora, 640 

Tubulipora, 639, 640 
obelium, Diplosolen, 640, 805 
obelium var. arctica, Diplosolen, 640 
occidentalis, Ascorhiza, 735 

Clavopora, 735,815 

Crisia, 677, 679, 680, 683,801 

Crisulipora, 685,686, 803 

Tubulipora, 650, 651, 652 
octoradiata, Disporella, 709, 718, 809 

Lichenopora, 718, 719 
Oncousoecia, 618, 619, 621, 624, 627,629, 

648 

abrupta, 626, 789 

canadensis, 625, 789 

diastoporides, 624, 625, 638, 791 

major, 621 

ovoidea, 626, 789 
Oncousoeciidae, 618 
operculata, Crisia, 678, 681, 801 
ovoidea, Disporella, 709, 713, 714, 809 

Oncousoecia, 626, 789 
Pachyegis brunnea, 774, 777, 821 

princeps, 777 
Pachystega, 614, 687 
pacifica, Crisia, 679, 680 

Fasciculipora, 665, 666, 799 

Heteropora, 693, 694, 695, 696 

Tubulipora, 649, 652, 795 
pacifica var. alaskensis, Heteropora, 668, 

695,696 
palmata, Anguinella, 738, 815 

Idmonea, 642, 643 
Paludicellea, 736 
papuense, Cylindroecium, 737 
papuensis, Vesicularia, 740 
parasiticum, Alcyonidium, 727, 729, 813 

Alcyonium, 729 
parvipora, Filifascigera, 671 
patina, Diastopora, 631 

Plagioecia, 630, 631, 632, 633, 634, 
635,636,637,641,805 



Tubulipora, 630, 631 
pectinata, Hornera, 688, 689, 803 
pedicellata, Farrella, 748 

Triticella, 748, 819 
Pedicellina, 761, 762 

americana, 763 

australis, 766 

cernua, 763, 823 

echinata, 763 

glabra, 763 

gracilis, 765 

hirsuta, 763 

nutans, 763 
Pedicellinidae, 759, 761,771 
Pedicellinopsis, 764 
pedunculatum, Alcyonidium, 727, 730 

731,813 
Pelagia eudesii, 690 
pelliculata, Heteropora, 693, 694, 695, 

696 

Tretocycloecia, 693, 694 
pellucidum, Zoobotryon, 742 
pellucidus, Zoobotryon, 742 
Penetrantia, 753 

concharum, 754, 819 

densa, 753, 754, 819 

sileni, 755, 819 
Penetrantiidae, 751, 753 
Peristomella (Escharoides) jacksoni, 

780 
Peristomoecia, 620, 621 
phalangea, Tubulipora, 657 
Pherusa, 734 
Pherusella, 733, 734 

brevituba, 734, 815 
Pherusellidae, 726, 733 
philippsae, Reptotubigera, 662 
pinnata, Hornera, 689, 803 
Plagioecia, 627, 628, 629, 630, 638, 641 

ambigua, 629, 631,638, 791 

anacapensis, 630, 637, 791 

grimaldii, 630, 634, 635, 791 

lactea, 639 

meandrina, 630, 635, 791 

(Microecia) tubiabortiva, 636 

patina, 630, 631, 632, 633, 634, 635, 
636,637,641,805 

sarniensis, 631, 632, 633, 636, 637, 
805 

tortuosa, 630,633,793 

tubiabortiva, 630, 636, 805 
Plagioeciidae, 628 
Platonea, 648, 661, 662 

elongata, 664, 797 

expansa, 663, 797 

veleronis, 662, 664, 797 
pluripora, Filifascigera, 671 
polyoum, Alcyonidium, 727, 728, 813 

Sarcochitum, 727 
Porella alba, 780, 781 

compressa, 775 



NO. 3 



INDEX 



833 



concinna, 781 

minuta, 780, 821 
porifera forma cancellata, Escharella, 

778 
Porina ciliata, 779 
praelucida, Cheilopora, 774, 821 
princeps, Pachyegis, 777 
pristis, Discoporella, 718 
proboscideoides, Entalophora, 668, 669, 

