INVERTEBRATE ZOOLOGY

-.

INTRODUCTION
No picture of organisms which ignores their physical and organic
environment can be even approximately complete. Studies of dead animals
or their parts or even of living animals in the laboratory, valuable
and indispensable as they are, give but partial pictures. In the studies
here contemplated we seek a firsthand knowledge of living invertebrate
animals in their natural setting, their behavior and interrelations,
their distribution within the habitat, the influence of physical conditions on this distribution and the correlation between their structures
and their behavior patterns on the one hand and the places they occupy
in the environment on the other.
Field trips are naturally of prime importance in such studies. The
more time spent in actual study of animals in the field the better.
Under most circumstances these periods must each be confined to a part
of a day. Experience has amply proved, however, that continuous studies
over a period of days increases the values received out of all proportion to the time spent. Appendix A gives specific information with regard to field trips and the schedules of such trips during the spring
and summer courses at Berkeley.
Such a field study might be thought to require previous courses
designed to give the student a knowledge of the animals which make up
the faunas to be studied. Certainly the animals must be recognized and
known by name if their behavior and ecologic distribution is to be
studied. In practice it is usually necessary and perhaps more advantageous to combine this type of study with the field study.
This portion of the work involves the laboratory. It falls naturally into four phases (1) the study of those external characters of
the various animal types which are used for their classification and
identification, (2) preliminary practice in the use of keys for the
identification of some member of each group, (3) a study of demonstration sets previous to field trips to give a preliminary knowledge of
the important animals to be expected in the particular fauna, and (4)
the identification of animals collected on the field trips.

THE IDENTIFICATION OF INVERTEBRATE ANIMALS

A necessary preliminary to the study of any animal is the determination of its scientific name or authentic common name. This is
necessary if only to make possible the satisfactory designation of the
animal in conversation or in class exercises. A correct determination
is especially important, of course, if the animal is to be made the
subject of a scientific investigation, since in no other way can the
facts be made available to others.
Few students of zoology realize the difficulties involved in
identifying with certainty most species of animals, particularly the
species of many of the numerous groups known collectively as invertebrates. A relatively few species are readily recognized because of
distinctive color, pattern, or structure. Such an animal, for example,
is the striped shore crab, Pachygrapsus crass ipes, which is abundant
in rocky crevices above low-tide mark along the Pacific Coast. In many
1

LABORATORY AND FIELD TMT IN INVERTEBM!CE ZOOLOGY

other cases, however, identification can only be approximated by the

beginner, and in still others even the specialist will find difficulty
in making identification. These difficulties are aggravated on the
Pacific Coast by the fact that the study of many groups of invertebrates has been greatly neglected. During the period when systematic
work was the vogue in zoology, there were very few zoologists on this
coast. With the change in emphasis in zoology these have largely abandoned this field and few others have entered it. For a few groups such
as the Amphipoda, and the littoral copepods, the fauna of the Pacific
Coast is largely unknown. In other groups a considerable amount of work
has been done, but much remains to be done.
Some groups, such as the decapod Crustacea, the marine Mollusca and
the echinoderms, are fairly well known to systematists. Even here there
remains much need for complete and careful systematic revision and monographing. But when all this has been done there will be necessary still
another step if this information is to be available to students of zoology or biology not specialists in the particular groups, and also to the
intelligent laymen. This is the production of manuals of faunas, containing brief diagnostic descriptions and keys to the species, illustrated, if possible, and accompanied by careful definitions of terms.
Familiar works of this type are the floras of various regions and
in the various mammals for the identification of birds. A Manual of the
Common Invertebrate Animals, by H. S. Pratt (Blakiston), is an attempt
at such a manual for the identification of the invertebrate animals of
the United States. Because of the imperfect knowledge of the local
fauna, the first edition was of very little value for identification
of the invertebrates of the Pacific Coast. The new edition (Blakiston)
is better, but still reflects the difficulties arising from the scope
of such an attempt and the lack of available information with regard
to the invertebrates of the Pacific Coast.
Freshwater Bioloa, by Ward and Whipple, represents a more satisfactory attempt in this direction for the animals of the United States
which inhabit fresh water. Since many of these tend to be widely distributed, the Pacific Coast species are better covered. In addition,
this work includes valuable discussions of the biology of the different
groups, as well as directions for their collection and study. Here
again, however, there is great need for revisions of the various groups
and a new edition is to be hoped for. Every student or teacher of zoology should own this book when the new edition is available.
Seashore Animals of the Pacific Coast, by Johnson and Snook, represents the only attempt at a comprehensive treatment of our local marine
invertebrates. It is a valuable eontribution and it also should he in
the library of every student or teacher of zoology or biology located
on the Pacific Coast. Its excellent illustrations make possible the
definite identification of certain common,forms, but the absence of
keys or any comprehensive treatment of the various groups makes it impossible to use it as a manual. Indeed, the lack of completeness of our
knowledge makes it impossible to produce such a general manual at the
present time, and if the knowledge were available it could not be compressed within the limits of a single volume.
Between Pacific Tides, by Ricketts and Calvin, published by the
Stanford University Press, will be of great value to students of the

INTRODUCTION

invertebrates of the Pacific Coast and its reading is required of all

students in this course. While it will not serve as a manual for the
identification of animals it will give a valuable picture of faunas
characteristic of the major environments of the seashore in this area,
and its excellent illustrations in conjunction with references to classification lists will make possible the identification of many common
animals.
Fortunately we have a slowly increasing number of well-illustrated,
authentic monographs on separate groups of California invertebrates.
Such are: Schmitt,
University of Cali
Fauna of the Pacific Coasts of North. Central. and Northern South America (1940, University of Southern California Press); Fraser, Hydroids
of the.Pacific Coast (1937, The University of Toronto Press), and others.
Some of these are in need of revision and others are too technical to
be available to the student involved in such a study as this.
As mentioned in the preface, the Illustrated Key to Nest American
Pelecy~odGenera, by Keen and Frizzell (1939, Stanford University Press),
is an excellent example of what can and should be done in this field.
Students are encouraged to obtain and use this valuable work. Unfortunately it stands practically alone,and even it leaves the species undetermined.
It will be seen, therefore, that if the invertebrate fauna of a
given area of the Pacific Coast is to be used for field studies, it becomes necessary, as a temporary measure, at least, to develop keys to
aid in the identification of the species concerned. Such is the purpose
of the keys here given, which are tentative, in many cases requiring to
be corrected and amplified as our knowledge increases. It will be noted
that in numerous instances the determinations are only to genus, and in
some cases even only to the family. This may be because there is only
one species concerned in the area, and the burden of scientific names
to be learned may be thus reduced, or because either the knowledge is
lacking to carry the identifications further, or the identifications
are too difficult for any except the specialist.
It cannot be too strongly emphasized that keys are short-cuts and
often very misleading; that their function is merely to clear the way
to an approximation, and that identifications made by them, if to be
of scientific value, must be reinforced by comparisons with descriptions and illustrations in monographs, if such exist, or by comparisons
with authentic named specimens, or by submission to a specialist.
The first two methods are available in a course such as the one
for which these keys are designed. Comparison with named specimens
should be considered as a last resort, however, since such named collections are rarely available and the gaining of experience in the use
of keys and works of reference is one of the important opportunities
offered by the course.
A list of the publications and manuscripts most available for
verification or identifications of species of Pacific Coast invertebrates will be found at the end of this volume.

LABOR4TORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

THE USE OF SCIENTIFIC NAMES

A scientific name consists of the name of the genus (capitalized)

followed by the name of the species (rarely capitalized by zoologists
and then because it is derived from a proper noun) followed by the name
of the original describer. The name of the author is properly placed
in parentheses if the generic name used is not that used at the time of
the original description, thus Hemigrapsus nudus (~ana). This usage is
under discussion at present and it has not been followed for all groups
discussed in this text. For the sake of convenience, the name of the
describer may be omitted in our use of scientific names but should
always appear on properly labeled specimens.
The genus name may be used alone, the specific name never, unless
it has.already been used with the generic name on the same page. Both
generic and specific names should be underlined. This indicates italics
to the printer and is of value in picking out the scientific names in
manuscript.
Great pains should be taken to spell scientific names correctly.
The correlation between careless, unscientific work, and careless use
of scientific names is very high.

THE USE OF COMMON OR VERNACULAR NAMES

Common or vernacular names are convenient but have many disadvantages. The rule of priority fixes the correct scientific name, which is
universal. Common names are local since there are no rules to determine
which of many such names is the correct name, and the same name may be
applied to very different species or types in different regions, or by
different persons in the same region. Vernacular names of American
birds and certain other groups of vertebrates are relatively uniform
owing to the united action of the workers in these fields, but no such
authentic common names exist for invertebrates.
It is necessary to make a sharp distinction between these two types
of names and to use the vernacular name only after having connected it
with the scientific name. ThusHemigrapsus nudus will be known to or
identifiable by all zoologists, but the purple shore crab would have no
meaning or a different meaning in areas in whichH. nudus is not found
or even in certain parts of the range of that particular species.

LABORATORY STUDIES
In this section materials are presented for the study, classification, and identification of the invertebrate animals generally encountered in the progress of the course. This material is arranged under
the headings of the various phyla of invertebrates considered. In general, the consideration of each group includes (1)a laboratory-exercise
acquainting the student with the general characters of the animals and
explaining terms used in the identification keys, (2) a key or keys for
the identification of specimens, and (3) a list of genera and species
which have been encountered in the localities visited in the course,
giving for each the full scientific name if it has been definitely de.termined
A careful study of the external anatomy of many types in each group
as well as of the internal anatomy would be an ideal preparation for the
use of keys in the identification of animals. Time does not permit such
an approach within the limits of this course. Some knowledge of those
characters, chiefly external, which are of taxonomic significance and
are commonly used in keys and descriptions is, however, an absolutely
necessary prerequisite to any successful attempt at identification such
as will be necessary in the faunal surveys and ecological studies to be
initiated later in the course.
The rapid studies of various groups outlined below are designed to
furnish this knowledge of characters of taxonomic importance in so far
as the very brief time available makes possible and must be entered upon
with this purpose in mind. They must by no means be considered as an attempt at any complete study of the external anatomy. A copy of some text
dealing with the group concerned will be a valuable adjunct to such
studies. Some such texts are (1) Invertebrata, by Borradaile and Potts;
(2) A Text Book of Zoolom, Parker and Haswell, Vol. 1; (3) Colle~e
Zooloas, by Hegner
Each such study should be followed by the use of keys and the consultation of specific descriptions in the-publications listed at the
end.

THE TAXONOMY OF THE INVERTEBRATES

The invertebrates include a large number of diverse types of animals including all but one of the phyla of the animal kingdom and even
some of the branches of that one phylum (Chordata). The classification
of this assemblage is therefore of cardinal importance in the study of
the invertebrates. It is, however, also a matter of considerable controversy. The student must be prepared to find that different writers
use different systems of classification. These differences are chiefly
of two sorts: first, the using of different names for the same group,
and seoond, the placing of the same group in different systematic categories, as, for example, the designation of a group as a class by one
writer, but as an order by another. Discrepancies of the second sort
are so abundant that in some cases it is advisable not to have a fixed
concept of the taxonomic rank of a certain group, but rather to remem-

LAI3ORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

ber only that it is a part of a certain superior group and can be divided into a number of subordinate groups. Thus it is not so important
that one decides whether the Crustaceae is a subphylum or a class as
it is that one knows that it is a major division of the Arthropoda and
that it includes the Malacostraca, Copepoda, etc. Appendix B gives a
synopsis of the classification used in this course. This agrees most
closely with that used by Borradaile and Potts, Invertebrata, and in
most features with that of Hyman, Invertebrates.
DIRECTIONS FOR THE STUDY OF THE VARIOUS GROUPS OF
INVERTEBRATES
PHYLUM PORIFERA (THE SPONGES)
References: de Laubenfels, 1932; Hyman, 1940
Terminolo~sand Instructions for Studp.-The sponges are structurally primitive animals, very ancient as a group and only distantly
related to the animals of other groups. They
are multicellular but have attained relatively
little histological differentiation either as
regards cell types or their integration in
layer and organs. Organs cen hardly be said
to exist, and whole tissues characteristic of
higher animals are lacking, such as nervous
and muscular tissues. Specialized gland cells
also are few or lacking. Indeed the body consists (fig. 2 , ~ )of a rather loose mesenohyme,
which produces the skeletal elements, covered
by external and internal layers of cells so
loosely organized as to be called epithelium
only by courtesy.
The sponges are in contrast to all higher
animals also in that; they lack a true mouth,
the body being pierced by numerous minute incurrent pores connecting with canals and cavities through which water flows to reach the
outside by larger openings figs. 1, 2) known
as oscula (singular osculum These features,
Fig. 1.-Surface of poras
well as the characteristic digestive cells,
t i o n of encrusting sponge
showlug oscula
(afier Guber- the choanocytes, are well seen in a schematic
diagram of the simplest type of sponge (fig.
let).
2,A) such as is exemplified by Leuoosolenia in
this region. In the complicated sponges which constitute the vast majority of those encountered at the seashore and all those of fresh water,
the choanocytes occur in small, scattered, spherical chambers (fig. 2,D).
Sponges are attached, non-locomotor, nearly always colonial, and
typically irregular in form, conforming to the surfaces on which they
grow (fig. 1). In all save the simplest sponges the colony is so closely
integrated that it is difficult or impossible to designate the individuals constituting it or at least to determine the limits of the individuals (fig. 1).
Except for a very few parasitic species all sponges are plankton

I.

feeders, the minute organisms and organic particles in the water currents being seized and taken in by the individual choanocytes. The
skeletal system of calcareous or silicious spicules serves as a protection against predators and the unpleasant odor characteristic of a

Fig. 2.-Diagrams
of types of sponge structure (after Eiyman, 1940). A. Diagrammatic
vertical seotion of simplest type of sponge (ascanoid type seen in ~eucosoleniq), showing cellular elements. B-D, sections of one wall; B, ascanoid; C, primitive syoonoidj
D, developed syconoid such as is seen in Grantial E, leuconoid, that of all local encrusting sponges. Choanocyte layer shown in heavy black. Arrows indicate course of
ter.

--

number of sponges would seem to indicate the presence of distasteful

chemical secretions. They lack, however, devices to protect against
intertidal exposure and hence they are confined to protected situations at, near, or below low-tide level. There they compete with other
plankton feeders such as the Bryozoa and tunicates as also with the
micropredators such as the hydroids and corals.
Until recently the identification of California marine sponges has
not been possible. De Laubenfels' treatise (1932) makes this possible,
but it requires study of spicule types and other internal characters
and is not usually feasible for the average student. A key to the more
common littoral species of central California, based on de Laubenfels'
descriptions, is given below.
See demonstration specimens of the calcareous sponges, Leucosolenia (asconoid type, fig. ~ , A , B )and Grantia (syconoid type, fig. 2,
C,D) and understand the arrangements of m t s characteristic of such
sponges.

LABORATORY AND FIELD TEXT I N INVERTEBRATE ZOOLOGY

Study pieces of a l o c a l encrusting sponge. Note i t s consistency.

Is i t hard and stony, s o f t and f r a g i l e , o r l e a t h e r y and spongy? What
i s t h e nature of t h e surface? Note t h e shape, s i z e , and d i s t r i b u t i o n
of the oscula and t h e pores. The oscula a r e t h e l a r g e r , excurrent openings ( f i g s . 1, 2,A); t h e pores o r o s t i a a r e t h e very small incurrent
openings. Remove a small but c h a r a c t e r i s t i c portion, mount on a s l i d e
i n a drop of water. Tease a p a r t s u f f i c i e n t l y t o make a c l e a r mount.
Examine under the microscope t o make out t h e types of s p i c u l e s .
Spicules with a s i n g l e point a r e termed monactinal ( f i g . 3,A,C,I),
those with two, d i a c t i n a l ( f i g . 3,H,J). Monaxon spiculeb a r e s t r a i g h t
rods, needles, clubs, e t c . Triaxons have t h r e e projecting points; i f
a l l a r e approximately equal they a r e t r i r a d i a t e o r t r i a e n e s ( f i g . 3,
E). Tetraxons may be quadriradiates o r c o n s i s t of t h r e e short rays
r a d i a t i n g from a long a x i s . Hexactinal s p i c u l e s have s i x ( o r f i v e )
rays r a d i a t i n g from a common center.

Fig. 3.-Types
of sponge spicules (after de Laubenfels, 1932). A, C,
subtylostyles; B, spiny-headed tylote; D, chelas; E, triaene; F, toxa;
G, sigma; H, acanthostyle; I, spiny subtylostyle; J, oxea.

S i l i c e o u s spicules, found i n a l l save the simple sponges, include,

usually, two very d i f f e r e n t types, l a r g e r menascleres which make up t h e
t r u e skeleton, and t h e smaller microscleres i n t h e s o f t e r t i s s u e s ( c f .
f i g . 3,A.B with D,F, and G ) . The megasclera may be
or
monactinal, c a l l e d r e s p e c t i v e l y t r i a e n e s ( f i g . 3,E), t s l o s t ~ l e s ,s u b t y l o s t y l e s ( f i g . ~ , A , B , I ) . Oxeas ( f i g . 3 , ~ a) r e s t r a i g h t o r somewhat
curved, more o r l e s s fusiform and gradually pointed a t t h e ends. Subt y l o s t y l e s o r monactinals have one of t h e ends rounded ( f i g . 3,A).
These p r i n c i p a l s p i c u l e s may be s p a r s e l y spiny a s i n Lissodendoryx
( f i g . 4 ) o r extensively spined acanthostyles ( f i g . 3,H). Tylotes a r e
d i a c t i n a l s with both heads rounded a s i n Plocamia ( f i q . 3,B).
Microscleres may be sigmas (S- o r C-shaped, f i g . 3,G) a s i n Lissodendoryx ( f i g . 4 ) ; chelas (dumbbell-shaped, f i g s . 3,D and 4 ) ; toxas
(broad V-shaped, f i g . 3,F).

a-,
a-,

10

LABORATORY AND FIELD TEXT I N INVERTEBRATE ZOOLOGY

......
........................

10
9. Usually with small c r a t e r s projecting above matrix
9. Vithout c r a t e r s
11
10. Lavender i n l i f e ; extensively encrusting surfaces a t low-tide
levels; height and s i z e of c r a t e r s variable: principal
spicules usually forming d e f i n i t e triangular pat terns; oxeas
only ( f i g 3, J )
Hal i clona perm01 1 is (p 120 )
10. Color i n l i f e not lavender
Haliclona spp.
11. Principal spicules tylotes and s t y l e s ( f i g . 4). (With a strong
sulphurous odor)
Lissodendoryx noxiosa (p. 76)

. . . . . .. .. .. .. .. . . . . . . .

........

Fig. 4.-The

spicules of

I J Y t o d w soxiosa. Frcn

r b m donnrrrdr tylotee ,
styles of three types (smooth,
apnnely spiny, and verg slender) , arouate chelae ( l e f t ) ,
and oontort sigma (after de
Laubenfels )

11. Principal spicules oxeas ( f i g . 3 , J ) . Orange t o green i n l i f e ;

usually massive; resembling crumb of bread when broken; spicul a t ion very irregular; "the bread spongen
Halichondria panicea (p. 56)
11. Principal spicules subtylostyles ( f i g . 3,A, C ) Closely and
densely surrounding objects so a s t o appear massive; usually
giving off burnt-umber color i n alcohol; tough; solid; finegrained
Prosuber i tes s isyrnus (p 54 )

.................

.............

LIST OF SPONGES I N KEY

Phylum PORIFERA
Class CALCAREA

M
BM
D *M

Leuconi a
Leuoosolenia eleanor Urban
Rhabdodemlla nutt ingi Urban

'1n t h i s and o t h e r l i s t s of species throughout t h i s s e c t i o n t h e

following symbols a r e used (placed before t h e names of specieej: D ,
found a t Dillon Beach, Marin County o r v i c i n i t y ; TB, found i n upper
end of Tomales Bay near Dillon Beach ( s i n c e the Tomales Bay region
i s r e l a t i v e l y new t e r r i t o r y many more epecies w i l l be found than
a r e indicated i n our l i s t a s occurring t h e r e ) ; M, found a t Moas
Beach, San Mateo County, o r other ocean beaches of t h e v i c i n i t y ; B,
found i n San Francisco Bay; F, found I n f r e s h water; T, t e r r e s t r i a l .
For f u r t h e r information concerning t h e l o c a t i o n of s p e c i e s i n the
San Francisco Bay Region, consult t h e check lists f o r t h e varioue
localities.
i n d i c a t e s t h a t the epecies i e r e l a t i v e l y abundant
The symbol
a t t h e l o c a t i o n indicated. It must be understood t h a t t4ese marks of
abundance cannot always be r e l i e d upon, owing t o f a u l t y o r incomp l e t e information, o r t o changes i n the r e l a t i v e abundance of species
from year t o year o r from locaAity t o l o c a l i t y .

Class DEMOSPONGIAE
Esperi opsis original is de Laubenfels
Fi cul i na
Hal i ohondria panioea Pallas
Haliclona penoll is Bowerbank
Haliolona spp.
Isociona 1 i thophoenix de Iaubenfels
Li ssodendoryx noxi osa de Iaubenfels
Ophl i tasporrg i a pennata Lambe
P1 ooamia karyki na de Laubenfels
Prosubsrites sisyrnus de Iaubenfels
Stelletta clarella de Iaubenfels
PHYLUM COELENTERPTA

References: Hyman, Invertebrates, I

Fraser, Eydmids of the Pacific Coast
hyer, Medusae of the World
The coelenterates resemble the sponges in that they are an ancient
group, relatively simple in structural organization, wholly aquatic in
habit, with their greatest development in the oceans, where they are to
be found from shore line to abyssal depths. At the seashore they are
confined, with few exceptions, to low tide level or below, as are the
sponges, Bryozoa and tunicates, because, like these groups, they are
not adapted to withstand intertidal exposure. In contrast to these
groups, which are all plankton feeders, all
coelenterates are ~redators.althouah manv
of them are stationary. Their contiiued &ccess seems to be explained by two devices
for food getting and defense, tentacles and
nematocysts, combined with effective means
of distribution, by ciliated larvae
and,
tulae or gastrulae known as planulaeIblaSin addition, in many g>oups (Scyphozoa,
Hydrozoa ) , a free-swimming sexual medusa
stage.
Representatives of three major types of
coelenterates occur regularly in the littoral fauna of California: (1) hydroids of
several species and their medusae (figs
7-9) and a species of hydrocoral (fig. 51,
( 2 ) a coral. and ( 3 ) several s~eciesof sea
anemones. 1n addition a scypho&dusan jellyfish, Aurellia, is occasionally washed in
~ i g . 6.-part
of a oolony
of
~ i , , h ~ = , and its asexual generation (scyphistoma)
a purple encrusting hydrooccurs at several points along the coast.
ooral. x2. (After Fisher,
Another
scyphomedusan occasionally encoun1938. )
tered is Pelagia. Velella (fig. 6), a purple
siphonophore (Class Hydrozoa), occasionally washes up on the beaches in
great numbers.

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

Fig. 6.-Diagramtic
representation of part of colony of
Velellg, a purple siphonophore. ( ~ f t e rBuchsbaum, 1938.)

As we have but one species of coral, Balanophyllia elegans, easily

recognized by its orange to red color, and of course by*its calcareous
skeleton (see demonstration specimens), no attempt will be made to
study the diagnostic characters of that group. In the case of the anemones also, the characters used by us for their identification are simple (see key on page 19). The hydroids, however, require more detailed
treatment.
Study or review Obelia. k k e certain of your ability to use the
rather numerous technical terms, defined below, which occur in the keys
and descriptions of specimens of the animals of this group, and which
unfortunately are used with different meanings by different authors.
Observe them on the mounts of Obelia, Gonothyraea, Syncoryne, and the
demonstrations of various other types.
Certain terms are widely used with regard to the parts of the
hydroid colony. Unfortunately, however, they by no means always have
the same meaning nor are they always consistently applied. We shall use
these terms in the sense in which they are used by C. McLean Fraser in
Hydroids of the Pacific Coast. In Pratt, for example, you will find a
different usage and again in other works. It will be necessary to know
these different usages. Figure 7 illustrates them as used in the key.
The hydroid colony consists essentially of a continuous cellular
tubular portion, the coenosarc (fig. 7). The coenosarc consists of a
layer of ectoderm (or epidermis), separated by a thin layer of noncellular meso~lea,from an inner layer of endoderm (~astrodermis)surrounding the continuous central cavity, the coelenteron or pastrovascular cavitx. Surrounding the coenosarc more or less completely is a
thin, chitinous, non-cellular layer, the perisarc (or ~eriderm)secreted
by the epidermis.

The colony arises by budding of new individuals from an original

zooid which developed from a zygote. It consists, therefore, of a number of individuals known in general as zooids. Actually no definite

Fig. 7.-Diagranrmatic representation of part of a colony and a

medusa of a thecate hydroid (Obelia), with complete metagenetic
cycle. One of the b l a s t o s t y l e s and one hydranth are shown longitudinally sectioned. (APter Buchsbaum, 1938.)

limits can be set to the individuals and much of the colony is made
up of connecting and rooting branches.
There is always a certain amount of polymorphism and the zooids
may be of several different types, named on the basis of the function
for which they are specialized, nutritive, generative, defensive, or
sensory zooids. The term hsdranth is used to designate the terminal
portion of a nutritive (vegetative) zooid but does not include any
associated perisarcal structures such as the hydrotheca. The hydranth

14

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

therefore is entirely coenosarcal, consisting of the bods, hypostome,

and tentacles. The term trophosome is used to include all the nongenerative zooids, typically nutritive zooids, of the colony together
with any perisarcal structures associated with them.
The term ponosome is used to include all the specialized generative zooids of the colony, with the perisarcal structures associated
with them, and is thus in contrast to the term trophosome which refers
to the rest of the colony. In thecate hydroids (fig. 7) the gonosome
would include those asexual generative zooids, the blastostyles, which
are specialized to produce the sexual zooids by budding, together with
the coverings of the blastostyles, known as 8;onawia (Fraser) or
gonothecae, and the sexual zooids to which the blastostyles give rise
by budding (fig. 7). Fraser uses the term gonosome in this sense, while
Pratt restricts it to the medusoid stage (sexual generative zooids).
The sexual zooids are termed gonophores by some (I prefer this usage)
or referred to in general as the medusoid stage in contrast to the hydroid stage. By others the term gonophore is confined to certain reduced sexual zooids which remain sessile. More commonly (e.g., Fraser)
the term is used synonymously with blastostyle including often the budding sexual zooids and protective coverings. Others would term this a
gonawium, a term which Fraser uses as synonymous with gonotheca. Such
an element of a colony (blastostyle with buds and protective covering,
if any) is often spoken of as a "fruiting body."
The sexual zooids, spoken of in general as the medusoid generation
in contrast to the hydroid generation, produce gametes from which by .
fertilization arise the zygotes which develop into the initial zooids
from which new colonies arise by budding. All the zooids of a given
colony are derived from a single zygote, and hence the sexual zooids of
a colony are all of the same sex and we speak of the colony as being
male or female (somewhat incorrectly of course).
Typical or highly developed sexual zooids are free-swimming
medusae. All intermediate conditions are found between medusae with a
long free life on the one hand and sessile, completely reduced; sexual
zooids which have lost all remnants of medusa structure on the other.
The latter have the appearance of being gonads of the asexual generative zooids (blast~st~les)
on which they arise. These extremely reduced
sexual zooids are spoken of as sporosacs, or, as noted above, as gonophores. We shall see them in Clava among athecate hydroids (fig. 8 , ~ )
and in thecate hydroids in Campanularia. Intermediate types, sessile
but still showing vestiges of medusa structure, are termed medusoids.
Such are seen in Gonothyraea (fig. 8,C) in contrast to the fully developed medusae of Obelia or the sporosacs of Campanularia or Clava. In
Gonothyraea the ripe sexual zooids (medusoids) are extruded from the
gonothecal aperture by continued growth of the blastostyle.
The term gonophore is used therefore, with the most diverse meanings and we must be certain to know with which of these meanings it is
used in each case. I prefer to use it for any sexual individual, whether
a free medusa, a sessile medusoid on a sporosac (thus used by H-).
Pratt defines it as any sessile reproductive individual (whether medusoid or sporosac, thus not including medusae under it; but in his keys
he seems to use it in the broad sense since on p. 116 we find, "The
gonophores are medusaen). We shall use it here as does Fraser for any

COEIi3NmTA

15

asexual generative zooid together with the sexual individuals budding

from it and its protective coverings, t h a t is, f o r what I would term a
gonangium,
bearing budding sexual zooids
and enclosed i n a
the whole a l s o spoken of a s a
f r u i t i n g body.

7
1-

Fig. 8.-fbrdroid types ( a f t e r Fraser. 1937, except D) A, s i n g l e hydranth of

l e u t o s t v l a shon)ng f i l i f o r m t e n t a c l e s , developing sporcsros, and ebeence bf perisarc.
B, hydranth of Ssmcome m i r a b i l i s showing s c a t t e r e d c a p i t a t e t e n t a c l e s and young
medusa buds. C , Gonothyraea c l a r k i i . ho gonophores (gommgia)~ l e f t , mature with
protruding medusoids; r i g h t , immature. D, Plumulrria. Portion of a branoh (hydrocladium) enlarged t o s h o r h y d r ~ n t h sand nenmtophores ( a f t e r blLPPn). $-G, & l a o ~ h e n i a
E, colony, 3/4 n a t u r a l s i z e i note B h s c a t t e r e d corbulae. F, a corbula
much enlarged. GI, portion of a hydrocladium, enlarged. H, S e r t u l h - .
I , same,
3/4 n a t u r a l s i z e .

.-

I'

I n c e r t a i n genera of thecate hydroids the f r u i t i n g body is made

even more complex by complicated protective s t r u c t u r e s derived from
other elements of the colony. Such is the corbula i n Aglaophenia ( f i g .
8,F). These would be included i n the gonosome but would be i n addition
t o the gonophores a s defined by Fraser.
This necessity of using terms whose meanings d i f f e r with the
author, while annoying f o r the moment, affords very excellent i n t e l l e c t u a l experience.
The t e n t a c l e s of the hydranth may form a single whorl a s in,Obelia( f i g . 7), o r two whorls a s i n rnbularia, o r may be m a t t e r e d over the
hypostome, the elevation which bears the mouth;, a s i n S p c o r y w (fig.
8,B). They may be f i l i f o r m (slender) a s i n Clava and Obelia o r c a p i t a t e

16

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

(knobbed) as in Syncoryne. In one large group of thecate hydroids, the

plumularian hydroids (Family Plumulariidae), there are present in addition to the typical hydranths numerous small individuals which function
entirely for offense and are known as nemato~hores(fig. 8,D). The small
branches
which bear the hydranths and nematophores are known as hydro- -.
cladia.
The hydrotheca may be stalked (Obelia, fig. 7) or sessile (Sertularia, fig. 8,H; Abietinaria, fig. 9; Plumularia, fig. 8,D; Aglaophenia,
fig. 8,F). They may be uniserial (Plumulariidae, fig. 8,D,F) or biserial
(Sertulariidae, figs. 8,H, 9). When biserial they may be opposite (Sertularia, fig. 8,H) or alternate (Abietinaria, fig. 9; ~ertularella).
Study mounts of various types of hydroids and, using the key below, identify severar of them. See also keys in Fraser.
Be able to recognize the following genera commonly encountered in
the local fauna; Szpcor.yne, Clava, Eudendrium, !l"ubularia,Obelia, Gonothyraea, Campanularia, Plumlaria, Aglaophenia, Sertularia, Pol orchis
(medusa only), Abietinaria, and Sertularella. Be able to make s etches
of each, bringing out diagnostic characters.

i:

......
........................

1. Fresh-water; solitary, or in small imperfect colonies

2
1. Marine; colonial
3
2. With tentacles; without distinct sexual stage but the polyps
bearing gonads which probably represent vestiges of such a
stage
Hydra
(For species see Hyman, Amer. Micros. Soc. Trans., 48, 49, 50)
2. Without tentacles; with well-developed medusae
Craspedacusta sowerby (= Microhydra ryderi)
3. Colony floating; in our common form blue in color. Order SIPHONOPHORA.
.'Ve]elJa
3. Hydroids; with attached, flexible colony
4
3. Hydrocorals; colony calcareous, encrusting; vividly purple with
j (fig. 5 )
scattered circular craters. Allopora poryphora (~ieher
4. Medusae relatively large, up to an inch in diameter; hydroid
stage undetermined
Polyorehis
4. Medusae very small or not formed as such; known by the hydroid
stage
5
6
5. Hydranths athecata. Suborder GYMNOBIAS!TEA OR ANTBOMEDUSAE..
5. Hydranths thecate. Suborder CALPPTOBLASTEA OR LEPTOUEDUSAE.
.14
7
6. Hydranth with a single basal whorl of filiform tentacles
.12
6. Hydranth with tentacles scattered
6. Hydranth with two whorls of filiform testacles; hydranths the
largest of any of our hydroids; medusae borne on long blastoTLIbularia
styles just above the basal whorl of tentacles
7. Stalks of hydranths without perisarc, unbranched, rising from a
Hydractinia
spiny calcareous crust
7. Stalks of hydranths with a perisarc (except Clava), usually
8
branched, not rising from a calcareous crust

...........................

...........
........................
............
.
.................
.............................
...
..
....
...............
....
...............

.........

8. Hydmnth with trumpet-shaped hypostome; perisarc stiff, brown,

closely and sharply annulated
Eudendrium
8. Hydranth with a conical hypostome; perisarc usually not sharply
annulated, or if so only on the smaller branches
9
9. Colony very small, stems not at all or very slightly branched
Perigonimus
9. Colony larger (up to several centimeters in length); stems
10
branched
Garveia
10. Hydranths relatively large, orange-colored, conspicuous
10. Hydranths smaller, not orange-colored
11
11. Hydranths very ;mall; colony finely branched; annulations of
perisarc almost entirely lacking
Bouqainvillia
11. Hydranths larger, colony with few branches; annulations distinct
at base of hydranths
Bimeria
12. Tentacles capitate (enlarged at tip); colony small, very
slightly branched; medusae arising among tentacles (fig
8,B)
8yncoryne
12. Tentacles filiform (tapering at tip); sexual zooids borne below
13
tentacles.
13. Hydranths, large, unbranched, arising from a filiform hydrorhiza; sexual zooids are sporosacs and colored, pink in male,
purple in female; no perisarc (fig. 8,A)
Clava
13. Hydranths small, colony very slightly branched; with free medusae; perisarc present
firri topsis
14. Hydrothecae stalked
15
14. Hydrothecae sessile (fig. 8,D-I)
18
Culycella
15. Hydrothecae tubular; colony extremely small
15. Hydrothecae bell-shaped; colony larger
16
16. Sexual zooids sporosacs; blastostyle not projecting
Campanularia
16. Sexual zooids medusiform but sessile; blastostyle and mature
medusoids projecting from gonotheca (fig. 8,C)
Gonothyraea
17. Medusae globular with 4 tentacles at time of liberation; colony
notmuchbranched.
Clytia
17. Medusae flatter with 16 or more tentacles; colony usually more
branched
Obel i a
18. Hydrothecae on one side of branches; nematophores present
19
(fig. 8,D-G)
18. Hydrothecae in two rows on o posite sides of branches; no
20
nematophores (figs. 8.H. 97
19. Colony slender and flexible, colorless; hydrothecae at considerable intervals, margins of hydrothecae not toothed or
Slumularia
sculptured (fig 8,D)
19. Colony large, stiff, brown or greenish-brown; branches of colony resemble feathers (fig. 8,E); hydrothecae close-set with
toothed and sculptured margins (fig. 8,F); "the ostrich
p ~ ~ m e ". . . . . . . . . . . . . . . . . . . . . . . A g ] a o p h e n i a

............
......
...........................
..........................
..
.............
.........
..................
.
.........................
.........................
.........
................
......................
................
.......
............
..
...
...................
........................
........................
................

................

18

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

20. Hydrothecae exactly opposite each other; operculum of two pieces;

colony small, delicate, without branches (fig. 8,H,I)

. . . . . . . . . . . . . . . . . . . . . . . . . . .Sertularia
.............

20. Hydrothecae alternate or subalternate

21
21. Hydrotheca with smooth margin; operculum of one piece; colony
larger, plume-like, with numerous side branches in one plane
(fig. 9)
Abietinaria

.....................

Fig. 9.--filioul.
Ellie and Solander. &lrly;ed
(from ~uberlet)

21. Hydrotheca with toothed margin; operculum of 3-4 pieces; colony

small, without side branches
sertularella

...........

LIST OF SPECIES OF HYDROIDS MOST LIKELY TO BE F O ~ D

INTERTIDALLY IN TBE SAN FRANCISCO REGION
I

(There are many other species which have been taken from shallow
water in this region, so that definite identification can be made only
by careful reference to Fraser, 1937. None of the species has been
marked abundant, because .of insufficient information. )
Athecates (Gymnoblastea or Anthomedusae )
B Bimeria franciscam Torrey
B Bougainvillia mertensi Agassiz
B Clava leptostyla Bgassiz
TB D B M adendrium californicum Torrey
D B M Garveia annulata Nutting
TB M Hydractinia milleri Torrey
B Perigonimus repens Wright
D B M Syncoryne mirabil is Agassiz
B M fibularia crocea Agassiz
M fibularia marina Torrey
B firritopsis nutricula McCrady
Thecates (Calyptoblastea or Leptomedusae)
D B M Abietinaria anguina Trask
D B M Abietinaria greenei lldurray

I
I

DBM
DBM
DM
B
BM
DM
B

BM
B
TB B M
B
B
M
B M

D BM
DBM
DBM

Aglaopenia strvthionides Murray

Calycella syringa Linnaeus
Campanularia jusiformis Clark
Campanularia gelatinosa Pallas
Campanularia urceolata Clark
Campanularia volubili sLinnaeus
Clytia edwardsi Nutting
Gonothyraea clarki krktanner-Turneret scher
Obelia conunissuralis McCrady
Obelia dichotoma Linnaeus
Obelia longissima Pallas
Obelia pl icata Hincks
Plumlaria goodei Nutting
Plumlaria lagenifera A l h
Plumlaria setacea E l l i s
Sertularella turgida Trask
Sertularia Jurcata Trash

OTHER EYDROZOA (See key)

Order Cal toblastea

Po yorchis penicillata Eschscholtz, common Pacific Coast medusa

Order Hydroida
F Hydra
Order Td7;;rallina

opora porphyra Fisher

Order Siphonophora
Velella lata Chamisso and Eysenhardt (often washed on shore)

Class SCYPHOZOA

Aurellia aurita Linnaeus

Pelagia
Class ANTEOZOA
KEX TO TEE SEA ANEMONES
(Modified from S
b! keys by A. S. Harrison and N. E. Kemp.
Nomenclature adapted t o that of Carlgren, a s reported on
p. 263 of Ricketts and Calvin. This group is very much i n
need of study i n the Pacific Area. )

......

1. Base attached t o rock, wood, o r other firm substmte

2
1. Base unattached-true burrowing anemones
Group Athenaria
(Genera and species unidentified )
2. Margin of tentacle-bearing disc deeply f r i l l e d o r lobed when
extended; tentacles short and very numerous; tentacle-free
area around mouth very small
Metridium dianthus

.....

.........

LABORATORY AND FIELD TMT I N INVERTEBRATE ZOOLOGY

20

2 . bkirgin of d i s c c i r c u l a r and not f r i l l e d when extended; t e n t a c l e s

3
fewer, leaving a l a r g e r a r e a f r e e around mouth

3.
3.

.......
Column thickly covered with p a p i l l a e ( s t i c k y t u b e r c l e s ~ a ' ~ b l e
of holding b i t s of s h e l l s and o t h e r d e b r i s t o t h e column . . 4
Column smooth o r , i f with p a p i l l a e , these arranged i n a few
v e r t i c a l rows and very small . . . . . . . . . . . . . . . . 8

4. With r e l a t i v e l y small p a p i l l a e , probably suckers, extending

over t h e upper two-thirds of t h e column i n longitudinal rows;
l i v i n g completely covered i n sand (but attached t o rock)
u n i d e n t i f i e d "sand anemone"
4. With p a p i l l a e p r a c t i c a l l y covering t h e e n t i r e column
5.
5. Column embedded i n mud: found along bay shore where mud and
"The Mud Anemone" ( u n i d e n t i f i e d )
rocks occur together
5. Column never embedded i n mud
6 .
6. Column dark red, o f t e n with brown patches; t e n t a c l e s blunt,
sometimes with t h e middle half of t h e t e n t a c l e rose-tinted
Tealia (= ~rticina)crassicornis .
6. Column green t o yellowish-white
7

.................

.....

It

.....
.................

................

................

7. P a p i l l a e i r r e g u l a r , branching, not conspicuously i n longitud i n a l rows; t e n t a c l e s uniform i n color, not tipped with pink;
solitary; large
Anthopleura (= Cribrina o r Bunodactis) xanthogrammica 7. P a p i l l a e round, arranged i n l o n g i t u d i n a l rows; t e n t a c l e s
u s u a l l y tipped with pink; c o l o r of column very variable;
a
usually i n close -se t aggregations
Bunodact is
Cri brina) elegant issima
8. L a t e r a l buds o r young anemones attached i n a c i r c l e t around t h e
column near t h e base
Epiaotis prolifera
9
8. No l a t e r a l buds o r young anemones attached t o t h e column

.....

.............
...........

(5

...

9. Found on underside of ledges on rocky ocean coast; column dark

Corynactis sp.
red; t e n t a c l e s white and c a p i t a t e
9.Foundmostlyinbaywaters
.10
10. Tentacles opposite each end of mouth-slit d i f f e r e n t l y marked
from others by having a yellow base i n c o n t r a s t t o t h e dark
reddish-brown base of t h e o t h e r t e n t a c l e s ; main portion of
the t e n t a c l e s yellowish o r banded with a l t e r n a t e yellowish
and brown bands; f a i n t r a d i a t i n g l i n e s from t h e mouth a l l
yellow; found on p i l i n g s a t F r u i t v a l e Bridge
"Yellow-striped Anemone" ( u n i d e n t i f i e d )
10. Tentacles opposite ends of mouth-slit not d i f f e r e n t l y colored
from o t h e r s
11
11. Column dark o l i v e green, o f t e n with v e r t i c a l orange s t r i p e s ;
small (extended column may reach 14 mm.); found a l l along
East Bay shore
"The Orange-striped Anemone, " Diadumene (= Sagart ia luciae)
11. Same a s D. luciae (may be a v a r i e t y ) except that every speclmen
has l o n g i t u d i n a l double rows of white l i n e s on t h e column;
found on p i l i n g s at F r u i t v a l e Bridge
"Double-striped Anemone" ( u n i d e n t i f i e d )

........
.................

.............
.........................

..

............

.................

11. Column never dark olive green

12
12. Column bright orange-red; inner base of each tkdacle with
white Bpot; oral opening surrounded by flaky white ring with
white lines radiating out to tentacles; on ocean coast
Sagartia (?) stimpsonii
12. Column and tentacles salmon to flesh color; column long and
slender when extended; surface of column with vertical lines;
on wood at Lake Merritt and Fruitvale Bridge
Sagartia leucolena

....................

.......................

CmCK LIST OF ANTHOZOA

Alcyonaria
Unidentified, rare
Sea Anemones
Anthopleura xanthogrammica Brandt
(= Cribrina or Bzlnodact i s ~antho~rammica)
Bunodact i s elegantissima Brandt
(= Cribrina elegantissima)
Corynacti s
Diadumene luciae Verrill
(= Sagartia luciae)
Epiacti s pro1 ifera Verrill
Metridium dianthus Ellis
Sagartia (? ) stimpsoni i Fewkes
Sagartia leucolena Verrill
Urticina (Tealia) crassicornis ~hrenberg'
"Mud Anemone" unidentified
"Yellow-striped Anemone" unidentified
"Double-striped Anemone" unidentified
"Sand Anemone" unidentified
Unidentified burrowing anemones
Corals .
Balanophyllia elegans Verrill
PHYLUM CTENOPHORA

The subspherical transparent jelly-like animals of this small

pelagic phylum which occasionally wash up on our shores (see figure
in text), belonging usually to the species Pleurobrachia bachei, are
variously known as cat ' s-eyes (by fishermen), sea gooseberries, or
sea walnuts. They are micropredators, capturing their prey by means
of the lasso cells of the long paired tentacles.

22

LABORATORY AND FIELD TEXT IN EUMENTARY ZOOLOGY

A ctenophore

PHYLUM PLATYHELMINTHES

The phylum Platyhelminthes is the lowest of the truly bilaterally

symmetrical phyla (the Bilateria). Two of the three classes, the Trematoda and the Cestoda, are parasitic and are not studied in this course.
The third olass, the Turbellaria, is composed of slow-moving but freeliving forms, all of which are predators.
The class Turbellaria contains four groups: (1) the Acoela, minute
marine forms without an intestine, not ordinarily encountered in the
local fauna; (2) the rhabdocoeles (Rhabdocoela), small, typically narrow forms in which the intestine is a single straight tube; (3) the
triclads (Tricladida), in which the intestine has three main trunks,
one forward from the pharynx and two backward (in marine species, mouth
in hinder part of body, intestine little branched); (4)the polyclads
(Polycladida), a large group of marine species of considerable size with
thin, leaf-like body and with numerous branches of the intestine.
The Turbellaria of the Pacific Coast are very poorly known. The
only important paper is that of Heath and YcGregor (1912). New Polxclads from Monterea Bay. California. Previous,tothat time but three
species of polyclads had been reported from the 7000 miles between
Panama and Asia. They report 18 species, all new, belonging to 9 genera, three of them new.

The common species in Central California seems to be Leptoplana

acticola (Ricketts and Calvin, p. 26) but the larger Planocera (Ricketts
and Calvin, p. 16) also occurs. Any who desire to carry further their
studies of the species of this very difficult group should consult
Zang, Die Polscladien des Golfes von Nea~el,and various works referred
to in Pratt.
The well-known triclad flatworms of the genus Planaria (properly
h g e s i a , according to Hyman) are frequently found in fresh water in
this region (especially in Golden Gate Park). Terrestrial flatworms of
two genera ( Bipal ium kewense Moseley and G'eopl ana mexi cana Hyman ) , also
of the Tricladida, are found on the Berkeley campus where they may most
easily be found feeding on earthworms killed during the spring rains.
Ricketts and Calvin mention three genera of rhabdocoeles, p. 265.
Of these, Syndesmis is very abundant in the intestine of the sea urchins.
The other two have not been identified from the areas here considered.
PHYLUM FEMEXl"l'A (NEMERTINEA)

References: Coe, 1. R., 1940, Revision of the Nemertean

Fauna of the Pacific Coast
Coe, W. R., 1905, Nemerteans of the Yest and
Northwest Coasts of America
Coe, W. R., 1904, Nemertinea, Harriman Alaska
Ex~edi
tion
Nemertean worms are readily recognizable by their relatively elongated and narrow, highly contractile, more or less flattened, unsegmented bodies as also, of course, by their long, slender, eversible
proboscis. This proboscis may be caused to evert by adding alcohol, a
few drops at a time, to a small amount of sea water containing the
worms or by putting them into fresh water. Like the polyclads they are
predators with a wide range of prey.
They are handled with difficulty because of their extreme contractility and the slimy mucus which they secrete in large quantities,
and should, therefore, be kept separated from other animals. For scientific purposes the> should be preserved in formalin.
Using the key below, identify your specimen. Check your identification with @oels description. Identify as many species as time permi.ts.

m TO m Mom

COMMON N E M E R U S

(For other species see plates of Coe: 1940, 1901, 1904, 1905.
Figure numbers refer to Coe, 1940. )

. .2
. . . . . . . . . . . . . . . . . . . .8
.................3
. . . . . . . . . . . . . . . . . . . .5

1. Entire body or dorsal surface solid color, no lines or rings

1. With longitudinal lines or transverse rings of color contrasting with ground color
2. Of same shade above and below
2. Dark above, light below

24

LABORATORY AND FIELD TEXT I N INVERTEBRATE ZOOLOGY

3. Minute, very slender; whitish, somewhat transparent; l i v i n g i n

d e l i c a t e parchment tubes under stones o r among algae
fibulanus p e l l u c i d u s (= Carinella pellucida)
3. Large f r e e - l i v i n g species with d i s t i n c t i v e colorations (red,
4
dark red, o r brown)
4. Bright red; very soft-bodied; l i v i n g i n o r on rocks
!Pubulanus polymorphus (= Car i n e l l a rubra)
4. Dark red t o brown; f i r m bodied; l i v i n g i n mud o r sand
Cerebratulus c a l i f o r n i e n s i s (Coe, 7 )
7
5. Color of head not noticeably d i f f e r e n t from r e s t of body
5. Head darker (Lineus) o r l i g h t e r (Amphiporus)
6
6. Broad f l e s h y species; head l i g h t , with two l a r g e , conspicuous,
dark brown t o black spots; oval t o t r i a n g u l a r i n shape (see
f i g u r e i n t e x t ) ; d o r s a l body c o l o r deep brownish-orange
Amphiporus bimaculatus

..........

.....................
...........
..............
...
.........
.....................

Two Comon Nemerteans ( a f t e r ~ o e ) .A , Micrura v e r r i l l i Coe. I t s distinctive white

caudal cirrus or t a i l has been l o s t as i s cPten the case. B ,
m u l e t u s Coe.

6. Very slender species; head darker than r e s t of body; body l i g h t

o l i v e green t o brown; a l i n e of 2-8 o c e l l i on e i t h e r s i d e of
a n t e r o l a t e r a l region of head; o f t e n with f i n e c i r c u l a r and
l o n g i t u d i n a l l i n e s (see elsewhere i n key)
Lineus vegetus (Coe, 1 3 )

....................
7. Very slender; uniform green above, w h i t i s h below
. . . . . . . . . . . . . . . . . . . . . . En lectonema g r a c i l e
7. Stouter; uniform brown, orange brown, o r purple grown on d o r s a l .
surface and s i d e s . . . . . . . . . . . . Paranenrertes peregrina
8. Very slender . . . . . . . . . . . . . . . . . . . . . . . . . 9
8.Stouterandlarger . . . . . . . . . . . . . . . . . . . . . . 1 0
9. Egtremely slender, a s much as one hundred times a s long a s wide;
a s i n g l e row of 2-8 o c e l l i on each a n t e r o l a t e r a l margin of
head; u s u a l l y with f i n e c i r c u l a r and l o n g i t u d i n a l l i n e s ( s e e
Lineus vegetus (Coe, 1 3 )
elsewhere i n key f o r those without)
9. Slender but l e s s so than above; d o r s a l surface with two narrow
longitudinal dark l i n e s on l i g h t background; two l a r g e o c e l l i
oneachside
.Nemertopsisgracilis
10. General body c o l o r deep brown; f i v e o r s i x very slender, o f t e n
broken, l o n g i t u d i n a l white l i n e s ; numerous narrow white r i n g s
spaced a t f a i r l y r e g u l a r i n t e r v a l s
!lbbulanus sex1ineatus (= C a r i n e l l a sex1ineata )

..

..............

.........

10. General body color ivory white, with a series of rectangular

dorsal areas of deep purple or wine color, separated by narrow
white bands (see figure in text ); posterior end with narrow
colorlees "tail," caudal cirrus (not shown in figure)

. . . . . . . . . . . . . . . . . . . . . . . . Micru:a verrilli

D B *M
TI3 D M

D- *M

M
TB D M
*B *I
*M
M

LIST OF COMMON NEMERTEA

Amphiporus bimaculatus Coe
ZZIbulanus polporphus Renier (= Carinella rubra Griffin )
Tubulanus sezl i neatus Griffin (= Cari nella sex1 ineatus
Griffin)
!l'ubulanus pellucidus Coe (= -Carinella pellucida ~ o )e
Cerebratulus californiensis Coe
Euplectonema gracile Johnston
Micrura verrilli Coe
llTemertopsis gracilis Coe
PHYLUM ROTIFERA

These minute animals chiefly plankton feeders are almobt always

present in freshwater cultures. Isolate a rotifer on a slide and anesthetize it for study or find one immobilized on a mount from a rich
culture. Study it under low and high power, making out the various
structures referring to figure 10. Figure 11 shows another type of
rotifer (Brachionus) very commonly encountered. Attempt to identify
your specimen, using the key below.

Fig. 11.-with two resting or winter egg8

(after Whitney)

Fig. 10.-Diagrammatic hemisection of a typical

rotifer (af'tar Shipley and MacBride).

26

LABOWTORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

Make yourself familiar with the chapter on rotifers in Ward and

Whipple and with the monograph of Hudson and Gosse (2 vols., 1889).

A KXY TO TflE ROTIFERA OF LONE TREE POND

1.
1.
2.
2.
3.
3.
3.

By A. B. Burch
Colonial
Megalotrocha alboflavicans
Solitary
2
Lorica pronounced
3
12
Lorica poorly developed or absent
Lorica laterally compressed
4
Lorica dorsoventrally flattened
5
Lorica more or less radially symmetrical, toothed anteriorly
with or without one long spine posteriorly; no foot

.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . . . . . . . . . . . .
.......................
...............
...................
................

. . . . . . . . . . . . . . . . . . . . . . . Anuraea cochlearis
4. Lorica with three dorsal furrows, toothed before and behind;
foot with two toes . . . . . . . . . . . . . Salpina spinnigera
4. Lorica with hooked head-shield, not toothed . . . . . . . Colurus
5. Lorica with distinct longitudinal furrows . . . . . . . . . . . 6
......... 8
5. Lorica without distinct longitudinal furrows
6. Lorica with one median dorsal furrow, teeth absent, one eye

. . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplois
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplax
8. hrica toothed anteriorly and posteriorly . . . . . . . . . . . 9
8. Not so toothed . . . . . . . . . . . . . . . . . . . . . . . . 11
9. hrica very convex dorsally . . . . . . . . . . Brachi onus bakeri
9. Lorica very flat . . . . . . . . . . . . . . . . . . . . . . . 10
10. With foot and two toes . . . . . . . . . . . Noteus quadricornus
.................
6. Lorica with one median dorsal furrow, teeth absent, eye absent

10. Without foot

.Anuraea aculeata
11. Lorica toothed anteriorly, with a median notch posteriorly;
foot short, two medium-sized toes, each with an oblique
Euchlanis mucrura
bristle at the base
11. Lorica toothed anteriorly, no notch posteriorly; foot long, with
one toe
.Brachionuspnotatus
Trochosphaera
12. Foot absent; body round
12. Foot present
13
14
13. Body highly contractile, telescopic
13. Not so
15
14. With three fine toes on very long, contractile foot, two spurs
Rotifer neptunius
midway of length of foot
14. With two toes
fiilodina roseola
15. Body more or less triangular in general outline; two anterolateral projecting styles
Synchaeta stylata
15. Not so
16

.............
.................
. . . . . .. .. .. .. .. .. .. .. .. .. .. .. .. .. . . . . . .
..............
............................

..........
.................
. . . . . . . . . . .. .. .. .. .. .. .. .. .. .. . . . . . . . .

16. Body very flat, quite round with a projecting corona; with a
small cylindrical foot, no toes; vase-shaped. Pterodina patina
16. Body not flat, but slightly flattened dorsally when swimming;
somewhat elongate; divided into a head and body region; foot
short,with two toes
.
. Triphylus lacustris

...... ....

(Some may, however, be found in M e Merritt when that water has

little salt. )
Names as in key and in Ward and
Names as in Harring, H. K. ,. 1913.
Whipple:
"Synopsis of the Rotatoria,"
hll. U.S. Nat. Mus., 81
Anuraea aculeata
Keratella quadrata Miiller
Anuraea cochlearis Gosse
Keratella cochlearis
Colurus
Colurella
Brachi onus bakeri
B. capsuliflorus Pallas
Brachionus punctatus Hempel
B. budapest i nens i s Daday
Diplax
Mytilina sp.
Diplois
Diplois
fichlanis maorura Ehrenberg
Ehchlanis macrura Ehrenberg
Megalotrocha alboflavicans
Sinantherina socialis Linnaeus
Ehrenberg
Platyias quadricornis
Philodina roseola Ehrenberg
Noteus quadricornis Ehrenberg
Philodina roseola Ehrenberg
Testudinella patina
Myti1ina wcronata spinigera
Pterodina patina Hermann
Rot ifer neptunius Ehrenberg
Synchaeta stylata Wierzejski
Synchaeta stylata Wierzejski
Salpina spinigera Ehrenberg
Synchaeta stylata Wierze jski
Enteroplea lacustris Ehrenberg
Triphylus lacrustris Ehrenberg
GASTROTRICHA

These microscopic creatures, often grouped with the rotifers but

structurally very distinct (fig 12), are commonly encountered in
ponds and in freshwater cultures and should be recognized.

F i g . 12.-A
p e t r o t r i c h , Chaetonotue maximum Ehrenberg (fram Weasenberg-Lund a f t e r Zelinka).

28

IDORATORY AND FIELD TEXT IN IllvER~BRBTEZOOLOGY

PHYLUM ANNELIDA

The annelids or segmented worn can be divided into the following

classes: (1)Archiannelida, a small, poorly known group with distinct
segmentation in our representatives, parapodia often poorly developed,
often lacking setae and with the two ends of the body often developed
for rapid locomotion through sand. When more completely laown they may
be found to represent modified forms of widely separated, already
known families. (2) Polychagta (fig. 13,A), which have many setae
(hence the name) and arapodia; they are almst entirely marine &d
are very numerous. (37 Oligochaeta (fig. 13,B.C). with a few setae and
no parapodia; they are predominantly freshwater or terrestrial. These
last two classes are often combined to form the single class Chaetopoda, but there is evidence that the two are not so closely related to
each other as each is to other groups in the phylum. (4)Hirudinea or
leeches (fig. 13,D), with no setae or parapodia and with important internal anatomical differences from the other classes; they are mostly
inhabitants of fresh water. (5) Eohiuroidsa, consisting of a few marine
species, of which Urschis caupo (fig. 13,F) is our local representative
(burrowing in mud). They have a few annelid characters, but are umegmented externally. (6) The Sipunculoidea (fig. 13,E), of very doubtful
affinities with the annelids. Like the echiurids, they are unsegmented
and they are likewise marine. The last two classea are probably more
properly considered as separate phyla.
Class POLYCHIlETB,
The names and characters of the major divisions of the polychaetes
are very much in dispute. One useful but perhaps not scientifically
justifiable division is into the Sedentaria or sedentary polychaetea,
chiefly tube dwellers, and the Errantia or free-living families. More
useful is the designation of family types (fig. 17), such ae nereid,
polynoid, serpulid, syllid, sabellid, cirratulid, spionid, eta.
On your specimen (Nsreis) make out the paired locomotor parapodia
(sing. parapodium) each (fig. 14) composed of an upper lobe, the notppodium, and a lower, neuro~odium.Remove and mount a parapodium to make
out the structures noted below. Each lobe t ically contains a bundle of
slender chitinous rods or spines, the setaeysing. seta) which project
from the parapodium, and buried in it a larger dark spine known as the
aciculum (pl. acicula) which functions as a fulcrum for morirq the parapodia and the setae. !Che shape, size, number, and position of the setae
are of importance in olassification. Arising from the b e of the notopodium above and the neuropodium below there are often present elender
flexible outgrowths (fig. 14), the dorsal cirrus (pl. cirri), and1cirrus respectively. The notopodium consists of a dorsal lobe and a
middle lobe between which the setae and aciculum project (fig. 14); the
neuropodium consists of a neuro-acicular lobe provided with setae and
an aciculum, and a ventral lobe.
The setae are of widely varying form and furnish very precise
characters for the determination of species. The use of such characters has been avoided where possible in the key but has been found
neceseary in some cases. !be common types and their names should be

I
I

1
i

D. A leech ( ~ i r u d i n e a )

A. Nereis
( Polyohaeta)

B. bolosama, a freshwnter oligoohaete

m.

C. An earthworn

8. Sipunculoidea

F. Urechis (~ohiuroidea)

(~li~oohaeta)

Fig. 13.-Typos of
(C, after Altuun; D , a f t e r Bwhsba~m;E, after
Fisher; F, a f t e r Bioketts and Calvin a f t e r k o O i n i t i e . )

30

LABORATORY AND FIELD TEXT IN lXVEFtTEBRBTE ZOOLOGY

recognized. Plate I (after Fauvel, 1922) gives some idea of the wide
variety of types. First we may distinguish simple setae (Plate'I, nos.
1-19) from composite or jointed setae (nos. 21-28). Long slender simple setae are spoken of as capillary setae (no. 6). The tips of setae
whether simple or composite may be entire or bifid (no. 8) or trifid.
If beni like a cycle they are termed falcate.
Some simple setae have bent, usually bifid ends. These are spoken
of as hooks or crotchets (Plate I, nos. 29-32). These are usually
relatively stout setae and graduate into broadened types known as
uncini (nos. 33-42), which are especially characteristic of the Sedentaria.

I
I
I.

II

notopodium

neuropodium

Fig. 14.-A parapodium of a nereid

polyohaete.

1
I

I1

Fig. 15.-Head of Nereis with everted proboscis: A, dors a l view; B , ventral view. ( 1 ) prostomium~ ( 2 ) palpi; ( 3 )
prostornial tentacles; (4) eyes; ( 5 ) perietomiumr ( 6 ) peristomial or tentacular c i r r i ; ( 7 ) jaws. I-VIII, areas of proboscis; V-VIII, on oral ring; I-IV, on maxillary ring.

Composite setae may be articulated as in the long bristles of the

Flabelligeridae (Plate I, no. 20), but are characteristically twojointed, composed of a shaft and a blade (nos. 22-28). This blade in
turn may have various shapes and may itself be a crotchet (no. 28).
The blade rests in a notch in the end of the shaft. If the two sides
of the notch are equal, it is spoken of as homogomph (no. 27); if
somewhat unequal, as hemigomph; and if distinctly so, as heterogomph
(no. 26). Finally setae, either simple or compound, may be embedded
distally within a clear matrix and these are spoken of as "hooded."
Thus we may have simple hooded crotchets (no. 29) or composite crotchets (no. 28).
Identify the following structures of the head; a pre-oral prostomium (fig. 15,A), provided at its anterior border with a pair of antennae
, and at the sides with paired, fleshy, biarticulated palui. Two
pairs of -,
more or less in rectangular arrangement are to be found
on the dorsal side of the prostomiurn, the more posterior pair sometimes
concealed by the peristomial ring. The pristomium is the first complete

Types of polyahaete setae (after ~auvel)

32

LAEORATORY AND FIELD TMT I N INVERTEBRQTE ZOOLOGY

r i n g o r segment. It may, however, represent two coalesced segments a s

i n Nereis. There a r e four p a i r s of slender, tentacular c i r r i (peristomial c i r r i ) on short s t a l k s a t the a n t e r o - l a t e r a l margin of the peristomium. Note t h a t the f i r s t parapodia a r e found on segment two (considering the peristomium a s one), and t h a t the f i r s t two p a i r s of
parapodia a r e uniramous, t h a t is, each contaias a s i n g l e aciculum
and s e t a l f a s c i c l e , and that a l l succeeding parapodia a r e birrnous.
W e a ventral longitudinal incision s t a r t i n g from the mouth and
extend the cut posteriorly through about twenty segments, Pin back t h e
f l a p s and identify the p a r t s of the proboscis. The following parts
( f i g . 15) a r e t o be found. (In the r e t r a c t e d proboscis the position of
o r a l and maxillary rings is i n reverse order): (1) an o r a l r i q which
is just external t o the mouth ( i n t e r n a l when proboscis is everted) and
bounded by the mouth on one end and the maxillary r i n g on the other,
d i v i s i b l e i n t o s i x areas, numbered from V t o VIII; and (2) beyond t h i s
a second ring, the maxillabounded on one end by the o r a l ring,
provided with stout, horny jaws at its other end and d i v i s i b l e i n t o s i x
areas numbered from I t o IV. Area V l i e s on the median dorsal l i n e of
the o r a l ring and i n Nereis is usually without the dark chitinous dent i c l e s termed paragnaths; areas V I a r e paired, lying on e i t h e r s i d e of
V, and normally a r e provided with a c i r c u l a r mount of t i n y pararmatha.
Area VII l i e s on the mid-ventral l i n e of the o r a l ring and areas VIII
a r e paired patches a t t h e sides of V I I . I n Nereis areas VII and VIII
usually have a continuous band of chitinous paragnatha. The areas of
the maxillary r i n g occupy positions similar t o those of the o r a l ring.
Area I l i e s i n l i n e with V, I1 with VI, I11 with VII, and I V with VIII.
Even numbered areas a r e always paired, odd numbered areas l i e i n the
median dorsal and v e n t r a l l i n e s .
Identify your specimen t o the group, using the key on page 34.
Study l i v i n g individuals of the serpulid Mercierella under the
dissecting microscope. Remove from tubes and identify: ( 1 ) a feathery
tentacular growp ( f i g . 16),
consisting of modified peristomial c i r r i and often called
4
branchiae, one branoh of which
i s specially modified t o form
1
F
the o~erc( f i g . 16). Ihe
m
G
prostomiurn i s g r e a t l y reduced
0
and made out with d i f f i c u l t y .
thomcio membrane
The & marks the ventral
side, and l i e s on t h e s i d e
opposite the operculum. h i s ing from the posterior border
of the peristomium is a broad,
t h i n membranous
open
dorsally, and r i c h l y supplied
with blood vessels and nerves.
Fig. 16.-Meroierella enignmtioa Fauvel,
Its function
is thought to be
the Lake Merritt tubtmora, removed Prom i t s
respiratorl.
~h~ collar
is
tube; sommhat enlarged.
coninued posteriorly a s the
U a c i c membrane along the posterior border of the thorax thus sharply
s e t t i n g off the thorax from a posterior region of the b o b , called the
abdomen.

a,

u,

POLYCHAETA

33

See and be able to recognize at this time the types given on the
list below.
Important Polychaete Tgpes (Families)
Nereid (figs. 13.8, 15)
Lumbrinerid (fig. 17,A and Plate V, no. 1)
~l~cerid
(fig. 17,~)
Phyllodocid (fig. 17,E and Plate PII, no
Polynoid (fig. 17,B)
Syllid (17,D and Plate BII, nos. 2 b 3)
Sabellid (fig 17,C and Plate I. no. 3)
Serpulid (figs. 16, 17,I)
Spionid (fig. 17,J and Plate IX, no. 2 j
Cirratulid (fig. 17,H and Plate IV, no.
Terebellid (fig. 17,G and Plate 111, no.