799 
Proboscina, 618, 619, 620, 627, 646 

boryi, 620 

fasciculata, 672 

incrassata, 623, 791 

lamellifera, 623, 791 

major, 621, 789 

sigmata, 622, 789 
(Proboscina) incrassata, Tubulipora, 

623 
prominens, Berenicea, 627 
Pseudidmoneidae, 687 
pugeti, Crisia, 679, 684, 801 
pulchra, Tubulipora, 649, 652, 653, 795 
punctata, Crisia, 685 
Pustulopora intricaria, 641 
pustulosa, Borgiola, 697, 698, 699, 805 
radiata, Discoporella, 714 

Lichenopora, 705, 706, 713, 714 

Melobesia, 714 
radicata, Diaperoecia, 644 
Radiopora, 717, 718 

irregularis, 718 
raja, Loxosoma, 761 
ramosa, Barentsia, 764, 765, 767, 769, 

823 

Fasciculipora, 665, 666 

Gonypodaria, 767 

Reptotubigera, 662 

Terebripora, 751 
raripora, Entalophora, 669 
Rectangulata, 615, 699 
regularis, Actinopora, 628 

Bientalophora, 671 

Entalophora, 670 

(Entalophora), Bientalophora, 671 
repens, Cylindroecium, 747 

Farrella, 750 

Lagenella, 750 
Reptotubigera, 661, 662 

neocomiensis, 662 

philippsae, 662 

ramosa, 662 
reticulato-punctata, Hippodiplosia, 779 
retiformis, Alecto, 623 
robusta, Barentsia, 765, 768, 823 

Filifascigera, 671 
rugosa, Borgiola, 697, 699, 811 

Canuella, 697, 698 

Diaperoecia, 644 
Sarcochitum polyoum, 727 
sarniensis, Berenicea, 632 



Diastopora, 628, 632 

Microecia, 632 

Plagioecia, 631, 632, 633, 636, 637, 
805 
Schizoporella condylata, 777 

(Emballotheca) stylifera, 777 

stylifera, 777 
Scleroplax granulata, 749 
Semitubigera tuba, 650 
separata, Disporella, 709, 717, 807 
seriata, Buskia, 747, 819 
serpens, Tubulipora, 661 
serrata, Acanthodesia, 774 

Crisia, 679, 680 

Crisina, 679 

Membranipora, 774 
serrilamella, Membranipora, 774 
serrulata, Crisia, 679, 682, 685, 803 
Sertulariacornuta, 675 

eburnea, 678, 682 

imbricata, 743 

lendigera, 740 

spinosa, 739 

uva, 745 
setigera, Aeverrillia, 745, 817 

Buskia, 745 
sigmata, Proboscina, 622, 789 
sileni, Penetrantia, 755, 819 
Smittia jacksoni, 780 
Smittina, 778 

cancellata, 778 
Smittinidae, 778 
socialis, Buskia, 748 
solida, Membranipora, 775 
spinifera, Alcyonidium, 732 
spinosa, Myosoma, 762, 823 

Sertularia, 739 

Vesicularia, 740 
spinulosa, Disporella, 709, 710 
Steghorneridae, 687 
stellata, Defrancia, 716 
stellata var. pacifica, Disporella, 709, 

716,811 
Stenolaemata, 613, 614 
Stenostomata, 614 
stipata, Nolella, 737, 815 
stolonata, Coriella, 771, 823 
Stolonifera, 745 
Stomatopora, 618, 619, 620 

diastoporides, 624, 625 

eburnea, 619 

fasciculata, 672, 673 

granulata, 619, 789 

incrassata, 623 

johnstoni, 645 

major, 621 
(Stomatopora) depressa, Diaperoecia, 

645 
(Stomatopora) expansa, Diaperoecia, 

645 
striata, Diaperoecia (Tubulipora), 645 



834 



INDEX 



VOL. 14 



striatula, Diaperoecia, 641 
stylifera, Emballotheca, 777, 821 

Escharella, 777 

Schizoporella, 777 

Schizoporella (Emballotheca), 777 
subpapyracea, Diaperoecia, 641 
subrigida, Barentsia, 764, 770, 823 
Supercytis digitata, 691 
symmetrica, Entalophora, 667, 799 
Tecticavea, 713 
tegeticula, Triticella,750 
Terebripora, 751 

comma, 752, 819 

ramosa, 751 
Terebriporidae, 751 
Terebriporina, 751 
timida, Barentsia, 766 
tortuosa, Plagioecia, 630, 633, 793 
transversa, Tubulipora, 649 
Trepostomata, 613, 614 
Tretocycloecia, 693 

magna, 693 

pelliculata, 693, 694 
trifolium, Amphiblestrum, 775, 821 

Flustra, 775 

Membranipora, 775 
Triticella, 748 

elongata, 749, 819 

flava, 748 

pedicellata, 748, 819 

tegeticula, 750 
Triticellidae, 748 
tuba, Semitubigera, 650 

Tubulipora, 649, 650, 651, 795 
tuba var. f asciculifera, Tubulipora, 

649,650,651,795 
tuberosa, Valkeria, 745, 817 
tubiabortiva, Microecia, 636 

Plagioecia, 630, 636, 805 

Plagioecia (Microecia), 636 
Tubipora flabellaris, 657 
Tubularia lobulata, 658, 660 
Tubulipora, 629, 638, 639, 641, 646, 648, 

662, 666, 699 

admiranda, 649, 656, 795 

atlantica, 654 

atlantica var. fiexuosa, 654 

concinna, 649, 655, 793 

dawsoni, 642, 643 

egregia, 649, 655, 793 

f asciculifera, 651 

fimbria forma pulchra, 653 

flabellaris, 649, 657, 795 

fiexuosa, 649, 653, 801 



(Idmonea) atlantica, 654 

intermedia, 646 

liliacea, 649, 661 

lobulata, 624, 658, 659, 660, 661 

obelia, 639, 640 

occidentalis, 650, 651, 652 

pacifica, 649, 652, 795 

patina, 630, 631 

phalangea, 657 

(Proboscina) incrassata, 623 

pulchra, 649, 652, 653, 795 

serpens, 661 

transversa, 649 

tuba, 649, 650, 651, 652, 795 

tuba var. fasciculifera, 649, 650, 
651,795 
(Tubulipora) labiata, Diaperoecia, 645 
(Tubulipora) striata, Diaperoecia, 645 
Tubuliporidae, 614, 615, 618, 619, 639, 

647,648,658,662,665,667 
Tubuliporina, 614, 617 
tubulosa, Flustra, 734 
turbinata, Lichenopora, 701 
umbellata, Discoporella, 736 
Unicavea, 712 

californica, 704, 711,712 
Unitubigera, 657 
uva, Sertularia, 745 
Valkeria, 745 

tuberosa, 745, 817 

vidovici, 741 
Valkeriidae, 745 
vancouverensis, Diaperoecia 

(Entalophora), 645 

Entalophora, 669 
veleronis, Platonea, 662, 664, 797 
verrucaria, Discoporella, 703 

Lichenopora, 702, 703, 807 

Madrepora, 703 
verticillata. Hydra, 742 
verticillatum, Zoobotryon, 742, 817 
Vesicularia, 739 

fasciculata, 739, 740, 815 

harmeri, 740 

papuensis, 740 

spinosa, 740 
Vesiculariidae, 739 
Vesicularina, 739 
vidovici, Amathia, 741, 746, 817 