Syllid

1-

'

Terebellid

Cirratulid

Fig. 17.-Tries of polychaetesi A , Arabella, a lumbrinerid; 8 , 8 . l o s v d n . a ~ 0 1 ~ noid; C , M~%ioola,a sabellid; D , S y l l i s , a s y l l i d t E, Eulalia, a phyllodmid; F *
Glroem, a glyoerid; G , Terebella, a terebellid; H, C i m t u l u s , a o i m t u l i d t I,
Serpula, a serpulid; J , Bmoardia, a spionid.

34

LBBORATORY AND FIELD

TMT IN INPERTEBIIBTE ZOOLOGY

1. Dorsal surface more o r l e s s completely covered by overlapping

scales ( e l y t r a ) (fig. 17,B; Plate 11, no. 2), paleae (modified
flattened-setae, f i g . 188), o r overlain by a thick f e l t y
layer (Plate 11, no. 1 ) (APHRODITIDAE, POLYNOIDAE, POLYODONTIDAE, SIGALIONIDAE, CflRPSOPETALII3AE)
2
14
1. Dorsal surface not covered with elytra, paleae, o r f e l t

.............

....

2. Dorsal surface more o r l e s s concealed

covers the elytra. APHRODITIaaE
2. Dorsal surface more o r l e s s concealed
P l a t e I , no. 2 ) .
2. Dorsal surface more o r l e s s concealed
CBRPSOPETALIDAE.........

a f e l t y layer t h a t
. . byby. Aphrodita
(Plate 11, no.
e l y t r a (fig. 17,B;
......................
by paleae ( f i g . 18,B)
..............

1)

4
3

Fig. 18.-A, head of Sthenelais &ua; B , parapodium

of a ohrysopetalid; C , elytrum of Arctono6 f r a a i l i s ; D,
neuropodial s e t a of same; E , neuropodial s e t a of &&brevisetosa.

3. Paleae of one kind; color i n l i f e , pale rust t o yellowish

Chrysopetalum
3 . Paleae of two kinds; color i n l i f e , glistening white o r
greenish
Pal eonotus

..........................
......................

4. Elytra and dorsal c i r r i a l t e r n a t e regularly from f i f t h segment

t o posterior end of body. POLYODONTIDAE. Neuropodia with composite setae; prostomium without median antenna; body short,
subrectangular, t o 12 mm. long
Peisidice
4. Elytra and dorsal c i r r i a l t e r n a t e i n anterior segments, but i n
the posterior segments e l y t r a a r e present on a l l segments and
e n t i r e l y replace the c i r r i . SIGALIONIDAE. Median prostomial
antenna ( f i g . 18,A) with broad base and l a t e r a l wings; elonSthenelais fusca
gate; dusky brown; among roots of e e l grass
4. Elytra and dorsal c i r r i a l t e r n a t e regularly t o about the 23d
segment, thereafter there a r e two e l y t r a followed by a dorsal
5
cirrus. POLYNOIDAE (Plate 11, no. 2. )

............

..

............

Polyohaete types (af'ter ~ c ~ n t o s h ) :1, A hrodita aculeata inn.) (~~hroditidae);

2, Barmothoe imbrioata (Llnn.) ( P ~ l ~ n o i d ~ ~ ~ vesioulatwn
a l o o a onta tap)

s(a1-.

36

5.
5.
5.
5.
6.

LABORATORY BND FIELD TEXT I N INVERTEBRATE ZOOLOGY

.. .. .. .. .. .. .. .. .. .. .. . . . . . . . . .
...............
...............

With 12 p a i r s of e l y t r a
Lepidonotus caelorus
With 15 pairs of e l y t r a
10
With 18 pairs of e l y t r a
Halosydna 6
7
I i t h more than 18 p a i r s of e l y t r a
Zlytra more o r l e s s firmly attached; neuropodial setae e n t i r e
a t end (fig. 18,E); common i n mussel beds, o r commensal with
Thelepus
Halosydna brevisetosa
6. Elytra e a s i l y shed; neuropodial setae b i f i d at end; free-living, under stones
Halosydna johnsoni
7. Elytra more o r l e s s completely covering the d o r m i n the anter i o r region; dorsal c i r r i and antennae club or pear-shaped;
neuropodial setae strongly f a l c a t e (sickle- shaped, f i g
Arctono8 8
18,D), the sides smooth
7. Elytni reduced i n s i z e throughout, those i n the posterior half
not noticeably smaller than those i n the a n t e r i o r half;
dorsal c i r r i and antennae cirriform; neuropodial setae s e r 9
r a t e d . . . . . . . . . . . . . . . . . . . . . . . . . . . .
8. Elytra strongly f r i l l e d at external 'margins (fig. 18,C); i n
Arctonolf f r a g i l i s
ambulacral grooves of Pisaster
8. Elytral margin smooth externally; a broad, transverse dark
band across anterior segments; i n branchial grooves of mollusks
A. v i t t a t a
8. Elytra s l i g h t l y undulate a t margins; commensal with holothurians
A. p l o h r a
9. Elytra i n pairs throughout length of body; notopodia without
Lepidasthsnia
setae; comme~maloften with tube-building worms
9. Some e l y t r a i n posterior region not paired; some a n t e r i o r notopodia have setae; commensal
Lepidamt r i a
10. Notopodial and neuropodial setae each of one kind
12
10. Notopodial and neuropodial setae each of two kinds, a stout
Hesperonob' l.l
and a slender form
11. Elytral surface smooth or with very few minute, low papillae; commensal with Urechis
Hesperonor3'adventor
11. Elytra with numerous scattered low papillae; commensal with
Callianassa.
H. ooomplanata
13
12. Neuropodial setae e n t i r e a t d i s t a l end
12. Some neuropodial setae with t i p s b i f i d
Harmothod (Plate 11, no. 2 )
13. Neuropodial setae l o w , slender, the d i s t a l ends hairlike.
Antinoif
13. Neuropodial setae with d i s t a l ends thicker, not hairlike.
fino8
14. Bnterior end completely concealed by feathery tentacles.
SBBELLIDAE (fig. 17,C; P l a t e I1,no. 3), SERPULIDgE (fig. 16). 15
14. Anterior end completely concealed by chitinous spines forming
31
an operculwn. SBBELI$RIIDAE ( f i g . 19,A), PECTINARIIDAE
14. Anterior end with long, strong, spinous setae, not forming an
operculum, but projecting forward and concealing the prosto33
mium (fig. 18,B and Plate 111, no. 1). naaELLIGWIDBE
[See opposite page. ]

................
..............

. . . . . . . . . . . . . . ..

.......

........................
......................

............
.......
...............
........
..................
............
...................
............................

...
...

14. Anterior end more or less completely covered by many long,

filamentous outgrowths concentrated on anterior region (fig
17,G); body usually with two distinct regions. BMPHBRETIDAE,
(Plate 111, no. 2) TERFMUIDBE tfig. 17,G; Plate 111, no.3 )
34
14. Anterior end more or less completely covered by many long,
filamentous outgrowtha as in category above, but also with
similar outgrowths continued along sides of body (fig. 17,H)
(except Dodecaceria); body not divided into two distinct re40
gions. CIRRARTLIRAE (Plate IT, no. 1)
44
14. Anterior end othemise; prostomium generally not concealed

. . . . . . . . . . .. ..

Fig. 19.-A,

PhI'aKmato~oma, Family Sabellariidae; B , a f l a b d l i g e r i d .

15. Thoracic membrane well developed (figs. 16, 17,I); operculum

16
usually present; in calcareous tubes.
15. Thoracic membrane little developed or absent; no operculum; in
sandy chitinized tubes. SAEELLIDAE (fig. 17,C and Plate 11,
n0.3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2
Serpuli due (see 14 and 15)*
16. Minute; in small neatly coiled tubes.
Spirorbis
17
16. Small to larger; tubes straight or twisted, but not coiled
17. Without operculum; tentacles few; forming small fragile tube
masses; red in life
Salnaoina
18
17. Vith operculum; tentacles few to numerous
18. Operculum bladder-like with m o t h stalk; tubes usually solitary, fragile, aometimes longitudinally ribbed
Vemiliopsis
18. Operculum funnel-shaped
19
19. Stalk of operculum smooth; tubes solitary, fairly large, cylindrical to somewhat twisted
Serpula
19. Stalk of operculum with lateral wings; tubes strongly ridged
longitudinally
Spirabranchus
20
19. Operculum with spines but without lateral wings on stalk
20. Spines of operculum produced from a single terminal plaque
(fig. 16); estuarine (Iake Merritt)
Mercierella enigmatica
21
20. Spines produced in a two-storied structure; marine
21. Distal series of spines with lateral projections
Hydroides
21. Distal series of spines with smooth sides
fipomatus

......

.........

..

. . . . . . . . . . .. .. .. .. .. .. .. . . . .

. . . . . . . . . . . . . .. .. . . . . .
..............
.................
...

...

......
. . . . .. .. ..

Figures thue given refer t o the categories i n the key through which
the particular family ha8 been arrived a t and permit of a quick review
of the charaatere etreeeed.

38

LABORATORY AND FIELD TEXT

IN INVERTEBRLITE ZOOLOGY

Sabellidae (see 14 and 15)

22. Without a thoracic c o l l a r ; tentacles without eye-spots; large,
t o 10 inches long o r more; white o r pale yellow i n l i f e ; i n
Mgxicola ( f i g . 17,C)
sand o r muddy sand.
22. With a thoracic c o l l a r ( f i g . ZO,A,B,C) ( r a r e l y absent ); with
23
( f i g 20,E) o r without tentacular eye-spot s

............

.........

Fig. 20.-Sabellidae: A , Pseudopotamilla

o c c e l l a t a . ( 1 ) Tentacular membrane with a dorsal
l o b e or f l a p . B, P. intermedia. ( 1 ) Dorsal edge
of tentacular membrane straight; ( 2 ) c o l l a r with
two dorsal lobes; and ( 3 ) two l a t e r a l lobes. C ,
Sabella. D , long-stenrmed uncinus from thoraoic
neuropodium of p b r i c i a . E, tentacular radiole
of Sabella o r a s s i c o m i s showing paired eye-spots.

. . . . . .. .. .. .. Schizobranchia
. . . . . . 24
24. Composite eyes a t d i s t a l end of some radioles . . . . . Megalomma
24. Without composite eyes i n such position . . . . . . . . . . . . 25
25. With tentacular eye-spots . . . . . . . . . . . . . . . . . . . 27
25. Vithout tentacular eye-spots . . . . . . . . . . . . . . . . . 26
23. Tentacular filaments divided several times
23. Tentacular filaments not divided

26. Tentacular filaments united f o r most of t h e i r length by a palmate membrane ( f i g . 17,C); thoracic c o l l a r conspicuous forming a smooth, e n t i r e membrane, open i n mid-line; thoracic
Chone
crotchets with a long stem ( f i g . 20,D)
26. Tentacular filaments not united by membrane o r a t most f o r a
r e l a t i v e l y small part of t h e i r length; thoracic c o l l a r not
strongly developed; thoracic uncini with short stem
Sabellamedia

..........

.........................

. . . . ...

27. Tentacular eye-spots disposed in pairs (fig 20,E); tentacular

bases not arranged in a spiral
Sabella crassicornis
27. Tentacular eye-spots not in pairs; tentacular bases in a spiral (Eudistylia) or not (~seudo~otamilla)
28
28. Base of tentacular megbmne conspicuously spiraled (&d i sty1i a)
29
28. Base of tentacular membrane not spiraled (~seudopotamilla)
30
29. Dorsal edge of tentacular membrane deeply cleft, the two lobes
thus formed overlapping; tentacles deep maroon except for a
pale band distally; in rocky crevices especially on reefs.
.&distyliapolymorpha
29. Dorsal edge of tentacular membrane with rectangular flaps; tentacular crown crossed with numerous bauds of maroon and pale
color; ventral lobes of collar produced; in sand or mud
flats
E. vancouveri
30. Dorsal tentacular membrane with notched flaps, dorsal collar
lobes short; tentacular eye-spots conspicuous
Pseudopotamilla occelata (fig. 20,A)
30. Dorsal tentacular membrane a narrow rectangular flap; dorsal
collar lobes long
P. intermedia (fig. 20,B)
31. Thorax with two pairs of pectinate (comblike) branchiae; caudal
appendage (tail) annulated and with setae; operculum formed
of a single series of a few large spines; constructs solitary, conical tubes (fig. 35,A), open at both ends. PECTIrn1DA.E (see 14)
Cistenides
31. Thorax without pectinate branchiae; with many filiform branchiae in longitudinal rows above the oral aperture; caudal
appendage smooth, slender; operculum formed of three whorls
of numerous closely-spaced spines; constructs masses of
tubes more or less firmly cemented tugether. SABELLBRIIDAE
(fig. 19,A) (see 14)
32
32. Opercular spines form a neat, black cone (fig. 19,A)
Phragmatopoma
32. Opercular spines form an open pattern, amber-colored. Sabellaria

..........
...............................
..

.....................

......................
..........

..................

....................
.........................
.

33.
33.

Flabelligeridae (see 14)

With only simple setae; inhabits U-shaped burrows in shales
Stylarioides (Plate 111, no. 1)
With some composite (jointed) setae; inhabits gelatinous or
chitinized tubes, or among the spines of Strongylocentrotus
.Flabelligera
Tentacles retractile into mouth, leaving branchiae exposed and
extending over the anterior end; prostomium often clearly
visible. AMPHARETIDAE (Plate 111, no. 2) (see 14)
Schistocomus
Tentacles not retractile into the mouth; prostomium and branchiae usually not visible except by lifting tentacles away;
branchiae dorsal. TEREBELLIDAE (fig. 17,G and Plate 111,
no. 3) (see 14)
35

................

.........................

34.

34.

..........................

.......................

40

LBBORATORY AND FIEU) 'DZT IN INVERTEBMTE ZOOLOGY

35. Without branchiae; dorsal peristomial lip large, broad. Polyoirrus

35. With branchiae on the dorsal side, posterior to the tentacles 36
36. Branchial filaments simple, cirrifom; thoracic setigerous
segments more than 20; color in life, salmon with bright
red branchiae; in coarsely constructed tubes on under sides
of stones; often with Halosydna brevisetosa . Thelepus (fig. 21)
36. Branchial filaments branched; thoracic setigerous segments 17
ormore. . . . . . . . . . . . . . . . . . . . . . . . . . . a 7

37. First few segments have broad, lateral flaps; branchiae elongate, dendritic, greenish; constructs tubes in rock crevices, the distal end of the tube with an open, reticulated,
sponge-like mass through which the tentacles are extended.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pista
............................

37. Not so.

38
38. With about 25 thoracic setigerous segments; branchiae three
pairs, much-branched, the first pair the largest
Terebella (fig. 17,G)
38. With 17 thoracic setigerous segments
39
39. Chocolate-brown in life; tentacles white, crowded; brauchiae
red; constructs soft mud tubes. , Anphitrite (Plate 111, no. 3)
39. Greenish; tentacles light tan; branchiae deep reddish-brown; in
Eupolyn~~ia
sandy tube6
Cirratulidae (Plate IV, no. 1); (See 14)
40. Body with few pairs of long, tentacular cirri; constructing
Dodecaceria
calcareous rock masses; dark to black in life
40. Body with more numerous tentacular cirri (fig 17,H), not
constructing calcareoue masses; reddish- to greenish-brown
inlife . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 1
41. Anterior end with a pair of stouter, grooved palpi (fig. 22,B)
l'haryx
in addition to the tentacles
(T. parvus on S& Francisco Bay mud flats; a different undescribed species at Moss Beach)
42
41. Anterior end without palpal structures (fig. 22.6).

.....................
.............

.....................
.

...

..............

......

1i

1
,

I
i

PLATE I11

Polyohaete types (after McIntosh) : 1, Cirriformia tentaoulata ( ~ o n t a ~ u(~irratu)

lidae)3 2, Sternas is soutata Bietsch (~ternaspididae)
j 3, Owenia fusifolmis Delle
ChiaJe (in d
n
1, Arenioola marina (L.) (~renlcolidT.).

POLYCHBETA

42. Tentacles extending as far anteriorly as setae (fig. 17,H),

prostomium with eye-spots on either side
Cirratulus cirratus
42. Tentacles first present on the third to fifth setigerous segment
Cirriformia (fig. 22.8) 43
43. Posterior segments with few, stout, thick, dark spines (fig. 23,B);
inhabits burrows or crevices in rocks
Cirriformia l m r i o s a
43. Posterior segments with more numerous, finer, pale spines
C. spirabrahoha
(fig. 23,A); inhabits sand among rocks

..

...............

...
....

Fig. 23.-Setae

Fig. 22.-Cirratulidae:
A , anterior end of
U r i f o I m i a ; B , anterior end of

m.

of posterior seg-

branoha; B , C . luxuriosa.

44. Prostomium without tentacles, although occasionally with a

lobed, flaring membrane (STERNASPIDU, OPHELIIDAE, ORBINIIDAE,
WITELLIDBE, blAINNIDAE, AREDlICOLIDU, OWENIIDU, IDMBRINERIDAE) . . . . . . . . . . . . . . . . . . . . . . . . . . 4 5
44. Prostomium with minute tentacles, which are usually inconspicuous unless rendered conspicuous because of reduced size
of head (fig. 17,F); prostomium usually reduced in size, or,
if large, partly concealed by the first few segments of
body; no palpi. m,
GLYCEEURAE, GONIADIDAE, AMPHINObIDAE, EDPHROSINIDAE
62
44. hstomium with tentacles more or less conspicuous, these
accompanied by a pair of thick, fleshy palpi (except in
Pbyllodocidae). SYLLIDAE (fig. 17,D), NEFU3IDAE (fig. 15,A),
EUNICIDAE, ON[JPBInBE, STAURONERTUDAE, PIiYLxODOCIDAE
73
44. Prostomium with a pair of long, prehensile, tentacular palpi,
somekimes accompanied by a pair of minute anterior antennae.
SPIONIDAE (Plate IX, no. 2) , MAGELONIDAE, CHAETOPTERIDAB
108
44. Prostomium with three tentacles; palpi slender, biarticulate
(Plate V, no. 4)
HESIONIDAE
Short; with long filamentous dorsal cirri (Plate V,
no. 4); purple to brown in life; in ambulacral grooves
of asteroids or under rocks
Podarke pugettensis
45. Posterior end of body provided with conspicuous sternal shield
(Plate IV, no. 2) bordered with stiff bristles; anal branchiae resent; body short, distended. STERNBSPIDAE (see 14
St ernasp i s (Plate IV, no. 2 )
and 447
45. Posterior end without such anal plates
46

...................

.....

..................

......

.............

............

44

LABORATORY AND FIELD TMT IN INVERTEBRATE ZOOLOGY

46. Anterior end f o m a flat plaque; (fig. 24.8)posterior end

terminates in a plaque provided with cirri; segments very
long, the segmental grooves node-like. MALDANIDAE (see 14
and 44)
Clymenella (fig. 24,A)
46. Anterior end with flat, lobed membrane (fig. 24,C). OWENIIDAE (= AWOCHARIDAE)
Oloenia (figs. 24,B,CandPlate IT, no. 3)
46. Anterior and posterior ends pointed or rounded; segments
usually not elongated
47

.................

...........

....................

m.

-.

m.

Fig. 24.-A,
B,
C , head expansion of
D , hooded crotchet
of Lumbrineris. E , mandible of Arobella. F, maxillary apparatus of brabella.

.....

52
47. Proboscis without chitinous jaw pieces; form variable
47. Proboscis provided with dark chitinous jaw pieces (figs
24,E.F); smooth, elongate, cylindrical, resembling an earthworm; parapodia weakly developed or, at most, simple lobes;
dorsal and ventral oirri often tiny
LIJMBRINEFUDAE (Plate Y, no. 1)(see 14, 44, 45, and 46)

.....

, I'
I

48. Parapodia with some hooded crotchets (fig. 24,D); prostomium

without eye-spots
Lumbrineri s 49
48. Parapodia without hooded crotchets; prostomium with or withouteye-spots
51

................
........................

PLATE

Polychaete types ( a f t e r ~ c ~ n t o s :h 1) , Arabella i r i c o l o r Mont. (~umbrineridae);

2 , Scolo 10s armiger ( 0 . F. IRiller) (0rbi3 , 0 h e l i a limacina (H. ~ a t h k e )
(OP*~
4, Podarke p l l i d a Claperkde ( 8 e s i o n i d i d . e

46

LABORBTORY AND FIELD l l E T IN INVXR'l'EBRBB ZOOLOGY

49. Parapodial lobes elongate, erect in posterior parapodia

. . . . . . . . . . . . . . . . .. .. .. .. .. .. .. Lumbriwris
. . . . . . ereeta
..

50
49. Parapodial lobes short throughout
50. Anterior parapodia with some composite hooded crotchets in
addition to pointed setae; prostomial lobe sp~~tulate
L. japonica
50. bterior parapodia without composite setae; prostomial lobe
bluntly conical
L. a o m t a
51. Parapodia with a stout acicular seta projecting from the lobe.

..........................
...................
. . . . . . . . . . . . . . . . . . . . . . . . . . Drilonereis
51. Parapodia without stout projecting acicular setae
. . . . . . . . . . . . Arabella (fig. 17,A and Plate V, no. 1)

52. Body consisting of 3 regions, an anterior and a posterior without branohiae, and a median region with conspicuous branchiae; parapodia with uncini hooks) sometimes of great size
(up to 50 cm.). ARENICOLIDBE see 14, 44, 45, 47)
Arenicola (Plate IV, no. 4)
52. Body consisting of 2 regions which are not always easily
distinguishable from each other (Plate IX, no. 3)
54
52. Body uniform; d o r m sometimes more or less completely concealed by interlacing, cirriform branchiae (fig. 25). no
podia. ORBINII~(= ARICIIDAE) (see 14, 44,
V, no. 2)
53

..................

......

................

Fig. 26.-Orbiniidae:
A , Winereis;
B, Sooloplos.

53. Prostomium terminating in a broad, spatulate lobe (fig. 25,A)

. . . . . . . . . . . . . . . . . . . . . . . . . . . Nuinereis
..................
....
..........

53. Prostomium terminating in a pointed cone (fig. 25,B)

~ c o l o p l o s(PlateV, no. 2)
54. No uncini; branchiae not restricted to posterior region; body
often broad
OPHELIMAE (fig. 26 and Plate V, no. 3) 57
54. Parapodia with tori bearing uncini; body slender; branchiae,
if present, only on a osterior portion. CAPITEUIDAE (see
14, 44, 45, 46, 47, 52j (Plate Lg, no. 3)
55

. . . . . . . . .. . . . . .

Dasybranchus
55. Iarger; with divided branchiae in posterior region
56
55. Smaller; without divided branchiae
56. Thorax consists of 9 setigerous segments, the first 7 with
capillary setae, the last two with uncini; constructs tubes
Capitella (Plate IX, no. 3)
in d flats
1Frrnotomstus
56. 'Phomx consists of 17 or 18 segments

...................

Opheliidae (see 14, 44, 45, 46, 47, 52, 54)

57. Rithout lateral branchiae; with dark eye-spots at sides bePolyophthalmus

tween the parapodia; resembling a nematode.
58
57. Nith lateral branchiae; lateral eye-spots present or absent
59
58. Ventral side with a well-plarked groove (fig. 26)
58. Ventral side without a longitudinal groove; resembling a
maggot.
Travisia
59. Ventral groove posteriorly only; color in life, bronze to purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G O
61
59. Ventral groove throughout length; color in life, otherwise
(fig. 26 and Plate V, no. 3)
Ophel ia
60. Branchiae simple
60. b ~ h i a ebranched
Pect inophel ia

...

..

......
.......................

..

. .. .. .. .. . . . . . . . . . . . .

Fig. 26.-Ophelia.

61. Vith lateral eye-spots between some successive parapodia

...........................

61. Without lateral eye-spots; ventral funnel greatly elongateArmndia

. . . . . . . . . . . . . . . . . . . . . . . . . . Ammotrypane

........

63
62. Prostomium quadrate (fig. 27,A) or subcircular
62. Prostomium pointed (fig. 17,F), provided with four tiny tentacles in a cross at tip: proboscis large, stout, terminat66
ing distally in dark chitinous jaw pieces (fig. 17,F)
63. With branohia-like structures in the form of a recurved cirrus
between the two r a m i of the parapodium (fig. 27,B); prostomium trapezoidal (fig. 27,A), the anterior part with two pairs
of small antennae; proboscis eversible, large.
(see
(Plate VI, no. 1)
64
63. With branchia-like structures in tufts above notopodia (Plate
VI, nos. 2 and 3); parapodial lobes not visible in dorsal
view, beoawe of thick, tufted, setal fascicles; a peculiar
dorsal appendage (caruncle) extending back from the prostom i u m . . . . . . . . . . . . . . . . . . . . . . . . . . . .
65

...

......................

7-

PLATE VI

64. Setae stiff; prostomium with extensive pigmented pattern (fig

27,C); recurved cirri usually present from fourth segment
posteriorly
Yephtys caecoides
64. Setae soft, flowing; prostomium with limited pigmented pat tern; recurved cirri present from third segment back

.................

.......................

N. californiensis

Fig. 27.-A,
Nephtys with everted proboscis; B , l e f i parapodium of
Nephtys showing reourved cirrus; C , Nephtya caeooides; D , l e f t parapodium o f Hemipodus; E, right parapodium of Glyoem robusta.

65. Setal tufts form transverse series, segmentally arranged

(Plate TI, no. 2), broad, almost reaching across d o r m ;
branchiae not conspicuous; EUPEROSINIDAE (see 14, 44, 63).
Bright red to orange in life; crawling over surfaces or in
crevices.
hbphrosine
65. Setal tufts limited to sides of body, not forming transverse
series (Plate VI, no. 3 ); branchial tufts conspicuous,
usually arborescent. AMPHINOMIDAE. Pale red or yellowish in
life; on under sides of stones or in crevices
Pareurytho'e'
66. Parapodia uniramous in anterior part, biramous and larger in
posterior part of body; distal end of proboscis with numer72
ous chitinous jaw pieces. GONIADIDAE (Plate TI, no. 4)
66. Parapodia similar throughout, either all uniramous or all bi67
ramous. GLYCERIDAE (see 14, 44, 62)

.....................

...

...
.............

PLATE VII

Polychaete types (after ~ c ~ n t o s rh )1, Paranatais roses ( ~ c ~ n t o s h(~hyllodocidae)

)
;
Gruhe (~yllidae)
8 3, Odontosyllis
~laperbde(~yllidae).

2. T m o s v l l i s

67. Parapodia uniramous with a single aciculum (fig. 27,D); setae

composite
Hemipodus
67. Farapodia bimous, with two acicula (fig. 27,E); setae both
simple and composite.
Glycera 68
68. Parapodia1 lobes broadest subdistally; segmental retractile
bnsnchiae emerge from the posterior faces of the parapodia.

......................
................

.......................

Glycera amsricanu

...
......

69
68. Without retractile branchiae; parapodial lobes otherwise
69. Parapodia with two presetal and two postsetal lobes
70
69. Parapodia with two presetal and one postsetal lobe
G. capitata
70. Parapodial rami long, with a blister-like branohiae on the
dorsal surface (fig. 27,E)
G. robusta
71
70. Parapodia without such branchiae
71. The two pasterior lobes of parapodium unequal, the dorsal the
G. alba
longer
71. The two posterior lobes of about the same length
G. tesselata

. . .. .. .. .. .. .. .. .. .. .. .. . . . .

........................
..
Goniadidae (see 14, 44, 62, 66)

72. Parapodia cbange abruptly from uniramous in anterior region to

biramous in posterior region; proboscis with chevron at base
(consists of 10 or more V-shaped pieces (Plate VI, rno. 4) Goniada
72. Parapodia ohange gradually; proboscis without chevron at
Glycinde
base
73. With fleshy, usually biarticulated palpi (figs. 15,A; 17,D)
78
73. Palpi relatively slender, tentacle-like (fig 28,A); one pair
100
of annulated prostomial tentacles
73. Without palpi: parapodia uniramous; dorsal cirri often large,
foliaceous, glandular; sometimes resembling the elytra of
a polynoid; proboscis without chitinous jaws. PHYUODOCIDAE (Plate VII, no. 1) (see 14, 44)
74
(&my of the genera are represented by several species; for keys to these reference must be made to
Bartman's papers)
74. With two pairs of peristomial cirri (fig. 28,C)
Eteone
74. With four pairs of cirri at the anterior end (on several segments) (fig. 28,B)
75
75. Segments 1 and 2 at least partially fused; color in life reddish-yellow or rust-colored
Genetyllis castanea
75. Segments 1 and 2 free from one another (fig. 17,E), although
sometimes reduced
76
76. The first three segments form complete rings; prostomium with
a median antenna
&1alia (fig. 17,E)
76. The first three segments dorsally reduced; prostomium with
median antenna; proboscis smooth
kmida
76. The first three segments dorsally reduced; prostomium withbut
median antenna but often with a nuchal papilla (fig. 28,B).
proboscis papillated
77

.........................

..
.
.............

.............

......
.....................
........
......................
.............
............
....................

52

--

LABORATORY AND FIELD TEXT IN INVERTEBRAE ZOOLOGY

77. Proximal part of proboscis diffusely papillated; yellow or rustPhylladoce

colored in life
77. Proximal part of proboscis with longitudinal rows of papillae;
.Amitides
bluishorpurple inlife

...................
..............

Fig. 28.-A, ETothria. Family Onuphidae; B , &itidea williamsi Hartman, Family

Phyllodocidaet C, Etaone dilatae Hartman, Family Phyllodocidae.

78. With an unpaired median prostomial antenna, with or without

paired peristomial tentacles; parapodia uniramous; dorsal
cirri often moniliform (fig. 17,D). SYLLIDU (Plate VII,
nos. 2and3).
78. With only aired prostomial tentacles and peristomial cirri
fig. 157; parapodia biramous; never with monilifo m
beaded) cirri. mIDAE (see 14, 44, 73)
79. Peristomium forms a conspicuous ventral flap that projects
forward under the prostomium; comnsal with pagurids
Cheilonereis (Plate VIII, no.
79. Peristomium does not form such a flap
80. Posterior notopodia have simple stout, uncinate setae (fig.
29,A); peristomial cirri elongate, extend posteriorly to
about segment 8;- tube-building, gregarious.
. . . . Platynereis (= ~ncinereis) agassizi (Plate VIII, no.
80. Posterior notopodia without simple, stout, uncinate setae

. . . . . . . . . . . . . . . . . . . . . . . . 92
. . . . . . . . . . 79

................

1)

. . . . . . . . . . . . . 80
2)

. . . 81

Fig. 29.-Eereida~: A , simple seta; B , homogomph falcigerous (composite) seta;

C , heterogomph composite seta; D , posterior parapodium of Neanthes brandti Malmgren;
E , Neanthes suocinea Frey and Leuckart.

PLATE VIII

California Nereidae: 1 , Cheilonereis cyclurus; 2 , Platynereis agassizi; 3,Lunereis lonqipesy 4 , Eunereis longipesj 5 , Ceratonereis tunicataei 6 , Nereis v e x i l losar 7 , Nereis latesoens; 8 , Nereis prooera; 9 , Nereis zonata; 10, Nereis mediator; 11, Nereis pelagica; 12, Perineis monterea; 13.14, Neanthes l i g h t i .

54

LBBORATORY AND FIELD THlT IN INVERTEBRATE ZOOLOGY

81. Paragnaths absent from one o r the other of the proboscideal

rings ( f i g . 15)
82
83
81. Paragnaths present on both rings
82. Paragnaths present only on o r a l r i n g
anereis (Plate VIII, nos. 3, 4 )
82. Paragnaths present only on maxillary r i n g
Ceratonereis (Plate VIII, no. 5)
83. Posterior notopodia with a few homogomph falcigerous setae
Nereis 84
( f i g . 29,B)
83. Posterior notopodia without homogomph falcigerous s e t a e
90
84. Posterior parapodial d o r s a l lobes g r e a t l y elongate, straplike
Nereis vexillosa (Plate VIII, no. 6 )
85
84. Posterior parapodial lobes not g r e a t l y elongate
85. Dorsum with a broken transverse pattern; small, usually l e s s
N. latescans (Plate VIII, no. 7 )
than 30 mm. long
86
85. Dorsum without broken transverse pattern; usually l a r g e r
86. Dorsal edge of posterior notopodial lobes convex; area VI of
proboscis with four large cones
N. callaona
87
86. Notopodial lobe otherwise

.......................
...............
................