Valkeria, 741 
Zoobotryon, 742 

pellucidum, 742 

pellucidus, 742 

verticillatum, 742, 817 



INDEX 

Index for Orders, Divisions, Families, Genera, and Species 



Acamptostega, 617 
Adeona, 441 

tubulifera, 442 

violacea, 441 
fifc^. Adeonidae, 441 
Aetea, 11 

anguina, 11 

ligulata, 13 

recta, 12 

truncata, 12 
/Itij. Aeteidae, 11 
Aeverrillia, 745 

setigera, 745 
Aimulosia, 352 

palliolata, 353 

uvulifera, 352 
irr**'!). Alcyonidiidae, 727 
Alcyonidium, 727 

disciforme, 730 

enteromorpha, 731 

maramillatum, 729 

parasiticura, 729 

pedunculatum, 730 

polyoum, 727 
Alderina, 59 

brevispina, 60 

smitti, 59 
Akx. Alderinidae, 58 
Amastigia, 126 

biseriata, 127 

rudis, 127 
Amathia, 740 

convoluta, 740 

distans, 741 

vidovici, 741 
Amphiblestrum trifolium, 775 
Anasca, 9 
Anexechona, 96 

ancorata, 96 
Anguinella, 738 

palmata, 738 
Antropora, 51 

claustracrassa, 53 

granulifera, 52 

tincta, 54 
Aplousina, 46 

filum, 47 

major, 47 
Arachnopusiidae, 95 
Arthropoma, 333 

cecili, 333 

circinata, 334 
Articulata, 674 
Ascophora, 271 
Aspidostomidae, 114 



Barentsia, 764 

discreta, 766 

geniculata, 769 

gorbunovi, 767 

gracilis, 765 

ramosa, 767 

robusta, 768 

subrigida, 770 
Bathysoecia, 658 

bassleri, 659 

hastingsae, 660 
Beania, 169 

alaskensis, 171 

Columbiana, 173 

hirtissima, 172 

magellanica, 171 

mirabilis, 170 
Bicellariella, 152 

brevispina, 153 

stolonifera, 153 
Bicellariellidae, 151 
Bicrisia, 676 

edwardsiana, 676 
Bidenkapia, 75 

spitsbergensis, 76 

spitsbergensis var. alaskensis, 77 
Bientalophora, 670 

cylindrica, 670 
Borgiola, 696 

pustulosa, 698 

rugosa, 697 
Bowerbankia, 743 

gracilis, 743 

gracilis aggregata, 744 

irabricata, 743 
Brettia, 16 

pellucida, 17 

tubaeformis, 17 
Bugula, 153 

avicularia, 160 

californica, 156 

cucullifera, 159 

flabellata, 157 

flabellata acuminata, 776 

longirostrata, 156 

minima, 155 

mollis, 158 

neritina, 154 

pacifica, 155 

pugeti, 158 

uniserialis, 159 
Buskia, 746 

nitens, 747 

seriata, 747 
Buskiidae, 746 



[835] 



836 



INDEX 



VOL. 14 



Caberea, 129 
boryi, 129 
ellisi, 130 
Caleschara, 103 

mexicana, 104 
Callopora, 63 
armata, 64 
aurita, 65 

circumclathrata, 65 
corniculifera, 66 
craticula, 67 
exilis, 68 
horrida, 69 
inconspicua, 70 
lineata, 68 
verrucosa, 71 
whiteavesi, 70 
Calyptrostega, 699 
Camptostega, 674 
Cancellata, 687 
Carbasea, 39 

carbasea, 39 
Carnosa, 726 
Catenicellidae, 286 
Caulibugula, 160 
californica, 162 
ciliata, 161 
occidentalis, 161 
Cauloramphus, 55 
brunea, 56 
cymbaeformis, 57 
echinus, 56 
spiniferum, 55 
variegatum, 58 
Cellaria, 116 
diffusa, 117 
mandibulata, 116 
veleronis, 118 
Cellariidae, 116 
Celleporidae, 492 
Cellularina, 119 
Cerioporina, 692 
Chapperia, 89 

californica, 91 
condylata, 90 
frontalis, 92 
longispina, 93 
patula, 89 
varians, 94 
Chapperiidae, 88 
Cheilopora, 464 
praelonga, 464 
praelucida, 464 
Cheiloporinidae, 463 
Cheilostomata, 8 
Chorizopora, 279 

brogniarti, 279 
Clavopora, 735 

occidentalis, 735 
Clavoporidae, 735 
Codonellina, 422 



anatina, 422 
anatina ligulata, 423 
cribriformis, 424 
Coilostega, 99 
Coleopora, 291 

gigantea, 291 
Colletosia, 187 
bellula, 188 
radiata, 187 
Conopeum, 30 

commensale, 30 
reticulum, 31 
Copidozoum, 71 
planum, 73 
protectura, 73 
spinatum, 74 
tenuirostre, 72 
Coriella, 771 

stolonata, 771 
Corynoporella, 163 

spinosa, 163 
Costazia, 504 
costazi, 506 
nordenskjoldi, 508 
procumbens, 509 
robertsoniae, 507 
surcularis, 510 
ventricosa, 511 
Cranosina, 48 

colombiana, 48 
Crepidacantha, 478 
poissoni, 478 
setigera, 479 
Crepidacanthidae, 478 
Cribrilina, 177 
annulata, 177 
corbicula, 178 
Cribrilinidae, 174 
Cribrimorpha, 173 
Crisia, 678 

californica, 685 
cribraria, 683 
denticulata, 685 
eburnea, 682 
elongata, 684 
maxima, 682 
occidentalis, 680 
operculata, 681 
pugeti, 684 
punctata, 685 
serrulata, 679 
Crisidia, 675 

cornuta, 675 
Crisiidae, 674 
Crisulipora, 685 

occidentalis, 686 
Cryptosula, 470 

pallasiana, 470 
Ctenostomata, 726 
Cupuladria, 33 
canariensis, 33 



VOL. 14 



INDEX 



837 



Cyclicopora, 285 

longipora, 285 
Cyclicoporidae, 285 
Cycloperiella, 296 

rosacea, 297 
Cyclostomata, 613 
Cylindroporella, 303 

tubulosa, 303 
Cystisella, 434 

bicornis, 435 

saccata, 434 
Cytisidae, 690 
Dakaria, 325 

apertura, 326 

biserialis,329 

dawsoni, 326 

ordinata, 327 

pristina, 328 

sertata, 329 
Dendrobeania, 165 

curvirostrata, 166 

laxa, 167 

lichenoides, 167 

longispinosa, 168 

multiseriata, 169 

murrayana, 165 

simplex, 169 
Desmacystis, 32 

sandalia, 32 
Diaperoecia, 640 

californica, 642 

claviformis, 647 

floridana, 644 

intermedia, 646 

johnstoni, 645 
Diastoporidae, 627 
Diatosula, 311 

californica, 312 
Diplosolen, 639 

obelium, 640 
Discocytis, 690 

californica, 690 

canadensis, 691 
Discoporella, 112 

umbellata, 113 
Disporella, 708 

alaskensis, 715 

astraea, 719 

californica, 711 

fimbriata, 709 

hispida, 710 

octoradiata, 718 

ovoidea, 713 

separata, 717 

stellata var. pacifica, 716 
Doryporella, 83 

alcicornis, 84 

spathulifera, 84 
Ectoprocta, 726 
Electra, 35 

anomala, 36 



bellula var. bicornis, 38 

biscuta, 37 

crustulenta, 35 

crustulenta var. arctica, 36 

hastingsae, 38 
Electrinidae, 34 
Ellisina, 49 

levata, 50 
Emballotheca, 322 

altimuralis, 324 

latifrons, 323 

obscura, 323 

stylifera, 777 
Enantiosula, 468 

manica, 469 

plana, 469 
Entalophora, 667 

capitata, 668 

proboscideoides, 668 

symmetrica, 667 
Entalophoridae, 666 
Entoprocta, 759 
Epistomiidae, 150 
Escharoides, 372 

jacksoni, 780 

praestans, 372 
Eucratea, 17 

loricata, 17 
Euritina, 114 

arctica, 114 
Eurystomella, 389 

bilabiata, 389 
Eurystomellidae, 389 
Euteleia, 288 

evelinae, 289 
Exechonella, 95 

antillea, 95 
Exochellidae, 372 
Farciminariidae, 119 
Farrella, 750 

elongata, 750 
Fasciculipora, 664 

pacifica, 665 
Fenestrulina, 387 

malusi, 387 

malusi var. umbonata, 388 
Figularia, 189 

hilli, 190 
Filicrisia, 676 

franciscana, 677 

geniculata, 677 
Filifascigera, 671 

clarionensis, 672 

fasciculata, 672 
Floridina, 101 

antiqua, 102 
Flustrella, 732 

corniculata, 732 

gigantea, 733 
Flustrellidae, 732 
Flustridae, 38 



838 



INDEX 



VOL. 14 



Frondiporidae, 671 
Gemelliporella, 359 
aviculifera, 360 
globulifera, 359 
inflata, 360 
Gemelliporidra, 337 
colombiensis, 338 