................

.....................

...........

....

........

.......

...

. . . . .. .. .. .. .. .. .. .. .. .. . . . . .
87. A l l paragnaths minute though dark . N. procera (Plate VIII, no. 8 )
87. Paragnaths not minute, some l a r g e r than others . . . . . . . . 88
88, Area V I of proboscis ( f i g . 15) with four o r f i v e large cones
89
disposed more o r l e s s i n a diamond
88. Area VI of proboscis with an oval mass of 6 o r 7 small cones
N. aonata (Plate VIII, no. 9 )
89. Cones more numerous and larger; posterior parapodia with dorsal and v e n t r a l s i d e s of d o r s a l lobes approximately parallel
N. mediator (Plate VIII, no. 1 0 )
89. Area V I I ( f i g . 1 5 ) without mass of smaller cones; p o s t e r i o r
d o r s a l lobe with dorsal edge s h o r t e r than v e n t r a l edge
N. pelagica (Plate VIII, no. 1 1 )

.............

.................

..............

................

90. Area V I with transverse paragnaths

................

Perinereis (Plate VIII, no. 12)

Neanthes 9 1
90. Area VI with only conical paragnaths
91. Marine; posterior parapodial lobes broad, foliaceous (fig.
29,D)
N. brandti
91. I n brackish water:
a ) Posterior parapodial lobes elongate ( f i g . 29,E); M e
N. succinea
Merritt and upper San Francisco Bay
b ) Posterior parapodial lobes short (Plate VIII, no. 13);
i n c o a s t a l streams and lagoons
N. lighti

........

........................

......
..........

Syllidae (see 14, 44, 73, 78)

92. Without v e n t r a l c i r r i
92. With v e n t r a l c i r r i

. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .Autolytus
. . . 93

93. Palpi fused throughout their entire length (fig. 30,C); minute,

93.
94.
94.
95.
95.

a few millimeters long; parapodial cirri inconspicuous or

hogone
absent
Palpi free from one another through most of their length;
larger
94
Prostomium partially concealed by a nuchal hood (fig. 30,B)
95
Prostomium without a nuchal hood
96
Pharynx unarmed; d o r m purple; in algal clumps
Hesperalia
Pharynx armed with a semicircle of papillae (fig. 30,A); dorsal surface pale, with black transverse stripes; in sponges,
and Bxyozoa
Odontosyll is (Plate VII, no. 3)

........................
...........................
..
...............
....
..........

Fig. SO.-Syllidaer
A,
s y l l i s ; C , Exonone.

B, Pippp-

96. Proboscis with a large tooth and a semicircle of hard papillae

Trypanosyll i s (Plate BII, no. 2)
(trepan)
96. Proboscis with a large tooth but no trepan
97

...........

97.
97.
98.
98.
98.

..........
Antennae and dorsal cirri amooth (fig. 30,B) . . . . Pionosyll is
Antennae and dorsal cirri articulated . . . . . . . . . . . . . 98
Parapodia with only simple setae . . . . . . . . . . Haplosyll is
Parapodia in median region contain both simple and composite
setae . . . . . . . . . . . . . . . . . . . Syllis (fig. 17,D)
Parapodia with only composite setae . . . . . . . . . . . . . . 99

Fig. 31.-!Two

types of o m p o a i t s s e t a e of ~ h l e r s i z .

...............
. ..............

99. Composite setae all alike

Typosyll is
99. Composite setae of two kinds, some with a short, some with a
long appendage (fig 31)
Ehl ersia

(Many other genera and species)

56

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

100. With one to five prostomial tentacles (fig, 32,A) which are
simple, cirriform or annulated, but without distinct basal
parts. EUNICIDAE
.lo3

....................

Fig. 32.-Eunicidae
( a f t e r artm man) : A , a n t e r i o r
end of m i c e ; B, r i g h t parapodium of m i o e l&c i r r a t a Webster; C , a n t e r i o r end of Maruhysa.

100. With one pair of long palpi and a pair of prostomial tentacles
that are simple cirriform or annulated (fig. 33,A). STAURONEREIDAE
107
100. With seven prostomial tentacles including 5 larger which
have distinctly annulated bases, and 2 small, simple, at
anterior margin of prostomium (fig. 28,A). ONUPHIDAE (see
14, 44, 73)
101

.........................

........................
Fig. 33.-Stauronereidae:
A , a n t e r i o r end of 5 t a u r ~ n e r e i s g&pilocero~ Moore;
B , r i g h t median parapodium
of S t a u r o n e r e i s a r t i c u l a t u s

Hartman.
A,

101. Without tentacular cirri on first segment dorsally; subintertidal

Hyalinoecia
101. With tentacular cirri on first segment dorsally (fig. 28,A) 102
102. Branchial filaments spiraled on the main stalk
Diopatra
102. Branchial filaments pinnately arranged on the main stalk Onuphis
102. Branchial filaments simple, cirriform
Nothria (fig. 28,A)

.......................

....

....

Eunicidae (= ~eodicidae) (see 14, 44, 73, 100)

. . . . . . . 105
. . . . . . . . . . . . . . . . . . . . . . . . . . Eunice 104

103. Second segment without dorsal tentacular cirri

103. Second segment with dorsal tentacular cirri (fig. 32,A)

104. Heavy acicular setae tridentate at free end; branchiae with

30 or more pinnae . . . . . . . . . . . . . Eunice hawaiensis
104. Heavy acicular setae bidentate at end, branchiae with about 10
pinnae; acicula yellowE. biannulata

.............

105. Branchial filaments not present before about the 100th segment
Palola
105. Branchial filaments present from about the 10th segment;
acicula black . Marphysa (fig. 32,C, and Plate IX, no. 1) 106
106. Branchiae simple, cirrifom throughout . Marphysa sty1obranchiata
106. Branchiae with four to seven filaments where best developed

.........................
.

. . . . . Stauronereidae
. , . . . . . .(see
. . .14,. 44,
. . 73,
. . .100)
? . M.californica

107. larger, brilliantly banded with cowl and cream; prostomial antennae thick, blunt (fig. 33,A)
Stauronereis moniloceras
107. Small (length 17 mm. or less), pale or colorless; prostomial
Stauronereis spp.
antennae long, slender, articulated
108. Body clearly divisible into two or more clearly defined regions (fig. 35,B)
116
108. Body not divisible into regions, although sometimes e single
anterior segment is specially modified. SPIONIDAE (see 14,

...
.....
.....................

4 4 ) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0 9

109. Sixth segment (fifth setigerous) modified (fig. 17,J) provided with special hooks
110
113
109. Sixth segment not modified
110. Branchiae present anterior to modified segment (fig. 17,J);
prostomium with sooty pigmentation; inhabiting narrow
Boccardia 111
crevices and soft shales
110. Branchia first present posterior to modified segment
Polydora (Plate IX, no. 2) 112
111. Branchiae on posterior fifth of body; 29-40 mm. long
Boccardia proboscidsa (= B. natrix)
111. No branchiae on posterior fifth of body; 8-20 mm. long
B. truncata
112. Posterior segments without large stout hooks; constructs soft
tubes; estuarine
Polydora ligni
112. Posterior segments with large stout hooks, burrowing in corralines, shells, etc.
P. a r m t a
113. Anterior end with two pairs of large tentacular structures
(fig 34,A) ; other branchial structures absent; estuarine

.................
.................

............
................
..............
..........................

...............
................

. . .. . . . . . . . . . . . . . . . . . . . . . . .Streblospio

113. bterior end without such structures; branchiae present more

posteriorly . . . . . . . . . . . . . . . . . . . . . . . . 1 1 4

Fig. 34.-A,
Streblos
lutinoola Hartman;
%
- d B , ho
hook of Nerine.

114. Branchiae present from fifth segment back

114. Bnrnchiae present from second segment back

. . . . . . . . . S io
. . . . . . . . . El5

Polychaete types (after ~clc'lntosh)

: 1, ldarphysa
(~ud.and Edw.) (~unicidae);
(~ohnson)(~pionidae)~
3 , Cavitella oawitata (0. ~abrioius)

POLYCHAETA

59

Ierine
. . . . . . . . . . . .. .. Laonice

115. Hooded hooks (fig. 34,B) in notopodia and neuropodia

115. Hooded hooks in neuropodia only

Magelonidae (see 14, 44, 108)

116. Prostomial pdpi greatly elongated, heavily papillated; body

consists of an anterior region of about eight segments, separated from a longer posterior region by a ninth segment
Magel o m
which is quite different. MAGELONIDAE
116. Palpi not greatly elongated, without such papillations; body
consists of two or more regions, each markedly different from
the others (fig. 35,B). CHBETOP!ElUnBE: (see 14, 44, 108). 117

........

Fig. 36.-A,
-,
family Pectinariidae, excavating in sand (after Meo~ntosh)8
B, Chaetovterucl rariovedatus Rbnier et Claparbde in tube (af'ter &c~initie).

1
i

117. Inhabiting large, coarse, chitinous, U-shaped tubes, both ends

of which are exposed; on sandy beaches
Chaetopterus (fig. 35.B)
117. Inhabiting large, coarse, somewhat curved, but not U-shaped
tubes; in eel-grass be&
Mesochaetopterus
117. Inhabiting smaller, annulated, more or less translucent tubes;
Phyll ochaet opte m s
usually in clusters

....................
..........
............

60

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

A LIST OF COMMONLY ENCOUNTERED POLYCHAETA

(Alphabetically arranged by families)

Ampharetidae

D M Schistocom~lshiltoni Chamberlin
Amphinomidae

D M Pareurythoe californicu (Johnson)

Aphrodi tidae

M Aphrodita s p. (typically subintertidal, occasionally

washed in?
Arenicolidae

TB B M Arenicola pusilla Quatref es

TB Arenicola marina (~innaemy
Capitellidae
TB *B M Capitella capitata (Fabricius)
TB M Dasybranchus giganteus Moore
M Eunotomastus sp.
TB Notomastus angulatus Chamberlin
TB B M Notomastus tenuis Moore
Chaetopteridae
TB M Chaetopterus variopedatus Renier
TB M Mesochaetopterus taylori Potts
D M Phyllochaetopterus pro1 ifica Potts
Chrysopetalidae
D M Chrysopetalum occidentale Johnson
D M Paleonotus chrysolepis Schmarda
Cirratulidae
D TB B*M Cirratulus cirratus Miiller
D B *M Cirriformia 1u;curiosa (Moore)
D TB B M Cirriformia spirabrancha (Moore)
D *M Dodecaceria pac ifi ca (Fewkes)
*B !7'haryx parvus Berkeley
M Cauleriella pac ifica (Berkeley)
Eunicidae
D Eunice antennata (Savigny)
D M Eunice biannulata Moore
D M Eunice hawaiensis Treadwell
D Eunice longicirrata Vebster
californi ca Moore
*MM Marphysa
Marphysu stylobranohiata Moore
D M finice (~alola)pallida Hartman
D M Nematonereis unicornis Schmarda
Euphrosinidae
D M Euphrosine spp. (Typically below the intertidal; occasionally washed in)
Flabelligeridae
D M Flabelligera affinis (Sars)

Plabelligeridae (Continued)
D *M Stylarioides eruca (ClaparBde)
D Stylarioides plunosa (0. F. ~uller)
Glyceridoe
TI3 B *M Glycera americana (Leidy)
TB Glycera convoluta Keferstein
M Glycera nana Johnson
TB B *M Glycera robusta Ehlers
TB M Glycera tesselata Grube
TB D B M Henipodia boreal is Johnson
Goniadidae
D M Goniada brunnea Treadwell
D M Glycinde armigera Moore
Hesi onidae
D I Podarke pugettensi s Johnson
Lumbrineridae
TB B *M Arabella iricolor (~ontagu)
TB M Arabella semimuculata Moore
TB M Drilonereis falcata Moore
TB Drilonereis nuda bore
TB D M Lmbrineris sonata Johnson
M Lurnbrineris erecta Moore
TB D M Lurnbrineris japonica Marenzeller
Magelonidae
TB M Hagelona longicornis Johnson
Maldanidae
TB M Clymenella rubrocincta Johnson
Nephtyidae
TB *B M Nephtys caecoides Hartman
TB B M Nephtys californiensis Hartman
Nereidae
D M Ceratonereis paucidentata (Moore)
D Ceratonereis tunicatae Hartman
TB M Cheilonereis cyclurus (Harrington)
TB M anereis longi es Hartman
TB D M Meanthes bran ti BlIalmgren
D Neanthes 1ighti Hartman
TB B Neanthes succinea (Frey & Leuckart )
D *M Nereis callaona Grube
D B M Nereis latescens Chamberlin
D M Nereis mediator Chamberlin
TB D M Nereis pelagica Linnaeus
D M Nereis procera Ehlers
TB *B *M Nereis vexillosa Grube
D M Nereis aonata blmgren
D M Perinereis monterea (~hamberlin)
)
TB D B *M Platynereis agassiai (~hlers

Onuphidae

TB M Dioptatra ornata Moore

62

LABORATORY AND FIELD TEXT IN N

IV
E
R
T
E
m
T
E ZOOLOGY

Onuphidae (Continued)
TB Piopatra splendidissimu Kinberg
M Hyal inoecia (subintertidal; rarely washed in)
Nothria elegans (Johnson)
D M Nothria geophiliformis Moore
D I Onuphis eremita (Audouin and Wne-Edwards)
Opheli idae
D TB M Ammotrypane aulogaster Rathke
D Anandia bi oculata Hartman
M A m n d i a brevis Moore
D M Ophelia limucina jkthke)
D Pectinophelia dil onensis Hartman
D Pectinophelia williamsi Hartman
D M Polyophthalmus pictus (Du'ardin)
D !l'horacophelia mucronata (!$readwell)
D M Travisia pupa Moore
Orbiniidae (= ~rici
idae)
TB D B M Nainereis laevigata Grube
TB D M Scoloplos ameceps Chamberlin
Ckoeniidae (= ~mmocharidae)
D M Oloenia fisiformis della Chiaje
Pectinariidae
TB M Cistenides brevicomu Johnson
Phyl1odocidae
D M Anait ides mediapapillata Moore
D Anaitides mucosa (OErsted)
TB Anait ides williamsi Hartman
TB M Eteone dilatae Hartman
B TB Eteone lighti Hartman
B TB Eteone 1onga Fabricius
TB M Xteone pacifica Hartman
D &la1 ia aviqulisetp Hartman
D M Eulalia virzdis (Linnaeus
TB D M Eumida sanguinea (OErsted
D M Gsnetylli s castanea (Marenzeller)
D H poeulal ia bil ineata (Johnston)
D M dyllodoce ferruginea Moore
Polyno idae
D M Arctonob fragil is

Polyodont i due
D M Peisidf ce aspera Johnson
Sabellariidae
D M Phragmatopom californica (Fewkes)
D B M Sabellaria cemsntarium Moore
Sabellidae
TB M Chone infundi bul ifonni
D *M lhdistyl ia polporpha
TB M &di sty1i a vancouveri
D M Fabricia sabella Ehrenber
D M Mega1onno roulei (~ravier?
TB M Mpicola infundibulum (~ontagu)
D M Pseudopotamilla intermedia Moore
D M Pseudopotanilla occelata Moore
D *M Sabella crassicorni Sars
Sabella media (~ush?
Schiaobranchia insignis Bush
Serpuli due
D M Wrpolnatus gracilis Bush
D M Hydroides uncinatus (Philippi )
*B Mercierella enignatica Fauvel
D M: S a l ~ o i mdysteri (Huxley)
D B M Serpula vermicularis Linn.
D,P Spirabranchus spinosug MooD B M S irorbis spirillun Llm., and other species.
M &niliopsis sp.
Sigali onidae
D M Sthenelais fusca Johnson
Spionidae
D B *M Boccardia proboscidea Hartman
TB M Laonice cirmta Sars
TB B Polydora ligni Vebster
D M Polydora armata Langerhans
M Pygospio californica Hartman
M Rwchospi o arenincola Hartman
M Spzo filicornis Muller
B Streblospio benedicti Webster
Stauronereidue
D M Stauronereis art iculatus Hartman
D M Stauronereis moniloceras Moore
D M Stauronereis minor Hartman
Sternaspididae
D M Sternaspis scutata (Ranzani) (subintertidal; rarely
washed in)
Syllidae
D B M Autolytus varius b a d w e l l
TB+M Ehlersia corputa Rathke
D B M &o one spp.
D TB M Hap osyllis spongicola (Grube)

64

LABORATORY AND FIELD

IN INVERTEBRATE ZOOLOGY

Sy11idae (~ont
inued)
D M Hesperali a californi ensi s Chamberlin
D M Odontosylli s phosphorea Moore
D M Pionosyllis gigantea Moore
D B *M S llis alternata Moore
D M Zllis elongata (~ohnson)
TB B *M Trypanosyllis adamanteus Treadwell
TB Trypanosylli s gemipara Johnson
D M Typosyllis armillaris (~ller)
D Typosyllis hyali nu (Grube)
Terebelli dae
D .M
TB M
D*M
D M
D*M
D TB ld

Amphi trite robusta Johnson

Eupolynnia heterobranchia (Johnson)
Pista elongata Moore
Po1yci rrus californica Moore
Terebella californi ca Moore
Thelepus setosus (Quatrefages)
Class OLIGOCHAETB

The oligochaetes which are likely to be found in this course are

(1) the earthworms (fig. 13,C) and (2) the usually much smaller freshwater worms (fig. 13,B). A key for the earthworms is given below. See
also Altman, 1936, Olinochaeta of Washinaton. For identification of
freshwater oligochaetes refer to Altman (1936).
KEX TO TEE TERFlESTRIAL OLIGOCHAETES (EARTINORIG) FOUND IN BERKEI;eP
By Marjorie E. Oliey

1. Clitellum beginning (anterior edge ) anterior to somite 15

MIEASCOLICIDAE
LUbBRICIDAE 72
1. Clitellum beginning posterior to somite 20

........................

.....

2. Prostomium completely divides peristomium, extending back to

boundarywith second somite
3
4
2. Peristomium continuow behind prostomium
3. Clitellum from somites 27-32
Lumbricus rubellus
Lumbricus terrestris
3. Clitellum from somites 32-37
4. Spermaducal pores (fig. 13,C ) on somite 13
Eiseniella tetraedus typica
5
4. Spermaducal pores on somite 15
Eisenia foetidus
5. Clitellum begins on or anterior to somite 26
6
5. Clitellum begins on or posterior to somite 27
Allolobophora chloroticus
6. Clitellum somites 29-37
6. Clitellum somites 27-34
Allolobophora caliginosus
Diplocardia s ingularis
7. Clitellum encircling body
7. Clitellum saddle-shaped, not extending to ventral side
Dip1ocard i a conununis

..................

............
.........
........

...................
.................
..
..........
.........
.........
.........
......................

Class H I R U D D W
From Lone Tree Pond and other fresh water: Glossiphonia stagnalis
a predaceous leech. Blood-sucking leeches are less common.
Class ECHIUROIDEA
See bibliographic list for references to Urechis caupo Fisher and
MacGinitie (fig. 17,F) found in tidal mud flats. This is the only
species of this group occurring in this region.
Class SIPUNCULOIDEA
Study externally a sipunculid (fig. 13,E). Note the absence of
any segmentation or true setae. The anterior portion is called the
introvert, as it oan be retracted into the rest of the body. Preserved
specimens usually have the introvert retracted. At the anterior end
of a specimen with extended introvert note the mouth surrounded by
tentacles. The anus is not posterior, but is situated on the dorsal
side of the body near the base of the introvert. This is one of the
reasons for doubting the relationship of the sipunculids to the annelids.
Sipunculids are abundant'at Moss Beach and vicinity, mostly living in holes and crevices in the soft sandstone rock and occur in
great numbers in mussel beds on Tomales Point. One undetermined species is found at Point Richmond in San Francisco Bay. The following
identifications and distinguishing characters are due to Dr. H. E.
Brown.
KEX TO THE SIPUNCULIDS OF MOSS BEACH

1. Body nearly cylindrical; conspicuous irregular black spots on the

introvert or on entire body; tentacles very short, unbranched,
extending only three-quarters of the distance around the
mouth
Physcosomu agassizi
1. Body pyriform, enlarged posteriorly; no black spots on body; ten2
tacles larger, fewer, branched, surrounding mouth
2. Dark-brown hooks on the anterior part of the introvert; tentacle
groups four, alike, each with a single base and with many
Dendrostoma petraeum (fig G F )
branches
2. No hooks on introvert; tentacles of two sets, an inner (nearer
mouth) of longer tentacles, an outer of shorter tentacles

...................

.......

.............
.
. . . . . . . . . . . . . . . . . . . . Phascolosoma procerum
LIST OF SIPUNCULOIDEA

B? !b D Physcosom agassiai (Keferstein)

M Dendrostoma pet raeum Fisher
M Phascolosoma procerum ( ? ) Moebius

(? )

66

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

PHYLUM ARTHROPODA

We shall consider four of the classes of Arthropods: (1) the

Crustacea, (2) the Myriapoda, (3) the Insecta, and (4)the Arachnids.
Of these the first three are much more closely related to each other
than they are to the Aracbnida, which differ in the possession of very
different head appendages. The Crustacea have two pairs of antennae,
the Myriapoda and Insecta one, while the hchnida have no antennae,
nor have they mandibles as do the others or appendages corresponding
to the two pairs of maxillae found in the other three subphyla.

The Subphylum Crustacea is distinguished in a general way from

the other arthropods on the basis of habitat in that the crustaceans
are almost entirely aquatic, while the other arthropods are predominantly terrestrial. One obvious morphological distinction of the
crustaceans is their possession of two pairs of antennae, other arthropods having one pair or none. Two major divisions of the Crustacea were
formerly recognized, the klacostraca apd the Entomostraca. The latter,
although it may still be used as a convenient term, is generally considered to be without taxonomic validity, since it comprises at least
fouP groups, each of equal rank with the klacostraca. Following is a
key to the divisions of the Crustacea considered in this oourae.

ICEP TO TBE UJOR GROUPS OF CRUSTACE4

1. Firmly attached, either to a solid substrate (Plate X, no. 3)
or rarely as a parasite which is partly internal (no. 2).
Class CIRRIPEDIA
2
1. Free living, or, if parasitic, generally external
3
2. Non-parasitic; protected by a calcareous shell of several valves
Barnacles (place X, nos. 1, 3 )
2. Parasites of an extremely simplified morphologic nature; usually
found on decapods (Peltogast er on hermit crabs; Saccul ina
on true crabs, Plate X, no. 2)
Order Rhizoce~hala
3. Body appendages leaf-like (Plate X, no. 4) and Ilumerous (except
in Cladocera)
Class BBBNCHIOPODA 4
3. Body appendages not leaf-like, generally differentiated into
two or more tgpes, relatively few in number
7
4. Without a cmpace (brine shrimp) Order Anostraca (Plate X, no. 4)
4. With a carapace
5
5. Carapace broad, shield-like .Order Notostraca (Plate X, nos. 5,6)
5. Carapace conspicuously folded; body laterally compressed
6
6. Carapace two-valved, hinged dorsally, envelopin& the entire body
and head; appendages very numerous
Order Conchostraca (Plate X, nos. 7,7,A)
6. Carapace apparently two-valved, actually not hinged; head uncovered
Order Cladocera (Plate X, no. 8)
by carapace; appendages few

.......................
........

.................

........
...............
..........
. . . . . . . . ... . . . . . . . . . . . . . . .
.
....

............

..

7a
Types of Crustacea. Cirripedia: 1, Acorn barnacle (half of s h e l l removed) ( a f t e r
Damin) j 2 , Saaoulina (Rhizooephalia) on a crab ( a f t e r Delage) j 3, Acorn barnacle
.
(half of s h e l l removed) ( a f t e r Claus); 4 , Artemia ( ~ n o s t r a c a )( a f t e r B ~ O M )Branchiopoda: 5 , Apus ( ~ o t o s t r a c a )(dorsal view; a f t e r B r o ~ 6,
) ~Apus (appendages f r a u t h e
s i d e ) ( a f t e r Lankester) j 7, Estheria ( ~ o n c h o s t r a c a )( a f t e r Lankester) 7a, Estheria
(one valve removed) ( a f t e r ~ a n k e s t e rj ) 8, &&&
la doc era) ( a f t e r Sara).

Types of Crustacba: 1 , Ostracoda ( a f t e r Sars); l a , Cypris (~stracoda)with l e f t

s i s (Mysidavalve removed (after Sars) 3 2 , C c l o s (Copepoda) ( a f t e r ~ a r t o g ) ;3,
a ) (
a otr8taoker); 4, 1s'-(after
Lankester) ; 6. Caprella &pods)
(after ~ a l e s;) 6 , Ampithoe humeralis (~mphipoda)

7. With a bivalve carapace enveloping body and head, externally resembling the Conchostraca, but differing in smaller size (not
more than 2 or 3 mm. in length) and in small number of appendClass OSTRACODA (Plate XI, nos. 1, la)
ages
7. Carapace absent, or present but covering at most the thoracic
8
region

..........
...........................

8. Flattened fish parasites with compound eyes, suctorial mouth and

limbless abdomen; often placed with next class
Genus Ar lus, Class BRCLNCHmRA (fig 47 )
8. With a median simple eye; consisting
-7 o very small swimtning
forms and of parasites which may be somewhat larger; abdominal
:lass COYEPODA (Plate XI, no. 2) 9
region without appendages
8. With paired compound eyes; swimming ur crawling f o m generally
much larger than in Copepoda; abdomen in swimmers almost always with swimming appendages
Class MALACOSTRACA 11
9. Body not abruptly narrowed behind; first antennae (antennules)
short, rarely of more than 8 or 9 segments (both prehensile
in male); largely bottom living
cticoida (fig. 43,D,E)
9. Body abruptly narrowed behind; antennules
Order long fig. 43,A and B),
usually with numerous segments
10
10. Bntennules very long, usually with more than 20 segments; antennae (second) biramous; a single, ventrally located egg sac
Order Calanoida (fig 43,B,C )
10. Antennules moderately long, with 8-20 segments; two laterally
or dorso-laterally located egg sacs
Order Cyclo~oida(Plate XI, no. 2; and fig. 43,A)
11. Carapace, if present, does not fuse with more than 4 thoracic
segments; oostegites present (plates on ventral side of body
of female which form a brood pouch for the hatching of the eggs
and the carrying of the young, figs. 52, 53)
.Subclass Peracarida 12
11. Carapace fusing with all the thoracic segments (fig. 7
r
oostegites
.Subclass Eucarida (including Order Decapoda)
12. Eyes stalked; thoracic limbs biramous
Order bsidacea (Plate XI, no. 3 )
12. Eyes not stalked; thoracic limbs uniramous
13
13. Body usually depressed (flattened dorso-ventrally) ; abdominal
appendages generally all alike, leaf-like, functiqning mainly
for respiration
Order Iso~oda(Plate XI, no. 4)
13. Body usually compressed laterally; abdominal appendages usually
of two sorts, the three anterior leaf-life, used for swimming
and respiration, the three posterior bent backward, used as
a leaping organ. The aberrant caarellids (Plate XI, no. 5)
do not fit this description, but are distinguished by the reduction of the abdomen to a mall stump.
Order Amphi~oda(Plate XI, no. 6)

............

...

.......

................

@=?-

...............

..................

........

.....................
....
................
.........

................

..........

70

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

Class BF$UCHIOPODA

Of the four orders of the Branchiopoda listed above, only one, the
Cladocera, will be frequently encountered during the course. The others,
chiefly to be found in seasonal ponds, may be briefly discussed as follows. The Anostraca are represented by the brine shrimp, Arternia salina
(Plate XI, no. 4), found in this region in the highly concentrated sea
water of pools for the commercial production of salt by evaporation.
The brine shrimp is much used as food for small fish in aquaria. Other
genera, commonly known as fairy shrimps (fig. 36). occur in enormow

Fig. 36.-Male

of Branchinecta

Lynch (after Lynch).

numbers at times in seasonal freshwater ponds. Living specimens of

Artemia will be seen in the laboratory. The Notostraca are represented
by Lepidurus, and Apus (Plate X, nos. 5, 6) also occurring in seasonal
freshwater ponds and will probably not be encountered in the coastal
area. A member of the Conchostraca, Estheria (nos. 7, 7a), is also
widely found in seasonal freshwater ponds and is often mistaken for a
small bivalved mollusk which its external appearance strongly suggests.
Order Cladocera
On a specimen anaesthetized with nicotine and on mounted stained
specimens make out (see fig. 37 )
carauace (not truly bivalved),
compound
, and the large antennae (second antennae). The head ends
in a depressed rostrum near which are the vestigial first antennae or
antennules. Above the origin of the antennae is a ridge known as the
fornix (see Pleuroxus, fig. 39). A notch or transverse groove, the
cervical sinus (fig. 37), marks the dorsal junction of head and body
in some species. A black spot, closely associated with the brain in
some species, usually below and behind the compound eye, is the single
eye o r ocellus.
The head may be in line with the body or depressed. The posterior
end of the shell-or carapace may bear a spine as-in Daphnia (Plate X,
no. 8). The spine may lie at the postern-dorsal angle or at the
postero-ventral (= infero-posteal) angle, when the posterior end of
shell is truncated. The postero-ventral angle may be acute, rounded,
smooth, or toothed. The nature of the shell surface is often of diagnostic importance. The number of setae on the various joints of the
two branches of the antennae beginning at the proximal end is made
into a formula of systematic value. The formula for Daphnia is
0 0 1 3 . M e it out for your specimen.
1-1-3

m,

......

bmin

-.-......
-.

ant ennule .-- - - -1st thor. app.

./$

s h e l l gland
tao. a*.

-- --- -....

3rd thor. app. ---

-.-A

,
,

terminal olaw

intestine

Fig. 37.-Simocephalue

(after Shipley and k c ~ r i d e )

Identify a specimen by means of the key in b d and Whipple. If

this is not immediately available use the key below but identify at
least one species using Tnard and Vhipple.
A KEY M !IEE COMMON GENWB OF FRESHWATER CLADOCERA

1. Antemules (fig. 37) of female very small (save in Moina, where

large, freely movable, and inserted near or behind middle of ventral surface of head, fig 38)
Daphniidae 2
1. Antemules of female low. immovable. inserted near anterior end of
ventral surface of head-and recurved forming a biakp(f ig.-39,~).
Bosminidae
Bosmina (fig.
. - 39.A)
- .
1. Antennulea of female long, freely movable, inserted at anterior
surface of head (fig 39,D )
.Family Macrothricidae
1. Antennulea of female inserted on ventral surface of head.v mrtlv
s covered by fornices which form a beak with the rost&
extending downward in front of antennules (fig. 39,B). Chydorie......................
la
.- .- .- .- .- .- . -la. Rostrum long and slender, often upcurved at tip; posterior margin short, often toothed
Pleuroxus (fig. 39,B)
2. Cervical sinus present (fig. 37)
3
2. Cervical sinus abaent
Daphnia (Plate X, no. 8)

...........

.................
.......
.

...............
..........

I
I

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

72
3. Antemules very small, hardly movable if at all; head small,
depressed, with a rostrum
4
3. Antemules large, freely movable; head large, extended, thick
in front; without a rostrum; valves small, not covering body;
long ciliated spines from dorsal side of postabdomen
Moina (fig. 38)

..................

.........................

Fig. 3 8 . - e

(after ~ a i r d )

4. Head moderately depressed; valves more or less quadrate

.....

5
4. Head greatly depressed; eye almost directly below bases of antennae; valves oval or round
Ceriodaphnia

...........

A. Bosmina loneirostris

B. Pleuroxis dsnticulatus

D. Macrothrix
Fig. 39.-Types

roses

of Cladocera (after Birge).

5. Large; valves subquadrate with rounded angles; body large and

heavy; yellow to yellow-brown
Sinocephalus (figs. 37, 39,C)
5. Smaller; valves almost rectangular? posteroventral angle of each
produced into a spine; color variable
Soapholeberis

...

.......

OSTRACODA

73

LIST OF COhlMON GENEEiA OF BRANCEEOPODA

(All except Artenia in fresh water)
hostraca

Cladocera

Artenia salina (in highly concentrated salt water )

Branchinecta

Notostraoa
Apus
Lepidurus

Bosmi na
Ceriodaphnia
Daphn i a
Macrothrix
Mo i na
Pl euroms
Scapholeberis
Simocephalus
-

Conchostraca
Estheri a

Class OSTRACORA
Examine whole mounts and living specimens. Note the truly bivalved
shell (Plate XI, no. l),which entirely encloses the head and body, and
the absence of segmentation or of leaf-like appendages (fig. 40 and
Plate XI, no. la), These characters differentiate the ostracods from
other entomostraca.
Remove one valve to identify such structures as time allows. Refer
to figure 40 of Asterope.

st t h o r , app.
Fig. 40.-An
ooetraood atero rope) with left
valve remwed (aPter Sara).