lata, 337 
Gemelliporina, 357 

monilia, 358 
Gigantoporidae, 303 
Harmeria, 281 

scutulata, 282 
Hemicyclopora, 439 > 

polita, 440 
Heteropora, 692 

alaskensis, 695 

magna, 693 

pacifica, 694 
Heteroporidae, 692 
Heteroporina, 692 
Hiantoporidae, 97 
Hincksina, 41 

alba, 41 

gothica, 774 

minuscula, 46 

nigrans, 42 

pacifica, 43 

pallida, 45 

polacantha, 46 

velata, 44 
Hincksinidae, 40 
Hincksipora, 282 

spinulifera, 283 
Hippaliosina, 475 

costifera, 476 

inarmata, 476 

rostrigera, 475 
Hippodiplosia, 338 

americana, 339 

cancellata, 778 

insculpta, 341 

pertusa, 340 

reticulato-punctata, 340 
Hippomenella, 363 

flava, 364 
Hippomonavella, 365 

longirostrata, 365 

parvicapitata, 366 
Hippopleurifera, 301 

mucronata, 301 
Hippopodina, 292 

californica, 293 

feegeensis, 292 
Hippopodinella, 467 

adpressa, 467 

turrita, 468 
Hippoporella, 348 

gorgonensis, 348 
hippopus, 350 

nitescens, 350 



rimata, 351 
Hippoporidra, 354 
granulosa, 357 
janthina, 355 
spiculifera, 356 
Hippoporina, 344 
ampla, 347 
contracta, 346 
porcellana, 344 
tuberculata, 346 
Hippoporinidae, 343 
Hippothoa, 276 

divaricata, 278 
expansa, 279 
flagellum, 278 
hyalina, 277 
Hippothoidae, 276 
Hippothyris, 363 

emplastra, 363 
Holoporella, 495 
albirostris, 497 
brunnea, 496 
hancocki, 499 
peristomata, 500 
quadrispinosa, 502 
tridenticulata, 498 
Hornera, 688 

pectinata, 688 
pinnata, 689 
Horneridae, 688 
Immergentia, 752 

californica, 753 
Immergentiidae, 752 
Inovicellata, 10 
Labioporella, 108 
sinuosa, 109 
Lacerna, 361 

fistulata, 362 
Lagenipora, 484 

admiranda, 491 
hippocrepis, 489 
lacunosa, 490 
marginata, 489 
mexicana, 486 
punctulata, 485 
socialis, 488 
spinulosa, 487 
Lepraliella, 452 
bispina, 453 
contigua, 452 
Lichenopora, 701 
buskiana, 704 
canaliculata, 702 
intricata, 707 
novae-zelandiae, 70j 
verrucaria, 703 
Lichenoporidae, 700 
Loxocalyx, 761 
Loxosoma, 760 