Be able to recognize ostracods when encountered. We shall not attempt to identify them to genus or species.
Class COPF9ODA
Laboratory Exercise.-Anaesthetize your specimen with a drop of
dilute nicotine solution. Note the enlarged anterior portion of the

74

LABORATORY BM) FIELD TlB!T IN J I I V I B m T E ZOOLOGY

body in front of the movable nrticulation, known as the metasome, and

the smaller posterior urnsome (fig. 41). These terms are safer than
thorax and abdomen, about the limits of which opiniom differ. The
(or furcae).
last or anal segment bears a pair of caudal or furcal
See the large uniramous first antennae (antennus&d the smaller,
often biramous, second antennae. How many pairs of thoracic legs (swimming legs) do you find (ref.ro fig. 41)? These legs, particularly the
last (fig. 44). are often of very great systematic importance.
ant ennul e

metas ome

uros ome

Fig. 41.-Diagrammatic figure of female oalanoid oopepod ( a f t e r Giesbrecht and Sohmeil).

Study stained mounts of appendages of a large freshwater species

(genus D i a p t o m s ) . Using both low and high power of the compound microscope see the antennules of male and female. Note the special features
of the right (clasping) antennule of the male, especially the process
on the antepenultimate segment (fig. 44,D,E) and the heavy spines on
segments 10, 11, and 13. On a swimming leg make out basi~oditeand two
distal branches. the exo~oditeand endo~odite.How many segments in
each of these?
Study the fifth legs of the female (fig. 44,B) noting that they
are greatly modified but symmetrical. Note the great asymmetry of the

COPEPODA

75

male fifth leg (fig. 44,A). The position of the spine on the outer side
of the second segment of the right exopodite is important in classification as also the nature of the medial apical process of the terminal
segment of the left exopodite (fig. 44,A).
Figure 42 illustrates the functions in copulation of the peculiarly
modified right antennule of the male, the male fifth legs, and the
usually asymmetrical urosome of the male.

Fig. 42.-Copulation
i n Diaptomus (after
wesenberg-~und): A, function of clasping antennule of male) B, function of urosome and
fifth legs of male.

See examples and learn the general distinguishing characters of the

three subclasses, the Calanoida (figs. 41, 43,~,C), Cyclopoida (fig.
43,A), and Harpacticoida (fig. 43,D,E). The calanoids are usually
larger (figures in 43 entirely out of scale), the antennules very long
(with 25 segments), the urosome sharply distinct from the metasome,
which is considerably longer and broader than the urosome (furcal rami
included in measurement, but not setae); the cyclopoids (fig. 43,A) are
smaller, the antennules shorter, usually with less than 17 segments,
the urosome and metasome distinct and of about the same length; the
harpacticoids (fig. 43,D,E) are very small, the antennules very short
with few segments and the urosome is not conspicuously different from
the metasome
A key to the orders of Copepoda is included in the key on p. 69,
but is repeated below, together with the identification of some local
genera and species,

76

LABORllMRY AND FIELD TEXT IN INVERTEBRQTE ZOOLOGY

A KEX TO TBE MOST COMMON COPEPODA OF

!E3
l SAN l?RANCISCO

MY REX4ION

1. Body not abruptly narrow behind; urosome nearly as wide as metasorne

(fig. 43,D,E); f i r s t antennae short, r a r e l y of more than 8 o r 9
segments (both prehensile i n male); l a r g e l y bottom-living.
Harpacticoida 2

.........................

Fig. 43.-Types
of copepods (variously magnified) : A, C clo s strenuue Fischer
(after Wesenberg-lund)i B, DLa t o m s W e r z e skii Richard ~$d-Wesenberg-Lund)g
C, D i a ~ t a u sgncilis Sara
Atthyella illinoisensis Forba
(after ~oker)

77

COPEPODA

Fig. 44.-Fifth
legs and r i g h t antennules of l o c a l species of
variously magnified: A, f i f t h legs of male of
oaduous Light; B, f i f t h legs of famale of same species$ C , r i g h t
f i f t h l e g of male of Diaptomus s i o i l o i d e a L i l l j e b o r g , much enlarged; D , antepenultimate segment of r i g h t antexmule of male of
Diaptomus novamexicanus Herriok; E , same of D. s i c i l o i d e s .

1. Body abruptly narrowed behind, urosome narrow; antennules long

(fig. 43,A-C), usually with numerous segments; chiefly freeswimming
3
2. Fresh water
Canthooalptus ( ~ t t h ~ e l l a(fig.
)
43,D,E)
2. Marine; orange red; present in high tlde pools

..........................

.....................

Tigriopus oal ifornicus

3. Bntennules very long, usually with more than 20 segments (figs.

41, 43,B,C); a single, ventrally located egg sac.
Order Calanoida 5
3. Antennules moderately long, with from 8-20 segments, usually
from 10-160 two laterally or dorsolaterally located egg sacs
(fig. 43.8)
Order Cyolomida 4

........................

.................

78

LBBORATORY AND FIELD TEgT IN INVWTEBRBTE ZOOLOGY

4. Freshwater; free-living; with r e l a t i v e l y narrow body.

Cyclops and related genera.
4. k i n e , on surface of Call ianassa; red; female bmad (figs. 45.46 1;
Clausidium (figs. 45, 46)
male much smaller (fig. 45)

...................
......

Fig. ----46.
Haddon: A , female
Fig. 45.-Clausidium vanoouverense Haddon, dorsal view of i n ventral v i m ; B , f i r s t thomoio l e g , magnified t o
olasping pair.
show modifioatiom; C , a portion of t h i s more highly
magnified t o show nature of suckers.

5. Urosome of female 4-segmented (fig. 41); metasome showing f i v e

d i s t i n c t somites behind head; l e f t f i f t h leg of male enlarged;
marine
Calaws
Commonest pelagic species
C. iJ' marchic u s
5. Urosome of female 3-segmented; metasome showing 3 o r 4 d i s t i n c t
somites behind head; exopod of right f i f t h leg of male tipped
Diaptows 6
with long, curved claw (fig. 44,A.C); fresh water
6. Spine of right f i f t h leg of male nearly s t r a i g h t , located near
d i s t a l end of second segment of exopodite (fig. 44,A); medial
process of l e f t exopodite of male f i f t h leg a slender curving, f l a t spine (fig. 44,A); t h i r d segment of exopod, of female
7
f i f t h l e g d i s t i n c t (fig. 44,B)

.........................
............
..

................

6. Spine of right fifth leg of male curved, much nearer middle of

segment (fig. 44,C); medial apical process of left exopod
digitifom; third segment of exopod of female fifth leg ves9
tigial, represented only by seta and spine
7. Antemules with setae more numerous than usual, two each on
D, caducus (fig. @,A)
segments 13-19, and four on segment 2
7. Antennules with but one seta on segments 13-19 and three on
8
segment 2
8. Larger; process on antepenultimate segment of right male antennule much longer than penultimate segment, slender distally;
right fifth leg of male with a prominent rugose swelling on
D. eiseni
median margin of second basal segment
8. Smaller; process on antepenultimate segment of right male antennule rarely ae long as penultimate segment, thicker distally;
no such swelling on right fifth foot of male
D. franciscanus
9. Antepenultimate segment of right male antemule without hyaline
lamella but with a short process abruptly outcurved near tip
(fig. 44,E); fifth leg of male with prominent hyaline lamella
on distomedial margin of basal segment of right exopod (fig.
44,C); this segment nearly twice as long as wide
D. siciloides
9. Antepenultimate segment of right male antennule with both hyaline lamella and a process; process with gentle distal curvature (fig. 44,D); basal segment of right exopodite of male
fifth leg wider than long, without hyaline lamella.
D. novamexicanus

..........

..
..........................

........
..

........................
LIST OF COMMON LOCAL COPEPODA

Order Haroacticoidg

F Atthyella
M Pigri opus cal ifornicus (Baker) (= T. triangulus Campbell)
Many other unidentified genera and species, freshwater
and marine. For marine species see Monk, Trans. American
1941, p.. 75.
Micros.
Order Calanoida
M Calanus finmarchicus Gunner
M Calanus spp.
Other marine species
F Diapt omus caducus Light
F Diaptomus eiseni Lilljeborg
F Diaptomus franciscanus Lilljeborg
F Diapt o m s novamexicanus Herrick (= D. washingtonensis)
F Diaptoms siciloides Lilljeborg
Order C~clo~oida
M Clausidiunr vancouverense Haddon
F Cyclops spp.
b y other freshwater genera and species

- &.,

80

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

Class BRANCHmRB or ARGULOIDEA

These animals (fig. 47) are ectoparasites of fish, freshwater and

marine. Two species are commonly encountered in this region, Argulus
japonicus Thiele (fig. 47) common on goldfish, and the much larger A.
pugettensis Dana common on the various types of marine perch.
The family Argulidae has been recently reviewed by Meehean (1940,
pot, U.S. National Museum). The group is variously considered to represent an order or subclass of the Copepoda or a separate class.
See the demonstration specimens and be able to recognize the two
common local species.
1 s t antenna

Fig. 47.-Ar
class s-ohi-er

ulua, the fish louse,

~aehrn).

Class CIRRIPEDIA
Order Thoracica, Barnacles
(Refer to Jirriuedia .ofPwet Sound, by Dora Henry, 1940. )
On a goose barnacle (~e~as,
fig. 48,A) make out the five platea
making up the armor, a pair of large scuta (sing. scutm) at the stalk
end, a pair of smaller terara (sing. t e r m ) at the free end, and a
single narrow median carina at the hinge side. The homologues of these
five plates are present in the sessile barnacles arr well as several
other plates. In Yitella, the leaf barnacle, many accessory platelets
are found, some of which may be homologous with the extra plates in the
sessile barnacles.
On the sessile or acorn barnacle, Balanus glandula (figs. 48,%C,

ala

tergum
oarina
scutum

rostrum

a r t i c u l a r ridge
oarina
carino-lateral
lateral
f o r depressor muscle
oavity

beak

purple patch
a r t i o u l a r ridge
a r t i c u l a r furror

f o r depressor muscle

s o u t a l margin

Fig. 48.-Types
Henry) 1 A ,
C , Balanus 3same inner view.

of barnacles and terms used i n t h e i r description (c-E seer

B,
Henry:
D , acutum of
-,
inner view: E , tergum of

a,
a goose barnaole;

Plate X, no. 1 ) - make out three calcareous regions: ( 1 ) the basis by

which it is attached, ( 2 ) the parapet o r &&, and (3) the movable
portion, the valves, shutting the aperture a t the f r e e end of the wall.
The wall of a s e s s i l e barnacle consists primitively of 8 plates,
each known as a co
ment The one f a r t h e s t from the head (dorsal
o r posterodorsal is the carina ( f i g s . 48,B,C), homologous with that
i n Lepas; that a b the opposite end is the rostrum. Those on e i t h e r side
of the oarina a r e the carino-laterals, and the pair next the rostrum the metro-laterals, while the middle compartments are known
a s l a t e r a l s . Various oombinations and eliminations of these primitive 8

.
f=?=-

82

LABORATORY AND FIELD TMT IN INVERTEBRATE ZOOLOGY

plates have occurred in different genera. In the genus Balarus (fig.

49,B) the rostro-laterals are fused with the rostrum. In Chthamalus
(fig. 49,A) the carino-laterals are lacking and in Tetraclita there
are only four plates, the rostro-laterals being fused with the rostrum
and the carino-laterals absent. The exposed median triangle of each
compartment (fig. 48,B) is known as a paries (pl. parietes.) The edges
of a compartment which overlap another compartment are known as radii

Fig. 49.-Schematic cross seotions t o show

arrangement of plates in I ; b t u ( A ) and
Balanus (B).

(sing. radius) if the lines of growth are different from the paries,
otherwise they receive no special designation. The edges of a compartThe rostrum
ment which are overlapped are known as aJgg (sing.
The lateral
and carina are symmetrical, typically each with two
compartments are symmetrical each with a radius on the posterior
(carinal) side and an ala on its anterior (rostral) side. In Chthamalus the rostrum is normal with two alae (fig. 49,~). In Balanus (fig.
49,B) and Tetraclita, however, it seemingly has two radii, but these are
really the radii of the rostro-laterals which are joined with it. The
carina has two alae in all cases since, while the carino-laterals are
lacking in some genera (Chthamalus and ~etraclita),they are never
fused with the carina as are the mstro-laterals with the rostrum
(Balanus and Tetraclita). The movable upper portion consists of two
pairs of plates, two longer anterior scuta (figs. 48,D and 49), and
two shorter posterior ter~a.Numerous characters important in the
classification are found on the inner bases of these plates (see fig.
4%D,E).
Identify your specimen using the key below. At some future time
attempt to identify other species by means of Henry's key (1940) or
using Pilsbry, U.S. Nat. Mus. Bull. 93. Our local barnacles are not
well known and species may be found which are not in the key.

a).

&.

m TO rn BARNACLES
I

....................
....................
.......................
.............

1. Stalked (goose barnacles)

2
1. Sessile (acorn barnacles)
4
2. With many plates and thickly scaly peduncle; attached to rocks
Mitella polymerus
2. Vith but five plates and smooth, slender peduncle; attached to
3
seaweed or floating wood (fig. 48,A)

CIRRIPEDIA

83

3. Plates t h i n and papery; carina ending below i n f l a t disc

Lepas fascicularis
3. Plates thicker; carina forked below
Lepas h i l l i
4. Very small, l e s s than 114 inch long, without v e r t i c a l ridges;
front and hind p l a t e s (rostrum and c a r i n a ) overlapped on both
Chthamalus fissus
s i d e s by l a t e r a l p l a t e s (fig. 49,A)
(rostrum) overlap4. Iarger; with v e r t i c a l ridges; a n t e r i o r
5
ping l a t e r a l p l a t e s (fig. 49,B)
5. Wall red, f i n e l y ridged, consisting of but 4 compartments, the
.
divisions between which a r e hardly discernible externally.
Z'etraclita squamosa rubescens

.......................
..........

......
...............

.................

5. Wall grayish o r white, variously ridged, consisting of 6 compartments, the margins of which may not be discernible externally
66. Outside of wall with numerous f i n e ridges (see f i g u r e below);
.Balanus cariosus
suture between scuta strongly sinuate
6. Outside of wall without o r with few ridges, these r e l a t i v e l y
heavy; suture between scuta feebly o r minutely sinuate
7
7. Small t o medium s i z e , usually about 1/2 inch i n diameter o r
smaller
8
7. Usually much larger; a t low t i d e mark
Balanus nubilis (fig. 48,D.E)
8. Commonly with d i s t i n c t ridge$ on wall; common high intert:dal
species
&lanus glandula
8. Val1 smooth; r a r e lower i n t e r t i d a l species
Balanus crenatus
(See Henry, 1940, and the special report of C. D. Michener f o r
more technical differences between species of Balanus)

...........................

.....

...
...........................
...

...................
...

LIST OF CIRRIPEDU

Order Rhi zocephala

Unidentified p a r a s i t i c forms (Sacculina, Peltogaster o r other
Rhizocephala ) on crabs, espec i a l l y on hermit crabs.
Order Thoracica

Balanus cari osus P a l l a s

Balanus crenatus Brugiere
Bal anus gl andul a Darwin
Bal anus nub i 1 i s Darwin
Chthamalus f i s s u s Darwin
Lepas h i l l i Leach
Lepas fascicularis E l l i s and Solander
Mi t e l l a pol ymerus Sowerbv
~ eracl
t i t a sq~amosarubescens
Darwin

The Thatched Barnacle, Balanus

cariosus Pallas (from ~ u b e r l e t )

84

LABORATORY BND FIELD TEXT IN INVERTEBRQTE ZOOMGY

Class UCOSTRACA

Three subclasses of the Malacostraca, the Isptostraca, the Hoplocarida, and the Spcarida, will not be encountered. The other two,
Peracarida and Eucarida, include many important elements of the fauruns
studied.
Subclass Peracarida
Two orders, Mysidacea (Plate XI, no. 3) and Cumacea (fig. 50),
include minute forms not important in such studies, although representatives of each may be encountered. A third order of m m y authors, the
Tanaioidea (fig. 51), is included in this syllabus with the Order
Isopoda as the Suborder Chelifera.

Fig. 50.-Diastylis

,a

cumroean ( a f t e r Sara).

Order Iso~oda
You will be given specimens of the five free-living suborders,
Chelifera, Flabellifera, Valvifera, Onisooidea, and Asellota. Distinguish them by means of the key below adapted from M. A. Miller (1940)
and Richardson (1905)

Fig. 51,--,

Suborder Chelifera

(= Order Tanaioidea) ( a f t e r ~ichardaon):

A , dorsal view; B , chela enlarged.

1. First pair of legs chelate (fig. 51); uropods terminal

.....................

Chel ifbra (~anaioidea)

1. First pair of legs without chelae; uropods terminal or lateral

~ p p

ISOPODA

85

. . . . . . . . . .3
. . . . . . . . . . . . . . .4

2. Uropods l a t e r a l ( f i g 58 ) o r v e n t r a l ( f i g 54 )

2. Urnpobe terminal ( f i g s . 52, 53, 61)

3. Urnpods l a t e r a l , v i s i b l e dorsally, with flattened branches

forming with telson a caudal fan ( f i g . 58)
Flabellifera f~ynothoidea)
3. Uropods ventral, not v i s i b l e dorsally, modified t o form l a r g e r
operculer p l a t e s hinged l a t e r a l l y and covering the pleopods
(fig. 54)
Valvifera (~dotheoidea)

...................

................

I
!

62i~Q*ia-

i s Say, Suborder
Hme)

i
4. Aquatic f o m ; pleopods branchial ; abdo~ninalappendages usually
covered by l a r g e r opercular p l a t e s formed by an a n t e r i o r p a i r
of pleopods; abdominal segments fused
Asellota (~selloidea)( f i g . 52)
4. T e r r e s t r i a l forms; pleopods uncovered, adapted f o r a i r breathing;
sometimes provided with tracheal t r e e s ; abdomen with 6 welldefined segments (the last fused with t e l s o n )
Oniscoidea ( f i g . 5 3 ) ~

.................

......................

86

LABORATORY AND FIELD TEXT

IN INVERTEBRATE ZOOLOGY

In your specimens note the characters of the order, the depressed

bodies, the long thorax with seven pairs of legs (peraeopods or periopods), in addition to the one pair of maxillipeds (figs. 52, 53), and
the short abdomen. The last abdominal segment is united with the telson and other reductions in visible numbers of abdominal segments will
be encountered (e.g., fig. 54). See the flat leaf-like pleopods functioning for respiration and for swimming (figs. 53, 54).

Thr'ee free abdominal segments

es

F i g . 54.-Idotea baltioa
Pallaa, Valvifera : ventral
view of abdomen with l i d l i k e uropod ranwed on one
side (af'ter ~ a l e s ) .

Fig. 53.-Porcellio scaber Latreille, Suborder Onisooidea ( a f t e r Van ~ a m e )

On a female make out the otistenits (figs. 52, 53), or marsupial

plates, arising from the inner bases of the thoracic appendages. These
form the mrsunium in which the eggs develop.
Study the mounts of the head appendages of Sphaerona. Make out
antennules, antennae, mandibles, maxillules or first maxillae, maxillae or second maxillae. The maxilli~edsare the first thoracic appendages.
Make yourself familiar with the following valuable works of reference listed in the terminal bibliography: Richardson (1905), Monograph
of Isopobs of North America (naturally far from complete); Miller,
M.A. (1938), Comparative Ecological Studies on the Terrestrial Isopob
of the San Francisco Bay region (includes a key to species); Van Name
(1936), Monograph of Land and Freshwater Isopods.

1. Legs of first pair cheliform (fig. 51), uropods terminal; pleopods,

when distinctly developed, exclusively natatory. Chelifera (often
considered a separate order, Tanaioidea)
2
1. First pair of legs not cheliform
4

............
................

Suborder Chelifera

2. Two pairs of pleopods; abdomen of 3 segments; uropods of 2 articles

Pancolus californiensis
2. Three pairs of-pleopods;abdomen of 5-6 segments; uropods of 3-7
articles
3
3. Abdomen of 6 segments
Anatanais
3. Abdomen of 5 segments
Z'anais (fig. 51)

..................
...........................
.................
..............
4. Uropods lateral (fig. 58) or ventral (fig. 54) . . . . . . . . . 5
4. Uropods terminal (figs. 52, 53, 61) . . . . . . . . . . . . . . -20
5. Uropods forming with terminal segment of abdomen a caudal fan
(fig. 58); most pleopods natatory (Cymothoidea or Flabellifem.... ..........................lo
5. Uropods valve-like, inflexed, arching over pleopods (fig 54)
which are largely branchial (Idotheoidea or Valvifera)

. . . . .6

Suborder Valvifera

6. Palps of maxillipeds with five articles,(fig. 55,A); epimera

of thoracic segments free (figs. 56,A-C)
Pentidotea 7
6. Palps with four articles (fig. 55,B), epimera of all save first
thoracic segment free
Idothea

.....
.................

Fig. 55.-A,
mandibular
palp of Pentidotea woaneaenskii showing f i v e arfic l e e ; B, same of Idothea
r e c t i l i n i a showing four

7. Posterior margin of telson concave, often deeply so, postero-

. Pent idotea resecata

. . . . . . . . . . . . . . . . . . . . . . . . . . .8

lateral angles greatly exteaded (fig. 56,C)

7. Posterior margin of telson convex, postero-lateral angles pot

extended

I1

88

LABORATORY ANJI FIELD 'IXrT IN INVERTEmTE ZOOLOGY

8. Telson broadly rounded at posterior end, with a very small

median tooth; commonly dark green or brown: found among seaPent idotea uosnesenski i (figs. 56,A,B)
weed
8. Telson more nearly rectangular, postero-lateral angles fairly
9
distinct, a distinct median tooth at posterior end

...........

......

.2

Fig. 56.-Idoth

oidea (after Richardson): A , B

woenesenskii Brandt ;
resecata ~timpsonrD, i d o t h r s r = Fnia ~ookington.

9. Sides of thorax approximately straight; tho&

not half again
as wide as telson; long diameter of eye about twice short
diameter
Pentidoteawhitei
9. Sides of thorax convex; thorax twice aa wide as telson; long
Pentidotea stenops
diameter of eyes five times short diameter

..................
.
Suborder Flabellifera

10. First pair of legs subchelate, larger and stouter than the
others; mouth parts sucking; uropods lateral and superior
Anthuridae (fig. 57 )
10. First pair of legs resembling others; uropods not superior;
mouth parts n o d
11

......................

.....................

.
11. Abdomen, including telson, of 6 short segments (often only 5
visible ) (figs. 58,C,D ) . . . . . . . . . . . . . . . . . . . 12
Fig. 67.-C
thura oarinsta xrdyer,
A n t h u r i w e r Iiiohardeon)

11. Abdominal segments reduced in number and imperfectly separated, ugually 2 visible segments (fig. 58,~);uropod
Sphaeromidae 14
with one immovable branch

...........

ISOPODA

89

12. !Pwo pairs of antennae unequal in length (fig. 58,C); uropods

well developed, forming a fan with the telson and last abdominal segment (fig. 58,C); speckled or dark gray in
color
Ci rol ana 13
12. Two pairs of antennae equal or subequal; uropods small, outer
branch rudimentary; body white, eyes black; boring in wood

.......................

. . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnoria

F i g . 58.-Sphaeroraidae (after Richardson) : A , FZoephaeroma thermophilum Richardson;

B , maxillary palp of Exoaphaerol~ashowing lobes; C-E, Cirolana harfordi Lookington;
F , Cirolana linguiformis Riohardson; G , Dynamenella.

13. Fifth abdominal segnibnt with lateral parts covered by fourth

segment; anterior margin of telson approximately straight
(fig. 58,C,E)
Ci rolana harfordi
13. Lateral angles of fifth segment free; anterior margin of telson with three angular projections (fig. 58,F)

................

. . . . . . . . . . . . . . . . . . . . . .Cirolana linguifrons
14. Posterior margin of telson entire (fig. 58,A) . . . . . . . . . 16
14. Telson with a median emargination (fig. 58,G) . . . . . . . . . 15
15. Surface of abdomen smooth; flagellum of second antenna with
10 articles . . . . . . . . . . . . . . . . . . . . Dynamenella
15. Surface of abdomen with tubercles; flagellum of second antenna
with less than 6 articles . . . . . . . . . . . . . . . Dynamene
16. Second to fourth articles of palp of maxilliped not produced
into lobes; surface densely granular, dark brown; in mud
flats, and in colonies ofdlercierella at Lake Merritt

. . . . . . . . . . . . . . . . . . . . . . . Sphaeroma pentodon
.....................

16. Second to fqurth articles of palp produced into lobes (fig.

58,B )
&osphaeroma 17
17. Body broadest posteriorly, thorax ridged with three longitudinal rows of small tubercles; branches of uropods expanded

. . . . . . . .. .. .. .. .. .. .. .. .. .. .. .. .. .EGcosphaeromaamplicauda
. . . . . . . . . . 18

17. Body otherwise

18. Extremity of telson produced in a rhomboid process; surface of

Bosphaeroma rhomburum
body punctate; color whitish yellow
19
18. Extremity of telson not produced in rhomboid process

...

.....

90

LABORATORY AND FIELD

TEgT

IN llVXRTEBRATE ZOOLOGY

19. Surface of abdomen with tuber ale^; all thoracic segmenta but
first marked with four conspicuous brown spots and with two
spots on first abdominal segment
kosphaeroma octoncum
19. Surfaoe of abdomen without tubercles; body ovate, twice as long
as wide: head three times as wide as long

....

. . . . . . . . . . . . . . . . . . . . Exosphaerom

oregonensi s

20. Littoral; abdominal segments fused (figs. 59, 60) (Suborder

Asellota)
20. Subralittoral or terrestrials abdomen consisting of 6 welldefined segrsents (Suborder Oniscoidea )

. . . . . . . . . . . . . . . . . . . . . . . . . . 21
. . . . . . . . . . . 24

Fig. 69.-lduxnm k h e r i Goodsir

(after Riohardson)

Suborder Asellota
21. Mite-like in appearance; last three thoracic segments short,
constricted
.Munna (fig. 59)
22
21. Not mite-like; last three thoracic segments not constricted
23
22. Second pair of antennae long, with dtiarticulate f
22. Second antennae short; flagellurn reduced (figs 60,B,lC
1

..................

..

. . . . . . . . . . . . . . . . . . . . . . .. . . . . Jaeropsis
23. Uropods tiny; first pair of antennae very tiny (fig. 60,A);
second pair without a scale . . . . . . . . . . . . . . . .Jaera
23. Umpods and first antennae well developed (fig. 60,D,E); second
antennae with a scale . . . . . . . . . . . . . . . . Janiropsis
Suborder Oniscoidea

. . . . . 25
. . . . . . . . . . . . . 28
. . . . . . . . . . . . . . 26
. . . . . . . . . . . . . . 27

24. Living just above water on the shore of ocean or bay

24. Truly terrestrial or near fresh water
25. Slow-moving; under rocks or debris; not actually entering
water; antennae and uropods short
25. Rapidly moving; living more in the open and entering water at
times; antennae and uropods long

ISOPona

aa

Fig. 60.-Jaeridae
( a f t e r Riohardson) ; A ,
Fabrioiuei B , aerouais lobata Riohardson; C , antennule
(small) und anienna of eame~ D. 1, Janirnpttis oalifomioa
Richardson.

26. Iarger; found only just above high-tide line under debris;
with characteristic large white spots
Alloniscus perconvexus
26. Small; found in other mgist locations; with uniform color or
at least not conspicuously mottled
29
27. ksal segment of uropods at least twice as long as wide (fig.
61,B)
L i g i a occidental i s
27. Basal segment of uropods about as wide as long (fig. 61,A)
.Ligiapallasii

..
.............
....................
........................

Fig. 61.-&&
(= L i ~ i d a ) (after Richardson):
A , L i i a (Ligia)
iBrandtr B ,
(l(ggllig~ooidentalie
Dam.

28. In or near streams, .generally under stones; flagellum of second

antenna multi-articulate; dorsal surface dark gray, glossy
Ligidium g r a c i l e
28. Not confined to streams; flagellum of second antenna with
three joints or less (fig. 53)
29

.......................

...............

92

LBBORAMRY AND FIELD TEX!t' IN INVER'l!EBRA'l!EZOOMGY

29. Abdomen abruptly narrower than thorax; brown or reddish; surface slightly granulated; usually under decaying vegetation
Porcellionides pruinosus
30
29. Abdomen not abruptly narrower than tho30. Body very convex, capable of being rolled into a ball; uropods
not extending beyond tip of abdomen. "The Pill Bug"
Armad illidium vulgare
30. Body more or less depressed, not capable of rolling into a
ball; uropods extending beyond tip of abdomen. "The Sow N s "

....................

...........

.....................

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 1
31. Body roughly granulate or tuberculate Porcellio scaber (fig. 53)

.
31. Body smooth or minutely granulate . . . . . . . . . . . . . . . 32
32. Body very smooth and shiny; posterolateral margin of first
four thoracic somites straight; terminal process of abdomen
(telson) pointed at tip, with a deep longitudinal dorsal
groove
.Porce1lio.formosus
32. Body smooth or slightly granular; posterolateral margins of
first four thoracic somites produced posteriorly; telson
rounded at tip and with but a faint dorsal groove
.Porcelliolaevis

..................

.......................
LIST OF ISOPODS

Suborder Chelifera (or regarded by some authorities aa a separate order, Tanaioidea )

M Pancolus californi ensi s Richardson
M Anatanais
M Tanais
Suborder Jalvifem (Idotheoidea)
*D M Idothea reotilinea Lockington
D D Pentidotea resecata Stimpson
D *M Pentidotea stenops Benedict
D M Pentidotea whitei Stimpson
*D *M Pent idotea wosnesenskii Brandt
Suborder Flabellifera (Oymothoidea)
Anthuridae
Cirolana harfordi Lockington
Cirolana 1inguifrons Richardson
Dynamene dilatata Richardson
Dynamenella glabra Richardson
Erosphaeroma amp1i cauda Stimpson
Exosphaeroma octonoum Richardson
hkosphaeroma oregonensi s Dana
Jkosphaeromu rhomburum Richardeon
Limnoria lignorum Rathke
Livoneca (clinging to gills of fish; not in key)
Sphaeroma pent odon Richardson

Suborder AseUota
M Jaera
Y Jaeropsis lobata Richardson
*M Jani ropsi s cal iforni ca Richardson
M hnna
Suborder gniscoidea
(NOTE:-Ligia and Alloniscus are almost as truly terrestrial as the others of this suborder, but live very near the
sea, just above the high tide marks, often descending into
the intertidal when the tide is out. )
TB M Alloniscus perconvexus Dana
*T Armdill idiunr vulgare Iatreille
*B D *M Ligia occidentalis Dana
*M Ligia pallasi i Brandt
T Ligidium gracile Dana
*T Porcellio fornosus Stuxberg
*T Porcellio laevis Iatreille
*T Porcellio scaber Iatreille
T Porcell i onides (~eto~onorthus)
prui nosus B d t
Order h ~ h i ~ o d a
The Amphipoda are divided into three suborders: Gammaridea (Plate

XI, no. 6), Hyperiidea (fig. 62), and Caprellidea (Plate XI, no. 5); the
first two with seven free thoracic segments, the third with six. The
Caprellidea are further distinguished by
a very narrow body and a rudimentary abdomen. The Hyperiidea which are commonly
parasitic and will not be seen by us are
further characterized by a very large head
(fig. 62) and large eyes. The Ganmaridea
are represented by the familiar "sand flean
of the seashore and a large number of species living on seaweed? under rocks, etc.
We shall confine our study to the GanrmaFig. BE. -Platyoelus, Suborder
ridea, but the student should be able to
Eyperiidoa, Order Amphipoda.
recognize caprellids
Using the dissecting microscope or low powers of the compound
microscope study fixed speoimens of Ganunarus and whole mounts of Gamnarus and Hyallela. Bdake out (fig. 63) antennules and antennae, known
also as first and second antennae. The elongate basal joints constitute
the peduncle, the small distal joints the fhellum. Note that there ie
a small secondary flagellum on the antem.de of some genera (fig. 71).
The mouth-parts will be studied later using mounts of the separated
appendages of Allorchestes. On the whole mounts m e the subchelate first
and second legs (2d and 3d thoracic appendages) known as first and w ond pmtho~ods,the five simple walking le known as perio~ods,the
three biramous pleopods with slender, mi?= tiarticulate rami and the
thee pairs of biramous leg-like ~ ~ O D O("springing
~ S
feetn).
The first two legs, called the first and second pathopods (fig. 63),

94

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

commonly terminate in expandea hands, and in one or both or neither the terminal segment (seventh segment or dactylus) closes on

antenna

Fig. 6 5 . - T a l i t m s lvatious Baewell, a gsm~nrrid

uphipod l a b e ~ h oharaoterlstio
o ~
8truotureS 8
Pleopoda 1-5, the appendages of t h e first three abdominal segments are omitted ( a f t e r shoemaker). S e p
ments of thorax and abdomen are numbered; 01, 0 2 s
f i r s t and second gnathopoda; pl-p5 = the nalking
legs or periopods; U1-U3 = the three posterior abdominal avvendages termed uropods; a - h e t h e s i d e
plates or 00x81 plates of the thoraoic appendages.

the expanded g& (pro~odugor sixth segment) as a claa~inafi-.