davenporti, 760 
Loxosomatidae, 760 



VOL. 14 



INDEX 



839 



Lunularlidae, 112 
Lyrula, 184 

hippocrepis, 184 
Malacostega, 14 
Mamillopora, 517 

cupula, 517 
Mamilloporidae, 517 
Mastigophora, 479 

pesanseris, 479 

porosa, 480 
Membranipora, 19 

annae, 774 

fusca, 25 

hastingsae, 29 

membranacea, 21 

pachytheca, 28 

perf ragilis, 24 

savarti, 27 

serrilamella, 22 

tenuis, 26 

tuberculata, 23 

villosa, 22 
Membraniporella, 174 

aragoi van pacifica, 174 

crassicosta, 176 

pulchra, 176 
Membraniporidae, 18 
Membraniporidra, 62 

porosa, 62 
Micropora, 105 

coriacea, 105 
Microporella, 375 

arctica, 779 

californica, 381 

ciliata, 377 

ciliata stellata, 378 

coronata, 386 

cribrosa, 380 

gibbosula, 386 

marsupiata, 382 

pontifica, 383 

setiformis, 385 

tractabilis, 384 

urabonata, 378 

vibraculifera, 379 
Microporellidae, 375 
Microporidae, 100 
Microporina, 106 

borealis, 106 
Mollia, 60 

patellaria, 61 
Mucronella, 435 

connectens, 437 

labiata, 437 

major, 438 

microstoma, 781 

ventricosa, 436 
Myosoma, 762 

spinosa, 762 
Myriozoella, 515 

plana, 516 



Myriozoidae, 513 
Myriozoum, 513 

coarctatum, 513 

subgracile, 514 

tenue, 515 
Nellia, 119 

oculata, 119 

tenuis, 120 
Nolella, 737 

stipata, 737 
Nolellidae, 736 
Oncousoecia, 624 

abrupta, 626 

canadensis, 625 

diastoporides, 624 

ovoidea, 626 
Oncousoeciidae, 618 
Onychocella, 100 

alula, 101 
Pachyegis, 313 

brunnea, 315,777 

princeps, 313 
Pachystega, 687 
Paludicellea, 736 
Parasmittina, 411 

alaskensis, 419 

californica, 415 

collifera, 416 

crosslandi, 418 

fraseri, 419 

jeffreysi, 414 

spathulata, 415 

trispinosa, 412 

tubulata, 420 
Parellisina, 75 

curvirostris, 75 
Pedicellina, 762 

cernua, 763 
Pedicellinidae, 761 
Penetrantia, 753 

concharum, 754 

densa, 753 

sileni, 755 
Penetrantiidae, 753 
Petralia, 290 

japonica, 290 
Petraliidae, 289 
Pherusella, 734 

brevituba, 734 
Pherusellidae, 733 
Phidolopora, 447 

pacifica, 448 

pacifica van catalinensis, 449 
Phylactella, 481 

alulata, 483 

aperta, 482 
Phylactellidae, 481 
Plagioecia, 629 

ambigua, 638 

anacapensis, 637 

grimaldii, 634 



840 



INDEX 



VOL. 14 



lactea, 639 

meandrina, 635 

patina, 631 

sarniensis, 632 

tortuosa, 633 

tubiabortiva, 636 
Platonea, 661 

elongata, 664 

expansa, 663 

veleronis, 662 
Porella, 392 

acutirostris, 394 

Columbiana, 398 

compressa, 393 

concinna, 396 

minuta, 780 

patens, 397 

porifera, 395 
Posterula, 309 

sarsi, 310 
Proboscina, 620 

incrassata, 623 

lamellifera, 623 

major, 621 

sigmata, 622 
Pseudostega, 115 
Puellina, 185 

setosa, 186 
Ragionula, 310 

rosacea, 311 
Rectangulata, 699 
Reginella, 178 

furcata, 179 

mattoidea, 182 

mucronata, 180 

nitida, 181 

spitsbergensis, 182 
Reteporelllna, 445 

bilabiata, 445 

denticulata van gracilis, 446 
Reteporidae, 444 
Retevirgula, 85 

areolata, 87 

lata, 86 

tubulata, 86 
Rhamphostomella, 424 

bilaminata, 427 

cellata, 431 

costata, 426 

curvirostrata, 430 

fortissima, 427 

gigantea, 433 

hincksi, 428 

ovata, 432 

spinigera, 429 

townsendi, 430 
Rhynchozoon, 454 

bispinosum, 455 

grandicella, 459 

rostratum, 456 