Such appendages are termed subchelate (fig. 64,B). The finger may form
with the sixth joint a forceps-like pinching claw, in which case the
appendage is chelate (fig. 64,A). Finally the finger may not close,
but remain stationary like a claw, as in the other legs, when the
gnathopod is said to be sim~le.
Note the flat extensioas of the chitinow bases of the thoracic
appendages which are known eg side dates or coxar plates (fig. 63).
Study mounts of the mouth-parts of Allorchestes. The most posterior appendages are the maxilli~eds(fig. 64,C) united by their two
basal segments. Each extends distally as a palp and bears internally
two endites or lobes born respectively as the outer and inner plats.
More anteriorly are two very runall appendages, the paired aecond and
first maxillae (figs. 64,X.F). In Allorchestes (in contrast to-the
condition in Ganunarus, fig. 64,E,F) the second maxilla is largest and
bears a tiny palp, while the first is d l and without a palp. The
mandibles are heavily ohitinized structures lacking a pal in Allorchestes (fig. 64,D) but having one in G a m n r s (fig. 64,~P.In front
of and behind the mandible are the fleshy upper and lower lips (fig.
64,H)
Sex dimorphiam is frequently marked and it is often necessary to
recognize the aex in order to identify. This mag often be done on the
basis of the $ze of the gnathopoda whioh are uaually larger in the
male. In sexually mature females the egg m s on the ventral surfaoe

of the abdomen enclosed in the marsupial plates (- otjstegites) is a

dietinguishing character.

Fig. 64.-Mouth parts of various amphlpods: A, f i r s t gnathopod o f Allomheates with

segments numbered, showing ohelate oondition; B , aeoond @thopod of same, subohelate;
C , maxilliped of Ganumlus ; D , mandible o f
loroheat s ( l a o n n g palp); E-(3, aeoond
maxilla, 1 s t maxinundible o f d % l e
wfth p l p ) ; 8, lower l l p of
hpitho'e', showing notohed fibnt lobes.

Using the key, identify the specimen given you and such others as
time allows. This will require a certain amount of dissection. Do this
with needles under the higher powers of the dissecting binocular. Mount
dissected parts under coverslips and study by means of the compound
microscope.
KEY TO C
! EE AMPHIPODS (Suborder Clammaridea)

. . . . . . . . . .. . . . .

1. Side plates 2-4 united, forming a large lateral shield Metopa sp.
2
1. Side plates 2-4 not united (fig. 63)
2. Uropod 3 (moqt posterior) vestigial, seemingly absent; species
Potiooerus spong i oolus (fig 65 )
living in aponge
3
2. Uropod 3 of at least moderate size, funational
3. Rostrum conspicuous, at least 1/3 ae long ae first joint of
antennule
4
3. Rostrum inconapiououa, lesa than 1/3 as long as first joint of
antemule
9

. . . . . . . ...
..........................
..........................

........

96

UBORAMRY AND FIELD TEXT IN 1NTWI"l'BRATE ZOOLOGY

4. Antennule with long accessory flagellum (fig. 66,A); legs with many
bristles, adapted for burrowing
Pontharpinia obtusidens (fig. 66)
4. Antennule without accessory flagellum; legs not adapted for burrowing
5

..,.............
............................

Fig. 65.-Podooerus
s p o w i c o l u s Alderman
( a f t e r b l d e r n n f i r s t and aeoond
gnathopoda of male; C-D, same of female.

antennule; By third periopod.

5. Integument indurated, bearing processes; mouth-parts prolonged,

Acanthonotozomatidae (fig. 67)
as if for piercing
5. Integument and mouth-parts not especially indurated
6

.......

.......

F i g . 67.-Pano~lea, family Acanthonotozomatidae ( a f t e r Shoemaker).

6. Antenna longer than antennule; telson cleft to base; abdominal

segments5and6fused
N o t o t r o i s t r i d e n s (fig. 68)
6. btennule longer than antenna; telson undivide& abdominal segments 5 and 6 not fused
7

......
....................

I
1

Fig. 68.-Nototro~ia t r i d e n s Alderman ( a f t e r

~lderman)t A, s i d e v i m j B , f i r s t gnathopod)
C , seoond j o i n t o f f i f t h periopod.

7. Body not laterally compressed but somewhat depressed

7.

. . . . . . . . . . . . . . . . . . Pleustes depressus (fig. 69 )

~ o d v l a t e r a l l ~ c o m ~ r e s s e.d .. . . . . . . . . . . . . . . . . 8

Fig. 69.-Pleuates d e ~ r e s s u a
Alderman ( a f t e r ~ l d e r m a n:
) A, s i d e
view; B , seoond gnathopod.

8. Flagellum of antennule 11-jointed, less than twice as long as

3
Neopleustes brevicornis
~edunclc
8. ~lkellumof antennule more than 11-jointed, more than twice as
Neopl eustes puget tensi s
long as peduncle
9. Body somewhat depressed (flattened dorsoventrally) ; usually living in tubes or in ascidian colonies, etc.
10
9. Body laterally compressed; not usually living in tubes
12
10. Flagellum of antenna with many joints
Ericthonius sp.
10. Flagellum of antenna with 3 or fewer joints (fig. 70)
11

...............
...........

&

:
!

.........
....
...... .....

98

LABORATORY AND FIELD TMT IN INVERTEBRATE ZOOLOGY

Fig. 7O.--Cor0 h i m aurvis in0.

+*o,.

......
. . . . . . . . . . . . . . . . . . . . . . Corophiunr insidiosum
12. Uropod 3 uniramous . . . . . . . . . . . . . . . . . . . . . . 13
12. Uropod 3 biramous, although one branch is sometimes very
small . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

Corophium bonellii
11. Flagellum of antenna with 3 joints
11. Flagellum of antenna absent, last joint of peduncle claw-like

Fig. 71.-,Uaroprotovu,us.

13. htennule with accessory flagellum; side plate 5 as. deep as 4

Mioroprotopus (fig. 71)
13. Antennule without accessory flagellum; side plate 5 much shallower than 4
14
14. htennule much shorter than peduncle of antenna
15
17
14. Antennule longer than peduncle of antenna
15. Gnathopod 1 subchelate in male; often on sand, but not burrowing in it
Orchestia traskiana (fig. 72,E)
15. Gnathopod 1 simple in male; burrowing in sand
16
Orchestoidea oaliforniana
16. Antenna longer than body
16. Antenna not more than half the length of body

....................
........................
. . . .. .. .. .. .. .. .. ..
...........
.........
.......
. . . . . . . . . . . . . . . . . . . . Orchestoidea cornioulata

(Perhaps intergrading varieties)

17. Inhabiting freshwater; size small; antennule, antem about
same length, flagellum of each with only 12 or fewer joints

. . . . . . . . . . . . . . . . . . . . Hyalella kniokerboohri
...........................

17. Marine; size larger, antennule and antenna generally with more
joints
18

AME'HIPODA
18. Bntennule short, l i t t l e more than half the length of antenna
19
18. AntenniilS longer, generally much more than half length of
21
antenna.........
19. Color black and gray with sometimes transverse white stripes;
s i z e f a i r l y large; found i n San Trancisco Bay
Hyalepugettensis
19. Color pink o r pale; size small
20
20. Color pink; found with red algae on ocean coast
Hyale rubm
20. Color pale whitish; San Francisco Bag shore Allorchestes angusta
21. Antenna with dense t u f t s of b r i s t l e s , b r i s t l e s being a t l e a s t as
long as width of flagellum; bay shore
A11 orchestes plumulosus
21. Antenna1 b r i s t l e s much fewer and shorter
22
22. Joints 4 and 6 of male gnathopod 2 i n contact ( f i g . 72,B)
Hyole sp. (fig. 72,A,B)
22. Joints 5 of male gnathopod 2 produced between joints 4 and 6
All orchestes sp.
(fig. 64,B)
(Character 22 is decisive i n distinguishing between the
genera Hyale and Allorchestesbut it is often d i f f i c u l t t c j
observe and consequently has not been used above where other
d i s t i n c t ions a r e possible )

..................

. . . . . . . . . . . . . . . .. .. .. .. .. .. .. .. . . . . . . . .
...

..
...........
....................
..................
.

Fig. 72.-A,B,

&
l a&
sp. ; C ,D, Aoroider californioa Alderman5 E, Orohertia tmskiana Stkpson, side plate 4.

23. Gnathopod 1 moh longer than 2, very long i n male

Aoroides cal ij'ornica (figs. 72,C,D)
(Aoroides columbiae reported common on rocky coast [see
Ricket t a and Calvin] )
23. Gnathopod 1 not larger than 2
24

..............

.................

100

LABORATORY AND FIELD TEXT I N INVERTEBRATE ZOOLOGY

24. Uropod 3 with prominent upward-turning hooks a t the end of the

outer ramus
25
30
24. Uropod 3 without such hooks
25. Antennule shorter than antenna; one hook t o each umpod
26
25. Antennule usually longer than antenna; two hooks t o each
uropod
27
26. Hind margin of joint 6 of male gnathopod 2 produced into
strong tooth; female gnathopod 2 much larger than 1
Jassa sp.
26. k l e gnathopod 2 without posterior teeth; female gnathopod 2
Ischyrocerus sp.
l i t t l e larger than 1
27. Palm of male gnathopod 2 transverse; flagellum of antennule
w i t h 3 5 o r m o r e joints
28
27. Palm of male gnathopod 2 oblique; flagellum of antennule
u s u a l l y w i t h l e s s t h a n 3 5 joints
29
28. Joint of 5 of male gnathopod 2 small; flagellum of antennule
with about 50 joints
Ampitho8 valida
28. Joint of 5 of male gnathopod 2 longer than joint 6; flagellum
with about 35 joints
Ampithog humeralis (Plate XI, no. 6 )
29. Flagellum of antennule twice a s long as peduncle; a prominent
Ampithob' vaillanti i
tooth defines palm of male gnathopod 2
29. Flagellum of antennule usually l e s s than twice as long as peduncle; a very small tooth defines palm
Ampithog rubricata
29. Flagellum of antennule l i t t l e longer than peduncle; a large
blunt tooth defines palm
Ampithod sinaulans
30. Antennule longer than antenna and possessing an accelssory f l a gellum
.31
30. b t e n n u l e not longer than antenna; accessory flagellum absent
except i n h'urystheus
35
31. Rami of uropod 3 very unequal, outer ramus much longer, o r at
least, much larger
32
31. Rami of uropod 3 about equal
34
32. Outer ramus of uropod 3 several times as long as the inner
ramus; eye round o r irregular, but usually not bean-shaped
Me1 ita cal ifornica (fig. 73)
(And other species of Melita. M. palmata reported abundant
by Ri cket t s and Calvin. )
32. Outer ramus of uropod 3 not more than 3 times as long a s inner
ramus i n our species, but the inner ramus is very much more
slender; eye bean-shaped
33
33. Inner ramus of uropod 3 at l e a s t 2/3 length of outer ramus
Gammarus pri bilofensis
33. Inner ramus of uropod 3 l e s s than 1/2 length of outer ramus
Gammarzls cory"srvico1us
34. Accessory flagellum of antennule 2-jointed; regular flagellum
Elasmopus ( f i g 74)
longer than peduncle
34. Accessory *flagellum 7-10 jointed; flagellum shorter than peduncle.
.Maera inaequipes

.........................
..................
....
...........................

.............

...................
..............
.............
...
..

..

..........
..........................
....................
.....................
.................

.................

..................

.....................
.....................
...........
....................

Fig. 73.-Melita
o a l i f o r n i c a Alderman ( a f t e r Aldernun): A, s i d e view; B , f i r s t gnathopod; C , second
gnathopod.

Fig. 74.-Elasmopus.

35. Legs 1-5 all with fingers pointing backward; abdominal segments 5 and 6 coalesced
Polycheria antarctica
35. Some of legs 1-5 with fingers not pointing backward; abdominal
36
segments 5 and 6 not coalesced
36. Side plate 4 concave posteriorly; gnathopods rather small
37
36. Side plate 4 convex or straight posteriorly; gnathopod 2
usually powerful, at least in male
39
37. Telson entire, pointed (fig. 75,A)
Kinonem sp. (fig. 75,A.B)
37. Telson cleft or emarginate
38

.........
......,........

...

.............
. . . . .. .. . . . . . . . . . . . .

Fig. 75.-A,

Kinoneme, t e l s o n and uropod; B , second gnathopod o f same with s i d e p l a t e

and ~ U s t e g i t e ; C , emarginate t e l s o n o f J'onto~enia.

LBBORAMRY AND FIELD TEXT I N INVER-TE

ZOOLOGY

38. Telson deeply c l e f t (fig. 75,C); side plate 4 deeply excavate

behind
.Pontogenia inenis
38. Telson emarginate, notch only 1/5 length of telson (fig. 76,A);
s i d e plate 4 with shallow excavation 01 igochinus sp. (fig. 76)

..................
.

Fig. 76.-A, t e l son: B , seoond periopod w i t h s i d e

p l a t e ; C , f i r s t gnathopod.

.......

39. Rami of uropod 3 about equal

arystheus tenuioornis
39. Inner ramus of uropod 3 much smaller than outer
40

........

Fig. 77.-Photis
oonohfoola Alderman: A, s i d e p l a t e S and seoond segment
of f i r s t p e r i o p o d ~B , s i d e p l a t e 4; C , s i d e p l a t e 6 and thiTd periopod.

40. Male has side plates 1 and 2 much shallower than others; female
has s i d e plate 4 broader than front lobe of 5

..................

m o t i s oonohicola (fig. 77)

40. Male has side plates 1 and 2 not m c h shallower than others;
female has side plate 4 narrower than front lobe of 5

. . . . . . . . . . . . . . . . . . . . . . . . Photis
LIST OF AMPHIPODS

Suborder Gammaridea
M Acanthonot ozomat idae
Allorchestes angustus Ihna
*BB A1
1orchestes p l u m 1 osus S t impson
M All orchest es ap.
Bd Ampithod humeral is S t impson

reinhardi

Suborder Gammaridea (Continued)

M Ampithod rubricatahntagu

M Ampithog simulans Alderman

M Ampitho8 vaillantii kcas

B *M
M
M
*B
B
F M
M
*MM
M
*B
*MB
*M
*B
*F
M
M
*M

M
*M

*MB
BM
*M
M

B
M
*M
*B *M
*M
M

M
M
M
M
M
M
M

Ampitho8 valida Smith

Aoroides californica Alderman
Aoroides columbiae Walker
Corophium bonellii Milne-Edwards
Corophium insid i osum Crawford
Corophium spp.
.Elasmopus brasil iensi s Dana
Elasmopus r a p Costa
Ericthonius sp.
Zurystheus tenuicornis Holmes
Ganunarus oonfervi colus S t impson
Gammarus pri biloj'ensis Pearse
Hyale sp.
Hyale mbra Thompson
Hyale pugettensis Dana
Hyalella knickerbockeri Bate
Ischyrooerus sp.
Jassa sp.
Kinonemc
hiaera inaequipes Costa
Melita californica Alderman
Me1 ita palmatu Montagu
Melita sp.
Met.opa sp.
Microprotopus
fleopleustes brevicornis Sars
Neopleustss pugettensis Dana
Nototropis tridens Alderman
Oligochinus
Orchestia traskiam Stimpson
Orchestoidea californi ana Brandt
Orchestoidea cornioulata Stout
Photi s re i nhard i &,dyer
Pleustes depressus Alderman
Podocerus spongicolus Alderman
Polycheria antarctica 'Stebbing
Polycheria osborni C a k
Pontharpinia obtusidens Alderman
Pontogenia ineais h y e r

Suborder Camellidea

*B
*M
*MM

Caprella aequili bra Say

Caprella kennerlyi Stimpson
Caprella acutifrons L a t r e i l l e
Caprella scaura Templeton

LABORATORY AND FIELD TEXT IN INVXRTEBRATE ZOOLOGY

Subclass Eucarida
This subclass includes two orders, a small group of small pelagic
shrimps, Order Euphausiacea, and the larger Order Decapoda including the
true shrimps, crabs, and other higher crustacea. We shall confine our
consideration to the Decapoda and to three suborders under it: the
shrimps, Suborder Caridea; the crabs, Suborder Brachyura; and the hermit crabs and relatives, Suborder Ano~nura.
Order Decauoda
The caridoid fascies or macrurous type, best exemplified by a
shrimp or prawn (fig. 78) best represents the baxic plan of structure
of the decapods. It should be familiar from earlier studies of the
crayfish or lobster. Review it if not. This type of body structure is
termed macrurous in contrast to the brachyurous type of the true crab
and the anomurous type of the hermit crabs.
W e yourself familiar with the brachyuran type of structure by
identifying on a crab all the appendages and all other structures and
regions labeled in figures 80432.
Key out your specimen, using Schmitt, also the key below.
The typical anomurous body plan is that of the hermit crabs in
which the abdomen is reduced asymmetrically and many of its appendages
are vestigial or lacking. Among the species which other characters show
to belong to the Suborder Anowa, however, are some which are very
crablike in appearance (fig. 94) and which may be termed brachyurous
anoruurans, and others again (figs. 95 and 96) which have large symmetrical abdomens and may be termed macrurous anomurans.
Using the keys in Schmitt's Decapod Crustacea of California, identify further (1) a shrimp, (2) a hermit crab, and (3) a brachyurous
anomran.
Be ready for a fifteen-minute test on terminology at the end of the
laboratory period.

1. Abdomen used for swimming, with well-developed tail-fan, and

pleopods; abdomen in our species with sharp bend; shrimps
(fig. 78)
1. Abdomen generally not used for swimming, with tail-fan and pleopods more or less reduced, except the Callianassidae (37) which
can be distinguished from the true shrimps by their large unbent abdomen and their burrowing habits
2. Rostrum very short, dorsally flattened; eyes free; hands subchelate (fig. 83)
2. Rostrum very small or wanting; eyes covered by carapace; hands
chelate, one or both powerfully developed
2. Rostrum distinct, usually well developed and spinose (fig. 87);
eyes free; hands weakly chelate

...........................2

............9

.......................3
. . . . . . . . . . .4
. . . . . . . . . . . . . . . .5

F i g . 78.-A

maoruran (shrimp-like) dsoapod (from ~ohmltt).

LABORBTORY AND FIELD TEXT IN INVERTEBRI1TE ZOOLOGY

Fig. 79.-Typioal third nmxilliped

of a maoruran deoapod (from Sohmitt).

Fig. 80.-Ventral

view of anterior part of body of a braohyuran decapod (from ~ c h m i t t ) .

Flg. 81.-Dorsal
braohvuran
- ---*
.

----

view of

oarnm
np. a
r---*.""-h d n s

regions used i n descriptions.

(eiopodth)

Bmnchm\/

'-.:~~i-?:.y.5y~

.....,

li.

'"

.-.

.?

Mg. 82.-Typioal t h i r d
maxilliped of a braohyuran
(orablike) deoapod (from
Sohmitt).

3. Sixth segment of abdomen with a large, cimular dark spot on

either side; a n t e d scale nearly three times as long as
wide (fig. 83,A)
Crago nigrilnaculata
3. No such spot; body more or l e s s covered with minute dark spots;
tail entirely black; scale hardly twice aa long a s wide
(fig. 83,B)
Crago nigricauda

..............
..................

Fig. 8 3 . - x o ;

A , hand and aoale of 2. ni~rimaoulataLookljlgton; B, hand and eoale

of C. niarioauda Stimpson.

4. One chela greatly enlarged, complex with dactyl above (fig.

84 ) "The Pist 01Shrimp"
Crangon
4. Chelae about equal, simple; dactyl below
Betaeus

................
.........

Fig. 64.-Chela

of Crangon dentipes Querin ( a f t e r Sohmitt).

5. Iarge; both legs of f i r s t pair simple; seoond pair of legs very

unequal, both with multiarticulate carpi (30 segments)
Pandalus danae
5. Smaller; both lega of f i r s t pair chelate (chelae very small):
second pair equal or nearly so, carpus with seven ,&gmentb7
Spirontocaris (figs. 85-87) 6

.........................
..................

F i g . 86.-Spirontooaris

paludioola Holmee. (After Rioketts and ~ a l v i n . )

108

LABORATORY AND FIELD TEXT IN lNVERTEl3RATE ZOOLOGY

6. Rostrum elongate (figs. 85, 87,A), reaching beyond middle of

ant ennal scale.
6. Rostrum short (figs. 86, 87,B), not reaching middle of antenna1 scale
,

.......................
........................ .

Fig. 86.-Spirontooaris

b r e v i r o s t r i s Dana (from Guberlet)

7. Rostrum not reaching as f a r as cornea of eye

......................

Spirontocaris taylori
Spirontocaris palpator
7. Rostrum reaching cornea or beyond
8. Rostrum reaching beyond middle of antenna1 scale but not into
d i s t a l quarter; greenish, semitransparent, with oblique reddish bands on carapace and crimson bars on legs; common i n
tidepools
Spi rontooaris picta (fig. 87,C )
8. Rostrum longer, reaching t o or beyond end of antennal scale;
uniform green; c o w i n eelgrass and on mud f l a t s
Spirontocaris paludicola (figs. 85, 87,D)

.....

............

............

Fig. 87.-Rostra
of ommum species of Svirontocaris ( a f t e r
Stimpaon; D,
Holmesl B, g. brevirostris Dana; C , g.

oh mitt): A, g. oarinata
g. paludioola Holmes.

9. Antennae situated i n t e r n a l t o eye, with short flagellum (figs.

80, 90); abdomen always small, folded under thorax, with uropods lacking; l a s t pair of legs generally about the same a s
others
Brachwra 10
9. Antennae situated external t o the eye, with long flagellum
(fig. 88); abdomen variable, of ten asymmetrical (fig. 98)
or much reduced, but generally with uropods present (exceptions under 29); last pair of legs usually much different
from others, often smaller
Anomura 27

......................

.............

Fig. 88.-Eead

of Papuras arliforniensie (after

oh mitt).

10. Carapace generally elongated, drawn out anteriorly between

the eyes as a rostrum (figs. 89, 91). Spider Crabs
10. Carapace squazvt or transversely oval, with very short rostrum
ifany . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 5

. . . . 11

Fig. 89.-Loxorhynahue

orisvatus Stimpson (after Rathbun)

11. Carapace roughly triangular in shape, broadest behind; very

hairy, with much foreign matter attached to hairs; ram
above the low tide mark. b e bsked Crab
Loxorhynchus crispatus (fig. 89 )
11. Carapace not triangular; generally not w hairy as above;
12
found also intertidally
12. Carapace about as hroad as long (fig. 90)
Yimulus 401 iatus (fig. 90)
12. Carapace longer than broad
13

................
...................
. . . . . . . . . . . . . .. .. .. .. .. .. . . . . . . . . . . . .

Fig. 90.-Mimulua

foliatua Stimpson (after ~ c h m i t t )

110

LBBORATORY BND FIELD TEXT IN T

E

ZOOLOGY

13. Surface of carapace smooth; distance between eyes less than

1/3 the width of carapace. The Kelp Crab

................

Ptrgettia (= ~ p i a l t u s )producta

13. Surface of cara ce tuberculated or spiny; distance between

eyes about $greatest
width of carapace (fig. 91)

. . . . . 14

A
Fig. 91.-Pu&tia

B
(after Sahniitt)~A,

E.

gmaili13 Dhn8# B,

11.

riohii Dana.

14. Smaller; carapace not expanded posteriorly; anterolateral

tooth broad, anteriorly directed (fig. 91.8)

.......................

Plrgettia g r a o i l i s

14. Larger; carapace distinctly broader po~teriorly;anterolateral tooth narrow, laterally directed (fig. 91,B)

........................

figettiarichii

15. Commensals with mollusks, annelids, etc.; eyes reduced,

carapace often poorly oaloified. Pinnotherid Crabs
16
15. Free-living; eyes and carapace normal
18
16. Oarapace much wider than long
Pinnixa
16. Carapaae not much wider than long
17
17. Carapace with two longitudinal grooves leading back from
orbit..
J'abia
17. Carapace without such groovea
Pinnotheres
18. Body square, sides nearly parallel, anterior edge nearly
transverse; palp of external maxilliped does not articulate
e
m (segment basal to palp)
25
near the inner angle of the m
18. Body usually round or tranwersely oval, with sides not parallel and anterior edge of carapace rounded; palp of external
(third) maxilliped articulates at or near inner angle of

. . . . . . . . .. .. .. .. ..
. . .. .. .. .. .. .. .. .. .. .. .. .. . . .
........................
............
..

m e r u s . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 9

19. Carapace much broader than long, with front (between eyes)
narrow, cut into five teeth (fig. 81)
19. C~irapacenot much broader than long; front wide, without
teeth.
b .

. . . . . . . . . . . . 20
. . . . . . . . . . . . . . . . . . . . . . . . -23

'i
1
I

!
I

111

DECAPODA

20. Front produced, with f i v e nearly equal teeth; fingers with

dark color. The Red Crab
Gamer productu s
21
20. Front not produced, with f i v e unequal t e e t h
21. Hairy; with red spots on ventral surface; antennae a s long as
width of front. The Rock Crab
Canoer antennarius
22
21. Not very hairy; not spotted below; antennae very short
22. Very large, widest at tenth tooth l a t e r a l t o eye, which is the
Cancer magister
largest tooth. The Edible Crab
22. Small, widest at ninth tooth; t e e t h a l t e r n a t e l y longer and
shorter
Cancer jordani
23. Fingers of same color as r e s t of chela (introduced from the
Atlanti c; now i n $an Francisco Bay and Lake Merrit t )
Rhithropanopeus harrisii
24
23. Fingers much darker than r e s t of chela; found on ocean coast
24. Hand of cheliped with a tooth on inner side of upper margin
?;ophopanopeus heathi
Lophopanopeus bellus (fig. 92)
24. Hand without tooth

. . . . . . . . .. .. .. . . . . . . .
........

....

........
.....................

....................

......................
........

7-

Fig. 92 .-Lophomnopeua

bellus Stimpson (from Guberlet)

25. Surface of carapace with numerous transverse lines.

Pachygrapsus crassipes
The Lined Shore Crab.
25. Carapace without transverse l i n e s
26
26. Color generally dark red o r purple; spots on chelae; no h a i r
Hemigrapsus w d u s
on legs. The Purple Shore Crab
26. Color generally greenish; no spots on chelae; legs hairy.
The Mud Crab.
Hemigrapsus oregonensis
27. Abdomen short, crablike, reflexed under thorax
28
27. Abdomen elongate, not reflexed (fig. 95)
36
28. F i r s t p a i r of legs chelate; carapace crablike (fig. 94)
29
28. F i r s t p a i r of legs simple, carapace subcylindrical with wings
which cover legs; second t o fourth legs with l a s t joint
curved and flattened. The Sand Crab
merits analoga
29. Fifth legs very small, folded under carapace, seemingly absent;
30
uropods lacking
29. Fifth legs reduced but held over the back; uropods present
32

..........
...............

.......

.............

........

...........
....

......
.......................
..

112

LBBORAWRY AND FIELD 'l%XT IN INVERTEBRATE ZOOLOGY

30. Carapace wider than long, covering legs; abdomen small,

flattened
Crypt01 ithodes si tohensis
30. Carapace as long as, o r longer than, broad; abdomen thick and
fleishy
31
31. Legs and carapace hairy; carapace as long ~ E broad
J
Hapalogaster
31. Isgs and carapace rough; carapace longer than broad
Oedignuthus inermis
32. Body and chelae thick; chelae unequal, tuberculate; legs
h a i r y . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3
32. Body and chelae t h i n (fig. 94); chelae equal and smooth; l e u
nude
34
33. Chelipeds granulated and tuberculate above; telson of 5 large
Pachyoheles rud i s
plates (fig. 93,A)
33. Chelipeds not tuberculate, granules hidden by short, thick
pubescence which f i l l s gap between fingers and extends almost
t o end of fingers- telson with 5 large and 2 very small
plates (fig. 9 3 , ~ )
Pachycheles pbescens

..............
...........................

......................

............................
..............

...........

34. Carpus of chelipeds about once and a half as long as wide;

s i d e s converging toward d i s t a l end; palp of t h i r d maxilliped
Petrol i sthes ci nct ipes
orange
34. Carpus longer, side^ parallel; outer edge of palp of t h i r d
maxillipedblue.
.35
35. Carpus rough; cheliped a s o l i d color on upper surface; proximal inner b e of finger of cheliped blue
Petrol isthes eriomerus (fig. 94)

.................
.....................

................

Fig. 94.-Petrolisthee

eriomerue Stkpson (Pram Quberlet).

35. Carpus r e l a t i v e l y smooth; a median row of blue dots on hand of

cheliped; proximal inner base of finger of cheliped orange
.Petrolisthes manimaculis
36.
36.

...................
Abdomen symmetrical ( f i g . 95 ), extended; burrowing i n mud
f l a t s . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Abdomen asymmetrical, s o f t and twisted (fig. 98), protected by
a s n a i l s h e l l . The Hermit Crabs . . . . . . . . . . . . . . . 38

Fig. 95.-Uporebia

37. F i r s t pair of
simple. The
37. F i r s t pair of
pair small,
Shrimp

(from ~ u b e r l e t )

legs subequal ( f i g . 95), subchelate, others

Upogebia pugettensis
Mud Shrimp
l e g s very unequal (fig. 96 ), chelate; second
chelate; f i f t h p a i r subchelate. The Ghost
Call ianassa cal iyorni ens is

..........

...............

Fig. 96.-Callianassa

(from ~ u b e r l e t )

38. Dactyls of walking legs with longitudinal s t r i p e s ( i n l i f e the

s t r i p e s a r e red, the ground color of the dactyl deep blue; i n
preserved specimens t h e s t r i p e s a r e dark on a l i g h t orange
39
background)
38. Dactyls of walking l e g s without longitudinal s t r i p e s ; aolor
not blue i n l i f e
40

.........................
......................

LABORATORY AND FIELD TEXT IN IIWEFFEBRATE ZOOLOGY

114

39. Antennae red i n l i f e : d i s t a l end of oron-"

-" walking legs
IUUi Ul
w i -th blue band; c
largin o
n with three subequal
in(ientat i cms, the l a t e r a,1 ones with m i nute, closely packed
tetath
, Pagurus sum
mae greenish brown i n l i f e , wlta yellow markings Ion some
guents; d i s t a l end of propodi with white band; d i s ,t a l mar.
1 of telson with f i v e indentations, the extreme l a ,t e r a l
.
3s small and without minute teeth; median indentat! nn 11s hirsu

a"-

....

..

TT

& Dana

F i g . 97.-&

carapac:e obscur e i by

40. Dactyls of walkine:

(Prom Gub e r l e t )

-I-..
LL.
znrougnout;
a n t e r i o r marain
.=-hairs;
Inedian
rowth
o:
heaw g
) s t r a l ) project ion short, blunt o r b:
rounded (fig.
,A)
rgums 5rranosimanus
40. u l s t a l t h i r d 01 aactyls whlte; a n t e r i o r margin of caral
,
icured k~y h a i r s, its r
;e
project:
.g. 98,E I )
.illi

Lars

U.L

-1-same c o
lor

..

......

...

rletaeus harfordi (Kingsle:

M Crago nigricauda (Stimpsol11

Crag o nigromaculata (Lock:ington )

M Cran:gon dent ipes (Guerin)

M ilrnn,
-,-.- gon cal iforniensis (Hc

.-

~ a n dalus
; danae Stimpson
Spi rontocaris brevirostri:r Dana
M.- Spirt
ontocaris palpator (bven )
ont ocar;is paludicola (Holmes(
ont ocariis picta (Stimpson)
ont ocariis taylori (stimpson)

LllBORllMRY AND FIELD TEXT

*B

TB M
DM
D *M

M
M
Ed
*D *M
*B TB D M

**MM
*M
D *M
D *M

*M
TI3 M

*B D M
M
B TI3 D M
*B M
TB M
B D *M
*B TB M
D*M

M
M
DM
TI3 M
M
DM
*I
TB D *Id

*L

IN IIWEBTEBRATE ZOOLOGY

Anomura
Callianassa californiensis Dana
Crypt01ithodes sitchensis Brandt
Emerita analoga (Stimpson)
Hapalogaster cavicauda Stimpson
Oed ignathus inermis (stimpson)
Pachycheles pubescens Holmes
Pachycheles rud is Stimpson
Pagurvs hirsutiusculus (Dana)
Pagurus granosimanus (stimpson)
Pagurus hemphi 11i (Benedict)
Pagurus same1is (Stimpson)
Petrolisthes cinctipes (Randall)
Petrolisthes eriomeris Stimpson
Petroli sthes manimaculatus Glassell
Upogebia pugettensis (~ana)
Brachvura
Cancer antennarius Stimpson
Cancer jordani Rathbun
Cancer productus Randall
Cancer magister Dana
Fabia subquadrata Dana)
Xemigrapsus nudus [Dana)
Hemigrapsus oregonensis (Dana)
Lophopanopeus bellus (Stimpson)
Lophopnaopeus heathi i Rathbun
Loxorhynohus crispatus Stimpson
Yimulus fol iatus Stimpson
Pinnixa faba (Dana)and other spp.
Pinnotheres concharum (Rathbun)
agettia gracilis Dana
Pugettia richii Dana
Pugettia (Epialtus) producta (Randall)
Rhithropanopeus harrisi i (Gould)
THE TERRESTRIAL ARTBROPODS

Most of the arthropods except the Cmstacea are adapted to terrestrial life, although some of them have returned to aquatic habits.
It is customary to group these under three subphyla (or nlnsses)
the Myriapoda, the Insecta, and the Arachnida, although the first
group, ths Myriapoda, is almost certainly polyphyletic. In addition
the groups Pycnogonida and Tardigrada, while placed by some in the
Arachnida, are better considered as of doubtful taxonomic relationships.
This enormous assemblage of animals can have but brief and fragmentary consideration here since our emphasis in these studies is
largely on the aquatic habitats. Our excursions into the terrestrial
domain are usually confined to one field trip to a near-by canyon

where varied environmental types including stream bed, stream banks

(riparian), tangled thickets, open forested hillsides, and grassy hilltop furnish a rich and varied fauna largely of mollusks and arthropods.
The student should recognize all of the types of terrestrial arthropods encountered and place them in the food cycles of the communities
studied. No attempt will be m d e to identify them further except that
the insects should be placed at least to the order.
Groups commonly encountered are the garden centipedes (Class
Sgmphyla), the true centipedes (Class Chilopoda), and the millipedes
(Class Diplopoda) (all of the subphylum Mpiapoda), the insects (Subphylum Insecta), and of the Subphylum Arachnids, the Scorpions (Class
Scorpionida), and the spiders (Class Araneida), the mites and ticks
(Class Acarina), the harvestment (Class Phalangida), and occesionally
)
the book scorpions (Class ~seudoscorpionida
The food habits of the terrestrial arthropods are extremely varied.
Certain groups, however, have relatively unifo m habits Thus the Symphyla and Diplopoda are plant feeders, while the Chilopoda and practically all Brachnida are predators.
The Insecta run the gamut as to food habits. Many are predatory
carnivors, others are parasites, while a vast number are plant feeders
and serve as key industry animals in important food chains. Such, for
example, are the aphids.

SUBPHPUTM'MYRIAJ?ODA
Class SYWEYIA, The Garden Centipedes
See mounts of Scutigerella. These are progoneate as are the Diplopoda but differ from them in many important features. They possess many
structural indications of relationship to the primitive stock from which
the insects have arisen. Note that there are only 12 leg-bearing somites.
See the peculiar coxal sacs at the bases of the legs below. There is but
one pair of tracheae opening at the bases of the antennae.
These tiny white wormlike creatures will be found in loam and among
decaying leaves. They feed on vegetation and cause damage of considerable importance to some crops.
Class DIPMPODA, The Millipedes
Note the rounded body of many segments (except in Polydesmidae).
The first few segments bear a single pair of appendages each, others
bear two grid each is really two fused segments. The appendages of the
seventh free segment of the male are modified for copulation, the
reproductive ducts opening near the anterior end.
The types commonly encountered are the relatively large dark-brown
cylindrical millipedes of the genus Tylobolus and the flattened species
of the family Polydesmidae. See special paper by Ellsworth for more
definite identification.
Class CHILOPODA, The Centipedes
Note the depressed body of many segments, varying in size, each
bearing a pair of appendages. Appendages of first postcephalic segment

LABOUO
!'l RY AND FIELD 'I'EXT IN INVERTEBRATE ZOOLOGY

118

modified as poison jaws (maxillipeds). Those of last pair are tactile.

Reproductive apertures (not easily made out) open on last true segment.
Most commonly encountered is a stout, flattened, reddish species,
Otocryptops sexspi nosus Say, an inhabitant of decaying wood, characterized by having 23 leg-bearing segments, and two prominent spines on the
first segment of the anal legs. For other species see special paper by
Gressitt

Time does not permit any adequate study of this enormoue group,
many of whose species are among the most important in terrestrial communities. An elementary knowledge is assumed. The student should know
at least the orders and where possible abundant species should be identified. For those who wish to attempt identification by means of keys,
Lutz's @adbook of American Insects is the most available, although its
examples are largelg Eastern species. For specific Western species,
Essig, Jnsects of Western North America, is recommended, although the
absence of keys, inevitable in such a work, makes identification more
difficult.
For aquatic insects reference may be had to A Guide to the St*
of Fresh-Water Biolws, by Needham and Needham (Comstock Co., 1930)
and to various other works listed under that headin@; in the bibliographs*
SWl?lRWM ARACBNIDA
Class SCOFQIONIDA
See the segmented body made up of (1)the prosoma, covered by a
carapace and bearing the six pairs of appendages characteristic of
arachnids, the chelicerae, ~ e d i ~ a land
~ i four pairs of &; (2) the
mesosoma, consisting of seven broad somites, the first of which bears
the reproductive organs covered by the genital operculum (seen between
bases of last two pairs of legs), the second the pectines, and the
next four stimata opening into book-lungs; (3) the metasom, consisting of five segments, and the telson modified as a caudal sting. Mesosoma and metasoma together constitute the o~isthosoma

Class BRBNEIDA, The Spiders

The body consists of cephalothorax and umegmented abdomen. See
the eight simple eyes. Identify the eight pairs of appendages. The
chelicerae are poison .jaws. See the modified terminal segment of the
serving to transfer sperm. Locate the openings to the
E
d (just posterior) the genital aperture. Near the tip of
the abdomen below see the three pairs of s~innerets.
For a general treatment see Comstock, The S~iderBook. For a key
to the local genera see special paper by hkehler.
Class &XlXA, Mites and Ticks
See the fused body without segmentation or subdivision. W e out
the appendages characteristic of the arachnids.

Mites are the only arachnids occurring also in marine and freshwater habitats.
Class P3AL$NGU]A, The Harvestmen or Daddy Longlegs
These are spider-like, but differ from the spiders in their segmented abdomen, chelate chelicerae, and number of eyes (2). One species
is frequently seen under pieces of wood or under leaves in the Berkeley
Hills. Because of lack of information concerning our local species, no
list ie given here.
The PYCNOGONlDA
The pycnogonids are a group of arthropods usually placed in the

Arachnids, but of decidedly doubtful relationship to that group. Several

species are common in the intertidal zone of ocean and bay. For the
morphology of this class see Borradaile and Potts, Parker and Baswell,
or another advanced text.

A KEP TO TBE FTCNOGONIDS OF C

! HE PACIFIC COAST OF NORTH
AbBRICA (ESPECIALLY OF CENTRAL CALIFORNIA*)
By Joel FV. Hedgpeth
This key is based on obvious characters without reference to their
value in indicating systematic relationships and is intended as an aid
to identification of pycnogonids in the field. The distribution of many
species is imperfectly known.
The characters used in this key should be visible under the medium
power of a dissecting microscope, although for many of the smaller
species it is difficult to make out details because of the debris adhering to the surface of the animal. This is particularly true of the spiny
f o m , and in some cases it will be necessary to brush the animal off
with a camel's hair brush, perhaps even to pick at it with a dissecting
needle. Specimens are best killed for study purposes by placing them in
fresh water for a few minutes, which causes them to die in an extended
condition. They are strongly thigmotropic, and should be kept separatelg
to prevent them from becoming inextricably tangled.
For terms used in the key see figure 99.

1. Proboscis one jointed (figs 99, 100)

..............

2
1. Proboscis two jointed; minute species with angular first coxal
joints; cheliforea and palpi present, chelifores chelate;
3-6 mm. long. m
a Beach, Santa Cruz Island
lRLrydyce spinosa Hilton

.....................

*For a complete key, including the species described i n Hilton's

preliminary papers, see Hedgpeth, J. W . , "A Key to the Pycnogonlds of
the Pacific Coast of North AmericaYnTrans. San Diego Soc. Nat. H i s t . ,
March, 1941.

LAKIRAMRY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

Fig. 100.-Pycnogonum

stearnsi Ives.

........
...............

3
2. Chelifores or palpi (fig. 99), or both, present
2. Chelifores and palpi lacking (fig. 100); stout-legged, knobby
form, oval in outline
Pyonogorurm
a. Body granular; dorsal tubercles inconspicuous or absent.
Littoral species, often on Gribrina or Metridiunr, and
Abietinaria. Body of female 6 mm. long, male smaller.
P. stearnsi Ives (fig. 100)
Alaska to San Diego
b. Body reticulated; prominent dorsal trunk tubercles which
are higher than eye tubercle; male 6-7 mm. long,
female smaller. Subtidal, wharf pilings and anemones.
Monterey Bay, Tomales Point
P. rickettsi Schmitt
3. Chelifores or palpi present, the other lacking
18
4
3. Both ohelifores and palpi present
4. Chelifores rudimentary or achelate in adult
6
4. Chelifores chelate in adult
5
5. Scape of ohelifore shorCer than proboscis; small; body compact;
male 5 mm. long, female 11.8 am. long. Central California
coast
Amothea ohelata Hilton
5. Soape longer than proboscis
Nymphon
a. Fingers of chelae short, thick; legs stout, with scattered
spines; neck short; 50 mm. long. m e t Sound-Bering Sea,
cirounnpolar
N. grassipes (0. Fabr)
b. Fingers of ohelae slender; legs slender; neck long; 80 mm.
long. Puget Sound, Departure Bay, B.C., offshore Southern
California
1. pixellae Scott
6. Body compact, discifom; segmentation indistinct (fig. 102);
legs short, stout
7
6. Body more or less elongate (fig. 101); lateral processes separated; legs long and slender
a. Proboscis oylindrical; ohelifores one-jointed stumps;
body 18-20 mm. long. On Abiet inaria, k i n County to
Lecythorhynohus marginatus Cole (fig. 101)
Laguna Beach
b. Proboscis elliptical; ohelifores three-jointed; legs without large tarsal claws; 8-9 mm. long. San Pedro Point,
Ammothella bi-unguiculata
(IDS Angeles) to Iaguna Beach
7. Proboscis broad, clavate or bulbous, long as body, sometimes
directed ventrally and apparently shorter from above
10
7. Proboscis tapered to a point or oonaiderably Bhorter than
finystylum
body: mall compact circular forma
8. Proboscis tapered to a point
9
8. Proboscis rounded; body very compact; 7 mm. long. Pacifio
Grove
T. occidental is (cole)
9. Light-brown hobby form with white articulations; 4-6 nrm. long.
Usually on Abiet inaria, Marin Oounty to Southefn California
T. californicuni Hilton (fig. 102)

.....

....
. . . . . . . .. .. .. .. .. .. .. ..
. . . . . . . . .. .. .. .. .. .. .. .. .. ..

. . . . . . . . . .. .. .. .. .. .. .. .. . . . . . . . .
............

................
......................

....

........
.................
..................

...............

*!Phi8 i e the common central California coast epeciea which has been
confuaed with T. Sntemdium. I t i a a l i t t l e larger, with etouter
lega, than T . intermediq. See Hedgpeth (1940) for more detailed
distribution, under T. Intermedium.

122

LABORILTORY AND FIELD !l!EXT IN IIlVTZTEBRATE ZOOLOGY

9. Delicate pale form with sharp spinelike protuberances on

lateral processes; female more rounded. On Obel ia San

Diego to Camel

..............

w
Fig. 101.-Lecythorhynchue

marginatus Cole.

I).

intermedium Cole

Fig. 102.-Tanvstvlum
c a l i f ornicum Hilton.

. . . . . . . . . . . 13
. . . . . . . . . . . . . . . . . . . . . . . . . . 11

10. Neither body nor legs conspicuously spiny

10. Body or legs spiny (fig. 103) or with conspicuous spiny
tubercles

Fig. 103.-N~n1~hoveie svinoeus Hall.

. . . . . . . . . . . . . 12

11. Without tall spiny tubercles on body

11. Three tall spiny tubercles on body; two rows of spiny
tubercles on tibia1 joints: coxae smooth; 10 mm. long. Sub-

tidal,

La@;una

Beach to Tomales Point

. . . . . . . . . . . Nymphops i s spinos i ssimus (Hall)

12.

(fig 103)
Chelifores and abdomen hobby; legs spiny; body compact; chelifores two-jointed; 16.m. long. Bering Sea, Monterey Bay
Ammothea latifrons Cole
Chelifores and abdomen not knobby; lateral processes separated;
spines or slender tubercles on trunk; chelifores threejointed; 10 mm. long. San Diego to &rin County
Ammothella spinifera Cole
15
Chelifores less than half as long as the proboscis
14
Chelifores at least half as long as proboscis
Tubercles on dorsal posterior margins of first two or three
trunk sements; proboscis broad as body; chelifores threejointed; 6-9mm. long. In holdfasts and on sheltered rocks,
Bdarin County to Laguna Beach
Ammothella tuberculata Cole (fig. 104,A)
lithout such tubercles; proboscis narrower than body; chelifores two-jointed; 18-20 mm. long. Hydroids, Puget Sound
Ammothea disco idea Exline

....................

12.

...................

13.
13.
14.

......
.........

............

14.

...................

Fig. 104.-A,

Amnothella tuberoulata Cole; B, Amnothea nudiuscula Hall; C, Ammothea

gracilipes Cole; D, Halosoma virldintestinalis Cole

. . . . 16

15. Proboscis elliptical or tapered at tip (figs. 104,~,C)

15. Proboscis blunt or truncate; 30 mm. long. Pribiloff Island

. . . . . . . . . . . . . . . . . . Ammothea pribilofensis Cole

16. Protuberance on dorsum of first coxal joint half as long as
first coxal joint . . . . . . . . . . . . . . . . . . . . . . 17
16. Protuberance less than half as long as first coxal joint;
proboscis broadly elliptical; 7 mm. long. San Francisco
Bay
Ammothea nudiuscula Hall (fig 104,B)
17. Protuberance half as long as coxal joint; body 13 mm. long.
Anunothea alaskens i s Cole
Alaska.
17. Protuberance t-hree-fourthsas long as coml joint; proboscis
narrowly elliptical; 8 mm. long. San Francisco, krin
County, Vancouver, B.C.
A. gracilipes Cole (fig. 104,C)

...........
................
....

124

LABORATORY AND FIELD T;MT IN INVERTEBRATE ZOOLOGY

18. Chelifores present, chelate (fig 105), usually overreaching

19
proboscis; palpi lacking
18. Chelifores lacking; palpi present; elongate; distinctly segmented; proboscis large, dilated; eye tubercle tall; 50 mm.
long. Offshore water, Southern California
Colossende is cal ifornica Hedgpeth
19. Chelae and proboscis not extended forward on a neck; chelae
small..
20
19. Chelae strong, heavy, flanking the rather blunt proboscis at
Call ipall ens
the end of a well-developed neck
a. Body rather compact; segmentation of last two trunk
segments indistinct; 9-10 mm. long. Littoral,
Laguna Beach
C. cal iforniensi s Hall
b. Body extended; lateral processes separated; body distinctly segmented. Offshore waters, Southern California
C. pacif'i ca Hedgpeth
20. First trunk segment not conspicuously overhanging proboscis
(fig. 105)
21

..................

...............

..........................
.........
...........

...............
.........................

Fig. 105.-Phoxicilidium
femoratum Rathke.

20. First trunk segment overhanging forward so that proboscis

appears ventral in origin
Anoplodactylus
a. Lateral processes separate at base; eyes present; 8-9
mm. long. Littoral, Monterey to Sen Diego, also
offshore waters
A. ersctus Cole
b. Lateral processes contiguous at base; eyes lacking;
15-16 mm. long. Littoral, Iaguna Beach
A. cal iforni ens i s Hall
21. Body extended, lateral processes separated; 8-10 mm. long.
Alaska to Laguna Beach; circumpolar
Phoxichil idiun fenoraturn (Rathke) (fig. 105)
21. Body compact; small delicate form with vivid green intestine;
7.5 m. long. krin County to La@;una Beach
Halosomu viridintestinulis Cole (fig. 104,~)

............

............

.................

..........

PHYLUM MOUUSCA

This great phylum includes many of the important animals of the

littoral marine and the freshwater and terrestrial faunas to be studied.
In the intertidal zone it ranks with the arthropods as a dominant
structural type.
Five classes, each with a very distinctive structure, constitute
the phylum:
1. Class Amphineura, the chitons
2. Class Gastropoda, the snails, whelks, limpets, slugs,
nudibranchs, etc.
3. Class Scaphopoda, the tooth shells
4. Claas Pelecypoda, the clams, mussels, oysters, pectens,
etc.
5. Claas Cephalopoda, the squids, octopi, and nautili.
We shall confine our attention to the animals of three of these
classes., the chitons, the gastropods, and the pelecypods. The tooth
shells are not found in our region. Of the cephalopods, the squid are
pelagic and only one species of octopus, Paroctopus appolyon ( = Polypus
hongkongensis ) is encountered along the central and northern California
coast and that only occasionally. For an interesting account of its
behavior and breeding habits see Ricketts and Calvin (p. 96-99).
The evolution of the various molluscan types has concerned chiefly
three plastic regions, the head, the foot, and the mantle (figs. 114,
115). The mantle is the body wall over the visceral region. This grows
out as a mantle fold (figs. 113,B and 114) enclosing a mantle cavity
or mantle chamber within which lie the gills, the anal opening and the
openings of reproductive and excretory ducts
The mantle secretes the shell and that typical molluscan structure
varies with the shape, size, position, and habit of growth of the mantle. The mantle cavity, primitively posterior, has taken on various
positions in the various molluscan types.
The characteristic feeding organ of the mollusk, known as the
radula, is a band of chitinous teeth (fig. 106) produced in the ventral
cavity of the pharynx from which it is extended by muscular action and
uaed to scrape away the food. It may be seen in use by watching freshwater snails feeding on the walls of an aquarium. The radula, which is
found in all classes of Molluaca except the Pelecypoda, is most commonly
uaed in feeding on large leafy plants or large algae (thus in some chitons and many Gastropoda) or for scraping off encrusting layers of
minute unicellular organisms (many chitons, the limpets, some snails).
In the Cephalopoda and some snails it has been adapted to predatory
habits.
The Pelecypoda are plankton feeders sifting the tiny organisms from
the water currents caused to flow through the large mantle chambers by
the action of cilia on the large, complicated gills. The radula, having
no function in such a habit of life, has been lost in the Pelecypoda along
with the sense organs of the head, and, indeed, the head itself since,
covered by the mantle folds and the shell, it has no direct contact with
the environment.

126

LABORATORY AND FIELD TEXT IN INVERTEJ3RATE ZOOLOGY

Class AMPHINEURB

The Aplacophora, shell-less forms, often considered to belong to

the Class Amphineura, are largely deep sea in distribution and will not
be encountered by us. The chitons, Order Polyplacophora, are important
and interesting members of the intertidal faunas and we should be
familiar with all the known species. Unknown species should be preserved for study by the specialist. They may be killed in extended
form by allowing them first to flatten out on glass slides to which
they may be tightly tied before fixation.

Fig. 106.-flnall

portions of the radulae of various Mollusoa (from various sources).

The characters used in the identification of chitons have chiefly

to do with the 8-valved shell (figs. 107-111) the width and ornamentation of the muscular integumental area between the shell and the margin,
known as the girdle (fig. 107)) and with the proportions of the body.
Each valve consists typically of two layers, a basal and more extensive articulomentum which extends under the other plates anteriorly
and is more or less covered by the girdle laterally, and the upper
mentum, covering only the exposed portion of the valve. In Crypotochiton
the valves are entirely covered by the girdle and the tegmentum is
lacking. The valves overlap posteriorly and the portion of a valve
underlying the one in front is often also designated the articulomentum
and that portion covered by the girdle, the insertion plate.
k i n e the girdle. Note its width in proportion to the width of
the exposed portions of the valves (cf figs. 110 and 111). Note also
its ornamentation (fig. 107). Is it smooth, rough and leathery, covered
with granules, or with scales, or with fine hairs, or spines, or long
stiff hairs? .
Note the proportion of width to length. Is it half as wide as long,
etc.?

'.-

u-

Dissect out the valves of your specimen. Place one of them under
the microscope to see the tiny pits which form patterns on the surface
of the tegmentum. In these pits lie characteristic sense organs known
as esthetes. Egamine the posterior margin of the last valve. The slit
or notch in it is known as a sinus.

coarse
hairs

Fig. 107.-Diagramnatio

figure of a ohiton i l l u s t r a t i n g some of the wtious typ_es

of g i r d l e ornamentation.

Using the key below identify your specimen. When Pratt is available identify another species to the family. See the figures in Tryon
and Pillsbry, A k u a l of Concholo~.
1. Girdle completely covering valves, thick, red, gritty; large
(6-8 inches)
"The Giant Chiton," Cryptochiton stelleri
1. Girdle two to three times as wide as exposed portion of valves;
"The Black Chiton,l1 Katharina tunicata
girdle smooth and black
2
1. Exposed portion of valves at least as wide as girdle
2. Valves marked with conspicuoua zigzag lines of color
"The Lined Chiton," Lepidochitona (~onicella)1 ineata
2. No such lines
. 3
3. Girdle (Observe under the lens!) covered with hairs (figs.
107,108).
4
3. Girdle covered with scales like a coat of mail (figs. 107, 109)
Ischnochiton7
3. Girdle with minute granules (fig. 107), minute scales or minute
papillae
.lo
4. Hairs coarse and stiff
5
4. Hairs lighter, flexible
6
5. h l in shape; girdle coarsely mossy, narrowed at ends; valves
often corroded, with dorsal ridge, ornamented with nodules in
lines
"The Mossy Chiton,'l Mopal i a muscosa
5. Nearly circular; girdle much elongated in front, with scattering
hairs; valves short and broad, without nodules
" m e Veiled Chiton," Plaoiphorella velata

.....

C,
}

......

......

.........................
..........................
.........................
..........................
.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..

...........

............

128

LABORBTOFR AND FIELD !CEXT IN INVERTEBRATE ZOOLOGY

6. Girdle conspicuously c l e f t posteriorly ( f i g 107); valves sculptured

i n concentric l i n e s (often covered with red deposit); h a i r s slender,
very pale, e a s i l y l o s t
"The Hairy Chiton," Mopal ia oil iata
6. Girdle not conspicuously c l e f t ; valves smooth, gray, green, o r
blue, with brown l i n e s slanting from apex of valves; h a i r s
thick a t base, slender and incurved d i s t a l l y , each r i s i n g i n
a l i g h t spot
Mopal ia lignosa ( f i g . 108)

...

............

Fig.

108.-Momlia

lianosa

7. Central areas of body valves s e t with raised r i b l e t s ; l a t e r a l

areas of body valves, usually a l l of head valve, and posterior
8
region of t a i l valve s e t with radiating rows of nodules
7. No nodules v i s i b l e t o the naked eye
9

....
...............

Fig. 109.-Ischnoohiton:

A , I. mertensii (Middendorf) 8 B. I. .cooperi Carpenter. x 1 1/2.

8. 'Red t o d u l l brown; nodules and r i b s of valves and scales of

g i r d l e conspicuous
"The Red Chiton," Isohnochiton mertensii (fig. lO9,A)
8. Gray; nodules, ribs, and m a l e s l e s s conspicuous
Ischnoohiton cooperi (fig. 109,B)
(These a r e probably v a r i e t i e s of the same species.)

.....
................

9. Body elongated, considerably more than twice a s long a s wide;

valves obscurely sculptured; l i g h t gray i n color
"The Gray Chiton," Ischnochi ton magdal enensis
9. Body about twice a s long as wide; valves apparently smooth;
with an i n t r i c a t e pattern of white s p o t s on a mottled bluegreen background
Ischnochiton regularis
10. Girdle with scale-like h a i r s (figs. 107. 110); g i r d l e broad;
valves nearly as long as wide
Nuttallina californica ( f i g . 110)
10. Girdle narrow, ornamented otherwise; valves short' and broad
(fig. 1 l l )
11

.........

............

...............
.........................

Fig. 110.-Nuttallina

californica ( ~ e e v e )

Fig. Ill-Lepidochitona
mondi Pilabry. x 2.

11. Broad oval, low; d u l l o l i v e green; g i r d l e covered with closes e t granules; valves covered with f i n e granulations and
l a r g e r ( .04 mm. ) wart -like granules on l a t e r a l areas
Lepidochitona hartwegi i
11. Narrower; darker, some almost black; usually small i n s i z e
(1/4 t o 1/2 inch); g i r d l e leathery, minutely papillose;
valves covered with very f i n e granulations and s l i g h t l y
Lepidochi tona raymondi ( f i g 111)
beaked

....................

............

LIST OF CHITONS
C l a s s AlbmmmJRA
Order Polyplacophora
Cryptochiton stelleri (Middendorff )
Ischnochiton cooperi (Carpenter)
Ischnochi ton ncrgdal enens i s (Hinds )
Ischnochi ton mertensi i (Middendorff )
Ischnoohi ton regularis (Carpenter)
Katharina tunicata (food)
Lepidochitona hartwegi i (Carpenter )
Lepidochitona (l'onicell a) 1 ineata (Wood)

130

LAEiORllTORY AND FIELD TEXT I N INVERTEBRATE ZOOLOGY

M Lepidochi tona raymondi Filsbry

D B "M Mopal ia cil iata (Sowerby)
M Mopal i a 1 ignosa (Gould)
D B *M Mopal ia m s c o s a (Gould)
D *M Nuttal ina cal ifornica (Reeve)
D M Placiphorella velata Carpenter
class GASTROPODA
These animals are the dominant and conspicuous animals over much
of the area which we are to study, and the number of common genera and
species which we must be able to recognize in the field is large. Bey
present themselves, naturally, therefore, as a group for careful study
of methods of identification. Although revisions are badly needed the
gastropods of the Pacific Coast have been extensively studied and
numerous valuable works of references are available. Unfortunately
none of these gives keys which are available for the fauna of this

Fig. 112.-A

marine snail, the whelk (~uooinum)(aPter ~uohsbaum).

coast, and we shall have to depend for identification upon the key
given below. Identification may be checked with the excellent illusby Tryon and Pilsbry,
trations in (1) Oldroyd, (2) Iknual of Concholo~?;g:
and (3) 'Packard. When still in doubt as to the correctness of your
identifications, reference may be made to authentic named collections
in charge of the teaching assistant.
There are five main groups or types of gastropods which are given
various names and are variously grouped by different authors. They are
(1) the Aspidobranchia (often called Scuti branchia or Di otocardia)
including the limpets of various kinds, the abalones, and a number of
typical snail-like forms of which Calliostoma is our best example;
(2) the Pect ini branchia (often called Ctenobranchia and ~onotocardia)
including the vast majority of marine shelled gastropods; (3) the
Tect i branchia, including the sea hares (no common Moss Ebach examples);
(4) the Nudi branchia, the nudibranchs; and (5) the filmonata, including
most land and fresh-water snails, and slugs, but with only two local
marine representatives, Gadinia and Arctonchis.
On the basis of the twisted condition of the nervous system the
first two of the groups above are often placed together in an order

GASTROPODA

131

(or subclass), the Streptoneura or Prosobranchia, and commonly spoken

of as prosobmchs or streptoneurous gastropods. The remaining three
groups are also often included in a single order (or subclass), the
Euthyneura, and spoken of as euthyneurous gastropods, those whose
nervous system is symmetrical. The tectibranchs and nudibranchs are
often included in the single order, Opisthobranchia, as in Hegner.

mantle c a v i t y ( opened)
espiratory aperture (cut)

mantle f o l d (turned back)

lood v e s s e l s
A
B
Fig. 1 1 3 . - g ,
a t e r r e s t r i a l pulmonate ( a f t e r ~ e l e n k a ) :A, w i t h s h e l l i n place;
B , s h e l l removedj mantle chamber ("lung") opened by c u t t i n g mantle f o l d a n t e r i o r l y and
t u r n i n g it back.

On the specimen given you, Helix mpersa, a pulmonate (fig. 113).

identify the foot, the head, bearing the tentacles, the fold of bodywall known as the mantle, and the mantle cavib, the marginal thickening of the mantle called the collar which in some species (Ilyanassa)
canal,
is drawn out to form the si~hon,lying, in life, in the ,si~honal
an extension of the shell. By means of forceps introduced through the
apex

oper'culum

proboscis

\mouth
Fig. 114.-Calliostoma,

a scutibranch

( a f t e r Eales).

Fig. 115 .-A

pectinebranch ( a f t e r Eales)

mouth pull out the characteristic feeding organ of the Molluscs, the
radula. M n e it under the microscope. See demonstration mount of
radula of a limpet.

132

UBORATORY AND FIELD TEXT IN I m T E B R A T E ZOOLOGY

The shell of a typical pectinibranch consists of the body whorl

(fig. 115), which opens by the aperture, and the whorls of the spire
ending in the apex. The groove between two successive whorls is known
as the suture. The axis of the shell around which the shell is coiled
is known as the colwnella (see sectioned shell); the hollow center
sometimes present in the oolumella is the umbilicus (fig. 122) and a
thickening partially or completely filling the opening of the umbilicus below as the callus.

canal

Fig. 116.-A

d e x t r a l l y c o i l e d gastropod s h e l l , Buccinum M m o r c h i a n u m F i s h e r
( a f t e r Oldroyd).

Here as in the bivalves identification depends upon characters of

the shell. To gain familiarity with certain important characters and
at the same time learn the names of certain species to be encountered
later, make out the following characters on the demonstration specimens :
Canal: (1) lacking (Tegula), (2) open (Thais), (3) closed (firpura).
Aperture: (1) smooth (Littorina), (2) crenulate (~ritonalia).
Inner lip: (1) toothed (Thais larnellosa), (2) corrugated (~earlesia),
(3) smooth (~ittorina).
Sutures distinct: (Calli ostom costaturn)
Shell: (1) low-spired (~omlo~orna),(2) moderately high-spired (Calliostorna), (3) acute spire (~~itoniurn
or Bittiurn), (4)with
spiral ridges (Calliostorna ), (5) with radiating ridges (Epitoniurn), (6)with slits (~al
iotis ), (7) with under-shelf
(Crepidula).
Using the key below, identify as many specimens as time allows.
Check some at least of your identifications with Mrs. Oldroyd's work,
or if Acmea, with Mrs. Grant's paper. Learn the names of as many as
possible of the genera found at Moss Beach.

XEY TO
MARINE SEEL;L;ED GASTROPODS
(See a l s o Vokes, 1934, f o r small species)

.........

A. S h e l l with more o r l e s s elevated, coiled s p i r e

B. Shell with flattened spire, not r i s i n g above l e v e l of surface

o r only s l i g h t l y so and with a row of perforat ions near l e f t

46
margin
C. Shell without a coiled spire, cap-shaped ( p l a t e s XII-XV)
25
D. I r r e g u b r l y twisted tube masses
Vermetidae

............................
....
............
1. Aperture rounded, e n t i r e ; no indication of a canal (figs. 121,
122) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
1. Aperture produced, forming a canal o r rudiment thereof ( f i g .
ll6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Canal rudimentary; s p i r e g r e a t l y elevated; 5-7 f l a t s p i r a l
threads on each whorl . . . . . . . . . . . Bittium ( f i g . 117,A)
2. Canal obvious . . . . . . . . . . . . . . . . . . . . . . . . . 3

Fig. 117.-A,

Bittium eschrichtii (~iddendorf); B, Nassarius (= ~lectrion)

3. Canal short, d i s t a l end of canal marked by a decided notch

(fig. 117,B)
3. Canal elongated, with o r without notch

4.

......................... 4
............. 9
Notch conspicuous; a deep s p i r a l furrow i n body whorl t o l e f t
of canal ( f i g . 117,B) . . . . . . . . . Nassarius (= Alectrion)
Notch inconspicuous; without deep furrow . . . . . . . . . . . . 5

4.
5. S h e l l with smooth, polished, evenly convex surface; body whorl
making up more than 3/4 of s h e l l ; aperture g r e a t l y elongated,
with a broad, conspicuous white c a l l u s on the columella and a
broad shallow notch a t base of aperture (canal)
Olivellabiplicata
5.Notso.........
6
7
6. Upper l i m i t of aperture a t o r above middle of s h e l l
8
6. Upper l i m i t of aperture well below middle of s h e l l

......................
....................
......
.......