spicatum, 460 



tuberculatum, 461 

tumulosum, 458 
Robertsonidra, 294 

oligopus, 295 
Rosseliana, 105 

rosselii, 106 
Savignyella, 288 

lafonti, 288 
Savignyellidae, 287 
Schizmopora, 492 

anatina, 493 

margaritacea, 494 
Schizolavella, 335 

vulgaris, 335 
Schizomavella, 330 

auriculata, 331 

auriculata acuta, 332 

auriculata ochracea, 331 

porifera, 332 
Schizoporella, 317 

cornuta, 320 

dissimilis, 321 

linearis van inarmata, 319 

trichotoma, 318 

unicornis, 317 
Schizoporellidae, 316 
Schizoretepora, 449 

tessellata, 450 
Schizotheca, 450 

fissurella, 451 

umbonata, 451 
Scruparia, 15 

ambigua, 16 
Scrupariidae, 15 
Scrupoceilaria, 130 

bertholetti, 133 

bertholetti van tenuirostris, 134 

californica, 135 

diegensis, 136 

ferox, 137 

harmeri, 138 

inarmata, 150 

macropora, 138 

mexicana, 139 

obtecta, 140 

panamensis, 141 

profundis, 142 

pugnax, 143 

regularis, 144 

scabra, 144 

scabra van paenulata, 145 

scruposa, 145 

spinigera, 146 

talonis, 147 

unguiculata, 148 

varians, 149 
Scrupocellariidae, 120 
Semihaswellia, 304 

sulcosa, 304 
Sessibugula, 163 

translucens, 164 



VOL.14 



INDEX 



841 



Smittia californiensis, 421 
Smittina, 399 

altirostris, 405 

arctica, 402 

bella, 403 

cordata, 407 

landsborovi, 400 

maccullochae, 405 

retif rons, 402 

smittiella, 404 

spathulifera, 401 
Smittinidae, 390 
Smittoidea, 408 

prolifica, 408 

reticulata, 409 

transversa, 410 
Steganoporella, 107 

cornuta, 107 
Steganoporellidae, 107 
Stephanosella, 367 

biaperta, 368 

bolini, 370 

vitrea, 369 
Stolonifera, 745 
Stomachetosella, 305 

abyssicola, 309 

cruenta, 306 

distincta, 308 

limbata, 307 

sinuosa, 306 
StomachetoselHdae, 305 
Stomatopora, 619 

granulata, 619 
Stylopoma, 336 

informata, 336 
Synnotura, 150 

aegyptiacum, 151 
Tegella, 77 

aquilirostris, 83 

arctica, 82 

armifera, 79 

magnipora, 80 

robertsonae, 81 

unicornis, 78 
Terebripora, 751 

comma, 752 
Terebriporidae, 751 
Terebriporina, 751 
Terminoflustra, 40 

membranaceo-truncata, 40 
Tetraplaria, 466 

veleroae, 466 
Thalamoporella, 110 

californica. 111 

gothica, 110 
Thalaraoporellidae, 110 
Trematooecia, 502 



hexagonalis, 503 

porosa, 503 
Tremogasterina, 98 

granulata van subspatulata, 98 
Tricellaria, 121 

erecta, 126 

gracilis, 124 

occidentalis, 122 

occidentaliscatalinensis, 122 

praescuta, 125 

pribilofi, 124 

ternata, 123 
Trigonopora, 442 

pacifica, 443 
Triticella, 748 

elongata, 749 

pedicellata, 748 
Triticellidae, 748 
Trypematella, 373 

umbonula, 373 
Trypostega, 280 

claviculata, 281 

venusta, 280 
Tubulipora, 648 

admiranda, 656 

concinna, 655 

egregia, 655 

flabellaris, 657 

flexuosa, 653 

pacifica, 652 

pulchra, 653 

tuba, 650 

tuba var. fascicuHfera, 651 
Tubuliporidae, 647 
Tubuliporina, 617 
Umbonula, 298 

alvareziana, 300 

arctica, 299 

patens, 298 
Umbonulidae, 298 
Valkeria, 745 

tuberosa, 745 
Valkeriidae, 745 
Veleroa, 473 

veleronis, 474 
Velumella, 103 

americana, 103 
Vesicularia, 739 

fasciculata, 739 
Vesiculariidae, 739 
Vesicularina, 739 
Vittaticella, 286 

elegans, 286 
Watersipora, 471 

cucuUata, 472 
Zoobotryon, 742 

verticillatura, 742