134

LABORATORY BND FIELD TEXT I N INV'ERTEBRATE ZOOLOGY

7. Sculpturing of conspicuous, uniform, s p i r a l ridges separated by

narrow, deep furrows ( f i g . 118); body whorl with about 10
s p i r a l s ; upper end of aperture near middle of s h e l l
!Thais canaliculata
7. Sculpturing (often obsolete o r obscure) of low, unequal,
s p i r a l , rounded r i b s , nodulated by growth l i n e s , separated
by wide, shallow depressions; upper end of aperture well
above middle of s h e l l
m a i s ewrginata ( f i g . 119,A)

.......................
......

Fig. 118 .-Thais canaliculata (Duo

8. Shell with inconspicuous longitudinal grooves; f a i n t (obsolete )

furrow on body whorl; color dark; mud-flat dweller
Ilyanassa

Fig. 119.-A, Thais emar inata Deshaysl B , T . lima ( ~ a r t p ) ;perhaps only a v a r i e t y

of T. emarg-Amphissa
colmbiana Dall. ( ~ 1 1from ~ u b e r l e t . )

8. Shell with smooth polished surface; s p i r e approximately half

a s long a s body whorl; s h e l l with 7 whorls; upper end of
aperture more or l e s s c l e a r l y angular
Afitrella (= Colunbella)

9.

.
Shell with three heavy flanges extending e n t i r e length of
s h e l l ; canal usually closed . . . . . . . . . . . . . . Pllrpura
No such flanges; canal usually open . . . . . . . . . . . . . . 10

9.
10. Edge of outer l i p bearing a prominent spine
10.Nosuchspine

. . . . . . Acanthina
. . . . . . . . . . . . . . . . . . . . . . . . . 11

11. Sculpturing of numerous, small, obliquely longitudinal ridges;

Amphissa ( f i g . 119,C)
l i p d i s t i n c t l y thickened near canal
11. Sculpturing absent, or, i f present, of l e s s numerous, larger,
12
and not oblique ridges
12. Inner margin of outer l i p corrugated transversely along i t s
whole length ( f i g . 120,A)
13
12. Inner margin of outer l i p smooth, o r soulptured with a few
14
discreteteeth
13. Corrugations of l i p extend f a r into aperture; s h e l l large
Searlesia ( f i g . 120,A)
13. Corrugations only suggested; s h e l l sculptured with f a i n t , s p i r a l
threads undulated by prominent, radiating ridges; s h e l l
"The Oyster Borer," Urosalpim
smaller

...
...................

..................
.......................
.....................

.............

Fig. 120.-A,

Searlesia

dira ( ~ e e v e ) )B ,

Thais lamellosa (Gmelin). ( ~ 0 t hfrom

~ u b e r l q

14. Shell large; outer l i p variable i n thiclmess; sculpturing of

oonspicuous, radiating lamellat ions, f l u t e d by widely separated s p i r a l r i b s (sculpture often obsolete ) ; aperture f l a r ing at top, pinched a t bottom; s p i r e about equal t o body
Thais lamellosa (fig. 120,B)
whorl
14. Shell small; outer l i p usually thin, s p i r e g r e a t l y elevated;
soulptwing of conspicuous, close-set s p i r a l r i b s , e i t h e r
without radiating sculpture, o r with rounded r a d i a l l y arranged
swellings-Tri tonal i a ( a d i f f i c u l t group of several species );
forspeciessee
47

...............

.......................
15. Spire g r e a t l y elevated; sculpture of 12-14 sharp-cut, radiating
ridges . . . . . . . . . . . . . . . . . . Epitonium ( f i g . 121)
15.Spire relativelylow . . . . . . . . . . . . . . . . . . . . . 1 6

LABORATORY AND FIELD !ED3 IN l2WBlEBRATE ZOOLOGY

136

..............
........................
.........
.......................

19
16. With pearly luster or white inside
16. lithout pearly luster inside; inner side of outer lip brown
17
or purplish.
17. Callus broad and flattened; umbilirms closed
18
17. Callus thin and thread-like; urnbilious open (fig. 122); shell
Lacunaspp.
tiny

Fig. 121 .-Epitonium.

Fig. 122 .-Lacum.

Much
enlarged, s h o q open
umbilicus.

Fig. 123.-Tegula

Punebralis

ad am^.

18. Larger; shell of three whorls; broad band of white cutting

diagonally across the face of the lip to its edge
Littorina planaxis
18. Smaller; higher in proportion to width, with four whorler in
Littorina scutulata
shell; no white band on lip
19. With small nodes or teeth on the columellar callus (fig.
. - 123)
20
19. Aperture smooth, without teeth or nodes on the callus
22
20. Color black; outer lip overlapping the whorl above (fig. 123)
Tegulafinebralis
20. Color brown, tan or gray; outer lip merely touching the whorl
21
above, not overlapping it
21. Umbilicus alosed; base rounded; whorls convex; shell turbanPepla bmnnea
like (more pyramidal in young)
21. Umbilicus open; base flat; whorls flat3 body whorl m a r ;
shell pyramidal
Tegula inontereyi
22. Umbilicus closed
23
22. Umbilicus open
2243
22a.Very large; whit iah
Polzjnices leloisii (fig. 124)
22a. Small; reddish
Hargarites spp.
23. Shell ivory to reddish-brown; large, over 3/8 of an inch in
diameter; ribs spiral, heavy, well-marked; apire moderately
high, acute
24
23. Shell reddish; small, less than 3/8 of an inch in diameter;
ribs spiral, fine, not prominent; spire low, broad; (msembling a young Tegula)
Homalopoma

.......................
........
.

.....

.......................
..................
.........
................
. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .........
. . .. .. .. .. .. .. .. .. .. . . . . . .

.........................
................

24. Body whorl rounded; base conTlex; sut

color reddish, with blue ti.ngesat; apex
ru-Cii)

..............

ULU

re or 1ess distinct;

Call
i ostoma costatu;
-

/Pin

125,A,

24. Body whorl acutely angular; 1lase f l a1t; sutu.res ind

color old ivory; shape p punidal

...

' a l l i o s t ona cana1 i c u l a turn (fi

Fig. 124.-

3s l e w i s i i (Gould:) (from ~ u b e r l e t r) A , s h e l l , considerably

reduced; B, sand I:ollar containing eggs.

25. Shell nb c
25. 1Shell pt

ucrluracau

UI,

apca

)ex (keg

a( a fLet Oldrc

C. oanalicu:
-

26. With a shelf om under side

26. Without; guch a. shelf
27. Shell b ~ a cor~aarK Drown, peaKea; apex recurves; common on
l'egula
Crep i dula adunca
27. Shell white; peak reduced or absent; under stones and in
crevices; variable in shape and proportions Crepidula rulmmaria

...
....................
.
I

>

138

LABORBTORY AND FIELD TEXT IN I N P E R ~ T EZOOLOGY

..
..............
........................
......
...
.................
.......................

28. S h e l l thick and very irregular; t n i c a l l y suboircular

Hipponit
29
28. Shell t h i n andmore o r l e s s regular
29. With no d i s t i n c t external o r i n t e r n a l color markings ( P l a t e
XIII, no. 8 )
30
35
29. With a d i s t i n c t color pattern, a t l e a s t i n t e r n a l l y
33
30. UsuaIly over 3/8 of an inch long; apex a t o r near center
31
30. Less than 1/4 of an inch long
31. Usually on Tegula
32
31. Usually on narrow e e l grass; brown; b i l a t e r a l l y compressed
Acmaea paleacea (Plate XII, no. 6; P l a t e XIV, no. 9 )
32. Black; broad oval; sides convex; on Tegula funebralis
Acmaea asmi (Plate XII, no. 1; P l a t e XIV, no. 1 )
32. Light i n color, usually covered by coralline; s i d e s p a r a l l e l ;
rare; on Tegula 6runnea (occasionally on coralline-covered
rocks). Acmaea t r i a n g u l a r i s (Plate XII, no. 7; P l a t e XIV, no. 4 )
34
33. Vhite inside and out
33. Dark brown inside and out; corneous; usually on Egregia
Acmaea insessa (Plate XII, no. 2; P l a t e XV, no. 1 3 )
34. Outer surface r a d i a l l y ribbed; s h e l l f l a t ; outline c i r c u l a r
Gadinia
34. Outer surface not ribbed; s h e l l high; outline nearly c i r c u l a r
Acnaea mitra (Plate X I I , no. 10; P l a t e
XIII, no. 8; P l a t e XIV, nos. 7 and 8 )
35. Apex l e s s than one-third of length from nearest ( a n t e r i o r ) end
(Plate XIV, no. 1 7 )
36
35. Apex a t l e a s t one-third of length from end
39
36. Anterior (short ) surface concave (Plate XIV, no. 17 ) unless
corroded by fungus
37
38
36. Anterior surface s t r a i g h t o r convex
37. Ribs broad and undulating, branching; margin crenate; i n t e r i o r
generally dark, with l i g h t e r owl -shaped a p i c a l marking; very
large and heavy; lower; apex very near a n t e r i o r end
L o t t i a gigantea
37. Ribs rounded, unbranched, prominent t o obsolete (Plate XIV, nos.
11-19); margin ruff led; inner surface with elongated, s o l i d ,
dark-brown area on white background; border dark (Plate 1111,
nos. 21, 22); higher; apex seemingly at l e a s t f a r t h e r from
a n t e r i o r end.
Acmaea d i g i t a l i s (Plate X I I , nos.
23, 32; P l a t e XIV, nos. 11-14, 16-19)
(See P l a t e XIV, nos. 23-27, f o r hybrids with pel t a )
38. Heavily ribbed, with projecting marginal points (Plate XII, no. 8 );
inner surface with i r r e g u l a r dark markings on porcelanous g r m d
(Plate ZIII, nos. 7, 11-13); r i b s often spinulescent ( P l a t e XII,
no. 2rQ)
Acmaea scabra (Plates X I I , X I I I , P l a t e XIV, nos. 20-22)
38. Ribs fine, thread-like (Plate XII, no. 9 ) ; posterior surface
markedly arched convexly (Plate 7 3 , nos. 18, 19); i n t e r i o r
milk-white, with plain, broad, dark border, and apical spot
Acmaea persona
(Plate EII, no. 23).

......
........

.....................

......
............................
.............

.....................

..........

.....................

..............

........................

........

.....

..............

S h e l l s of v a r i o u s species
b e t i c a l l y by species :

1, A . asmi
20,30, A. cona
23,32, A. d i g i t a l i s
13,15, A. f e n e s t r a t a c r l b r a r i a
2, A. i n e s s a
18, A . i n s t a b i l i s

a in
-

d o r s a l view. Natural s i z e . Listed alpha-

LL,LG ,25,28,31, A. l i m a t u l a
1 0 , A. m i t r a
1 9 , A . ochracea
6, A . paleacea
3,26, A. p e l t a
4,9,17. A . persona

5 , A. rosacea
8,16,24, A. scabra
21,22,29, A. scutum
1 6 , A. t r i a n g u l a r i s

140

lX3ORBTORY AND FIELD TEXT I N INVERTEBMTE ZOOLOGY

39. Height a t apex l e s s than one-third of width (Plate XV, nos. 10,
40
16)
42
39. Efeight a t apex at l e a s t one-third of width

40.

.............................
..........
Width d i s t i n c t l y more than three-fourths of the length of s h e l l
(Plate XII, nos. 29, 31); s h e l l very low . . . . . . . . . . 41

40. Width about three-fourths of length of s h e l l ; s t r i a t i o n s

straight, unbranched, continuous from apex t o margin, with
interspaces equal i n width t o s t r i a t i o n s (Plate X I I , 19);
external color pinkish o r dark, spotted with white; i n t e r i o r
blue and brown; s h e l l very f r a g i l e
Acmaea ochracea (Plate XII, no. 19; Plate
XIII, no. 2; Plate XIV, nos. 28, 29)
41. Shell with thread-like r i b s (Plate XII, no. 14), with interspaces
wider than r i b s ( r i b s often obsolete ); external coloration plain
dark, or dark and white i n varying patterns (Plate X I I , nos.
21, 22, 29); internal color bluish-white with brown markings
(Plate PII, nos. 17, 18); large; common
Acmuea scutum (Plate XV, nos. 10-12)
41. Shell with imbricated r i b s alternating with 2-4 r i b l e t 8 (Plate
XII, nos. 25, 31); margin f i n e l y s e r r a t e (Plate XV, nos. 14,
15); rare
Acmaea limatula (Plate XIII, nos. 9, 10, 14, 15)
42. Sides compressed, making e l l i p t i c a l outline (Plate XII, no. 18 );
bottom rocker-like; external surface grooved; color brown,
often marked by white spot a t apex; inner surface white with
pale brown spot a t apex, and narrow brown border; rare; on
ami in aria ~ n d e r s o n i i Acmaea i n s t a b i l i s (Plate XII. no. 18: Plate XIII, no. 28)
42. Sides not compressed (except i n young-of A. pelts on seaweed);
43
outline subcircular t o ovoid
43. Outer surface with large, rough ribs, unless corroded; dense
white within; usually with narrow, scalloped, brown border;
often with brown blotch behind apex; large; common; very
Acmaea pelta (Plate X I I , no. 26; Plate XIII,
variable
nos. 20, 27, 29-31; Plate XV, nos. 1-6)
(See Plate XIV, nos. 23-27 f o r hybrids with d i g i t a l i s )
43. Outer surface corroded and satiny, or very f i n e l y s t r i a t e ; inner
surface suffused with brown, and highly nacreous; r a r e
Acmaea fenestrata (Plate X I I , nos. 13, 15;
Plate XIII, nos. 4, 6; P l a t e XV, nos. 7-9)
44. Perforation one-third a s long a s shell, with a l a t e r a l pro jection of the s h e l l modifying otherwise perfectly oval slit;
s h e l l l e s s than 2 cm. long, low, narrow; i n l i f e , s h e l l
nearly hidden by mantle protruding from perforation; mantle
surface a t .least 4 times a s great a s s h e l l surface
Megatebennus spp.
44. Perforation l e s s than one-twelfth a s long as shell; mature
45
s h e l l more than 25 mm. long; apex elevated

...........

..............

...

...

................

.....

...........

.......................
.........

Inside v i m of s h e l l s of various s p e c i e s of Acmaea. Natural s i z e . Listed alphab e t i c a l l y by species:

28, A. i n s t a b i l i s
23, A. persona
26, A.
9,10,14,15, A. l i m t u l a
25, A. rosacea
3 , A.
21,22,24, A. d i g i t a l i ~
8 , A. mitra
7 , l l - 1 3 , A. scabra
16-18, A. scutum
4 . 6 , A. fenestrata
1 , 2 , A. ochraoea
19, A. ins20,27,29-31, A. pelta

PLATE XI1

Shells of various species and one hybrid of t h e genus Acmaea seen i n l a t e r a l

view. Natural s i z e . Listed alphabetically by s p e c i e s :
1,

A.

5,6,

gg&

A. cons

3.10-14,16-19, A. d i g i t a l i s
7 , 8 , A.
28,29, A. ochracsa

9 . A. paleacea
23-27, A. Delta x A. d i p i t e l i s
2 , A. rosacea
15,20-22, A. scabra
4 , 11. triangularis

PLATE W

" ~ - 37

s of various species of Acmaea i n lateral view:

1-6, A. pelta
7-9, A. fenestrata cribrs
10-12. A. scutum

13, A. insessa
14-17, A. lirrmtula
18-19, A. persona

144

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

45. Perforation below apex, extending down anterior slope of shell,

wide anteriorly, narrowed posteriorly; shell white, heavily
Puncturella spp.
ribbed, with ribs projecting beyond margin
45. Perforation at apex, eliminating peak, subcircular in shape,
truncated posteriorly; shell gray or white, with darker gray
Diadora
radiating color streaks, margin crenulate
46. External color red
Haliotis rufescens
46. External color black
Haliotis cracherodii
47. Height of spire equal to or greater than height of body
whorl..
48
47. Height of spire distinctly less than that of body whorl
49
48. Sculpturing of spiral ribs crossed by longitudinal ribs of
equal size and prominence, producing effect of basket
Tritonulia interftossa
weave
48. Sculpturing of flattened spiral ribs without pr~minentlongiTritonulia gracillima
tudinal cross ribs
49. Sculpturing of regular, flattened, spiral ribs not markedly
interrupted by longitudinal growth lines; grooves shallow;
Tri tonal ia luri da
shell regular in outline
49. Sculpturing of elevated, rounded, spiral ribs, undulated by
prominent, rounded, revolving longitudinal swellings;
grooves deep and sharp cut; outline irregular

..

.......
..............
............

..........................
....

..................
...........

..........

. . . . . . . . . . . . . . . . . . . . . Tritonalia circumtexta
KEX TO THF: MORE COMMON NUDIBRANCHS

1. lith a circlet of branchiae on posterior dorsal surface (fig.

126,A)
1. lithout a circlet of branchiae
2. Branchial plumes retractile into cavity
2. Branchial plumes not retractile
3. Dorsal surface tuberculate, with scattered dark spots
3. Dorsal surface tuberculate, without scattered dark spots
3. Dorsal surface velvety, with dark rings (fig. 126,A)

............................2
. . . . . . . . . . . . . . . . . 12
............ 3
................ 8
. . . . . 45
....

. . . . . . . . . . . . . . . . . . . . . . Diaulula sandiegensis
4. Color lemon-yellow, with patches of black tubercles
. . . . . . . . . . . . . . . . . . . . Archidoris montereyensis
4. Color bright orange, with scattered dark spots and patches
. . . . . . . . . . . . . . . . . Anisodoris nobilis (fig. 126,B)
5. Small red or reddish brown species . . . . . . . . . . . . . . . 6
5. Large yellow and white species . . . . . . . . . . . . . . . . . 7
6. lho conspicuous black spots, one in front of branchia, and one
behind sense organs . . . . . . . . . . . . . . A1 di sa sanguinea
6. No conspicuous black spots, but with minute red' or brown papillae; often found on red sponges

........................

Rostanga pulchra

h
&

7. Yellow with white-tipped, scattered papillae; rhinophores

dark; branchiae whitish yellow
Dendrodorisfulva
7. Pale yellowish white with dark rhinophores and a row of lemonyellow spots on low tubercles on each side from rhinophore
Cad1 i na flavomaculata
t o branchial plume
7. Pale with numerous tubercles each tipped with 1emon:yellow; a
band of lemon-yellow outlining foot and mantle ( f i g . 126,C)
Cadlinarnarginata

.......................
...........

.......................

Fig. 126.- C h o n nudibranchs (from ~uberlet): A, Dialula sandieaensis (cooper);

B, Anisodoiis nobilis (Mao~arland)~
C, Cadlina marginaka MscParland; D, Hermissc

orass ieornis (~schsoholtz)

. . . . . . . . . . . . Polycera at
....................... Y
9. Body brown with many small bluish-white spots dorsally
. . . . . . . . . . . . . . . . . . . . . . . . Triopha maculata
9. Body pink or white . . . . . . . . . . . . . . . . . . . . . . 10
. . . . . . . . . . Hopkinsia rosacea
10. Body e n t i r e l y pink .
. . . . . . . . . . . . . . . . . ..ll
10. Ground color white .
t
d . . . . . .Lailacockerelli
11. Papillae long, club-like, r e~ d -ippe
11. Papillae short, red . . . . . . . . . . . . . . Triopha carpenteri
12. Without cerata o r dorsal g i l ~ ---tie
~ ,
dark red with f i n e white
longitudinal l i n e s . . . . . . . . Pleurophyllidia californica
12. With g i l l s o r c e r a t a (fig. 126,D) . . . . . . . . . . . . . . . 13
13. Without cerata; with ~ l u m e - l i k eg i l l s on ridges along e i t h e r
. . . . . . . . . . Tritonia festiva
side of back . . .
. . . . . . . . . . . . . . . . . . 14
13. With c e r a t a ( f i g . 12E
8. Body s t r i p e d longitudinally
8.Bodynotstriped

1".

14. Cerata b r i l l i a n t l y colored; body with b r i l l i a n t l y colored

areas
Herrni ssenda crass icorni s ( f i g 126,D )
15
14. Cerata and body without b r i l l i a n t coloring

..........

..........

146

LABORAMRY AND FIELD TEXT IN IITVXRTEBRATE ZOOLOGY

15. Cerata very numerous; feeding on sea anemones

. . . . . . . . . . . . . . . . . . . . . . . Aeolidia papillosa

15. Cerata larger, less numerous; feeding on %bularia

. .Alderia sp.

A KEY TO THE COMdON Gl3NERA OF FFUSJ3IATER SNAILS

.........................
........................
..........................

1. Shell dextral
2
1.Shellsinistral
4
2. Shell spired
3
2. Shell planispiral; very small; flat or slightly depressed above,
deeply umbilicate below; aperture somewhat expanded

. . . . . . . . . . . . . . . . . . . . Planorbula (= planorbis)
3. Iarge; operculate . . . . . . . . . . . . . . . . . . . Viviparus
3. Small; not operculate . . . . . . . . . . . . Fossaria (= ~ynuaaea)
4. Spired; spire acute or depressed; aperture contracted above,
rounded below . . . . . . . . . . . . . . . . Physella (= Physa)
4. Planispiral; base funicular; aperture suddenly expanded and
thickened; fewwhorls . . . . . . . . . . . . . . . . .Helisoma
IIEPTOm~LANDPuLMONATES
(For figures see paper by Nicholson)

A. Shell minute:
1. Elongate:
1'. Length 2$ mm.; dark brown to black; dull

....................

2 ' . Length 6 mm. ; yellowish-brown; glossy

Vertigocalifornica

. . Cochl icopa lubrica

1. Depressed:
1'. Widely umbilicated; 3 whorls; diameter 1.5 mm.; ash-gray
to white in color
Striatura milium
2'. Moderately umbilicated; 4 whorls; diameter 2 mm.; dark
brown (horn) in color
finctum conspectum
3 ' . Slightly umbilicated; 4i whorls; diameter 2 mm. ; light
horn to white in color
Pristiloma nicholsoni
B. Shell 6-8 mm. in diameter; depressed; moderately, but deeply,
umbilicated; 5 whorls; light to dark brown in color;
Polygyraloricata
oblique
C. Shell 14-18 mm. in diameter; depressed; moderately umbilicated;
6 whorls; dark to yellowish-brown; hirsute; some with parietal
tooth . . . . . . . . . . . . . . . . . . . Polygyracolumbiana
D. Shell 20-30 mm. in diameter; depressed; widely umbilicated; 5
whorls; greenish-yellow in color . . . Haplotrema vancouversnsis

.............
..........
.......

...................

E. Shell 25-40 mm. in diameter; depressed.

1. Imperforate (no umbilicus); 4 whorls; globose; yellowish to
russet; banded; diameter 25-40 mm.

........................

Helixaspersa
2. Minutely umbilicated; 5 whorls; pale horn color to ash-yellow;
superior chestnut narrow band; diameter 25-40 mm.
Helminthoglypta nickliniana

.................

3. Moderately umbilicated
1'.6$ whorls; brown on top, black beneath; body whorl
angulated; 35-38 mm.
Helminthoglypta infimata
2'. 7 whorls; light brown to yellowish in color with superior chestnut band; body whorl rounded; often dark
brown; 34-40 mm.
Helmi nthogl ypta arrosa

......

.........

FIELD KEY TO TBE MCAL LAND SLUGS

(For figures see paper by Waste, 1938)

1. Respiratory orifice anterior to middle of mantle

1.
2.
2.
3.
3.

. . . . . . . . . . . . . . . . . . . . . . Prophysaon andersoni
.......
...
...............
...
........
.......
..............
...
..................

Respiratory orifice posterior to middle of mantle

2
Tip of tail excavated, with a mucus pit
Ariolimax columbianus
Tip oftailconica1,not excavated
3
Yilax gagetes
Central oval area on mantle marked by a groove
Mantle without demarcated central area, but ornamented with
closely concentric grooves over whole surface
4
L i m maxims
4. Conspicuous irregular stripes on body
4. Body at most spotted, never striped
5
5. Color pattern a mosaic of black, gray, or buff-colored areas
L i m flaws
bounded by anastomosing grooves; slime yellow
5. Body coloration and reticulation of the integument not in correspondence; slime not yellow
6
6. Larger, adults about 40 mm. long; secreting abundant milky slime
when touched; color of back variable usually gray, or ocherous
buff with dark gray patches shading rarely to almost uniform
black
.Agriolima;l:agrestis
6. Smaller; hardly reaching a length of 30 mm., secreting very
Agriolimux campestris
little milky slime; color uniform

...................
....

LIST OF SPECIES OF LOCAL GASTROPODA EIICUJSIVE OF lEIE PULMONATES

Order Streptoneurn (Prosobranchiata)
Suborder &s~idobranchiata (Diotocardia )
D *M Acmaea asmi Middendorf
D *B *M Acmaea digitalisEschscholtz
TB D B M Acmaea fenestrata (Reeve)
D M Acmaea insessa (Hinds)

148

LABORATORY AND FIELD TEXT I N INVERTEBRATE ZOOLOGY

Suborder Amidobranchiata (Diotocardia) (Continued)

M Acmaea instabilis (Gould)
M Acmaea limatula %enter
D *M Acmaea mitra Eschscholtz
M Acmaea ochracea Dall
D M Acmaea paleaoea (Gould)
D *B M: Acmaea pelta lechscholtz
B M Acmaea persona Eschscholtz
D *B *M Acmaea scabra (Gould)
D B *M Acmaea scutum Eschscholtz
M Acmaea triangularis Carpenter
M Astraea inaequalis (brtyn)
D M Calli ostoma canaliculatum (&rtyn)
D *M Calliostoma costaturn (Bbartyn)
M Calliostoma tricolor Gabb
D M Diodora aspera (Eschscholtz)
D M Hal i otis cracherod i i Leach
D M Hal i ot is rufesosns Swainson
D *M Homalopoma oarpenteri (Pilsbry)
D A! Lott i a gigantea Sowerby
*Id Margari tes 1i rulatus (carpenter)
*M Margari tes pupillus (Gould)
M Margari tes salmoneus (Carpenter
M Margarites succincta (Carpenter
M Megateberms birnaoulatus (Dall)
D :M Tegula brunnea Philippi
D M Tegula funebral is Adams
M Tegula ligulata (Menke)
M Tegula montereyi Kiener
M Tegula pull igo (btyn)
Suborder Pectinibranobiata (Monotocardia )
D *M Acanthina spirata Blainville
M Aletes spamigems (Carpenter)
M A1vania hartmanae Vokes
D M Amphissa bicolor Dall
M Amph i ssa oolumb i ana Dall
M Amphissa versicolor Dall
D M Barleeia ha1 i ot iphila Carpenter
M Barleeia oldroydi Bartsch
D *M Bi ttium eschrichtii (Middendorff)
M Bivonia compacta Carpenter
TB M Cerithidea californica (Haldeman)
M Cingula californica Tryon
M Cinguia montereyensis Bartsch
M Clathrodrillia inoisa (Carpenter)
M Cows californicw Hinds
D *M Crepidula adunca Sowerby
B M Crepidula convexa glauoa Say
Id Crepidula lingulata Gould
D *M Crepidula wmmaria Gould

Suborder Pectinibranohiata (Monotocardia) (~ont

inued)
Cypraeolina pyriformi s (Carpenter)
Diala acuta Carpenter
Epi tonium i nd i anorum (Carpenter)
Epitonium tinctum (Carpenter)
Epit onium wroblewski i (Morch)
Hipponix antiquatus (Lim~eus
)
Hipponix tumens Carpenter
Ilyanassa (~asscrius)obsoleta (Say)
Lacuna marmorata Dall
Lacuna porrecta Carpenter
Lamellaria
Littorim planaxis Philippi
Littorina scutulata Gould
Marqinella ( ? )
M ~itFa
D *M Mi trella (= ~olurnbella)carinata (~inds
)
*M Mitrella gouldii (Carpenter)
* bfitrella tuberosa (carpenter)
*M Nassarius
mendicus Gould
M Nassarius perpinguis Hinds
M Odostomia oregonensis Dall snd Bartsoh
TI3 *Y 01ivella bipl icata (sowerby)
!b Phasi anella compta Gould
M Pseudomelato m torosa (Carpenter)
D*M hrpura foliata k t y n
D M Searlesia dira (Reeve)
M Spiroglyphus 1i tuellus brch
D*M Thais canaliculata (hclos)
D M Thais emurginata Deshayes
D *B*M Thais lamllosa (Gmelin)
Tritonali a ci rcumtexta Carpenter
D
*M Tritonalia foveolata Hinds
M Tritonalia gracillima Stearns
*I Tritonali a interfossa Carpenter
D *M Tritonalia lurida Eddendorff
M Tritonalia painei Dall
M Tritonalia subangulata Stearns
D M Trivia
*BM Turbonilla
Urosalpinx cinereus (Say)
M Velutina 1aevigatalVhrller
F Piviparus japonicus von Martens
F Vi viparus malleatus Reeve
Order Opisthobranchiata
Suborder Tectibranchiata
TB Actaeon punct ocaelata (Carpenter)
M Haminoea vesicula (Gould)
TB Phyllaplysia taylori Dall
B Tethys californica (cooper)

MBORATORY AND FIELD TEXT I N INVERTEBUTE ZOOLOGY

150

Class PELECYPODA
On a f r e s h o r preserved clam l o c a t e t h e hinge at t h e d o r s a l side,
t h e a e a t the v e n t r a l s i d e of t h e s h e l l , t h e s i ~ h o n s ,exhalent
( d o r 3 and inhalent, e i t h e r separate, a s i n M a c o w , o r u n i t e d t o
dorsal margin

resilium

cardinal hinge teeth

posterior lateral

muscle scar

muscle scar

anterior margin
pallial sinus

chondrophore

I
ventral margin
Fig. 127. The pelecypod shell, terminology (after Keen and Frizzell): A, oroas
section; B, right valve.

form a

"neck," a s

i n Mzja o r Schizothaerus. Remove one valve t o see t h e

t h e r e l a t i o n s of t h e siphon t o them. Within t h e mantle
chamber make out t h e g i l l s two p a i r s of l a b i a l m l u s , located on e i t h e r
s i d e of t h e mouth, t h e v i s c e r a l mass and foot, making up t h e body
proper, and t h e a n t e r i o r and ~ s t e r i o adductor
r
muscles running between
t h e two valves of t h e s h e l l .
The major d i v i s i o n s ( o r d e r s ) of t h e Pelecypoda a r e based i n p a r t
a t l e a s t on t h e s t r u c t u r e of t h e g i l l s . We s h a l l d e a l with two orders.
The Fil ibranchia a r e characterized by g i l l s i n t h e form of p a r a l l e l
filaments and by a h o l d f a s t , t y p i c a l l y of numerous threads, known a s a
byssus. Here belong t h e j i n g l e s (Pododesmus, f i g . 129,A), t h e marine
mussels (Mytilus and Modiolus) and t h e pectens o r s c a l l o p s (Pecten,
Hinnites, f i g . 129,B).
The g r e a t majority of pelecypods belong t o t h e order Eulamellibranchia. This includes t h e oysters, t h e clams, t h e freshwater mussels,
t h e boring clams and t h e shipworms (yeredo). They a r e characterized by
lamellar g i l l s composed of completely joined filaments.
See s l i d e s of permanent mounts of s e c t i o n s of Mya and Mytilus t o
show these two types of g i l l s t r u c t u r e and study pieces of t h e g i l l s of
each from f r e s h l y collected specimens. On t h e s e last note a l s o t h e
powerful c i l i a r y a c t i o n .
The common names o r d i n a r i l y used f o r groups of pelecypods a r e not
derived from t h e names of t h e o r d e r s but a r e t r u e vernacular names,
such a s oysters, scallops, rock o y s t e r s o r jingles, sea mussels, f r e s h water mussels, clams, e t ~ .
C l a s s i f i c a t i o n of pelecypods t o genus and species is based l a r g e l y
on characters of t h e s h e l l . On t h e s h e l l given you, o r i e n t yourself by
remembering t h a t t h e h i w e is d o r s a l and t h a t t h e &
p
l i e s nearest t h e
mantle f o l d s and
--

ELErnM

The cardinal teeth are directly under the umbo and the lateral
teeth either in front of or behind the umbo, if in front they are
antero-laterals, if behind posterior-laterals
Understand the following commonly encountered terms which are
illustrated by demonstration specimens; inteari~alliate,lacking a
pallial sinus (Cli nocardium); sinurwlliate, with a pallial sinus
(Protothaoa); dimrian, with two acauctor muscles; iso arian, with
adductor muscles approximately equal in size (Protot-eromvarian, with adductor muscles strikingly unequal (Mytilus ); monowarian, with a single adductor muscle (~strea,Pecten); chondro~hore
(fig. 121), shelf-like process of-left valve to which is attached the
ligament (Hya ); myophore (fig. 128), a more or less extended, inwardly
directed, process to which muscles are attached in the boring clams.
Identify your specimen by means of the key which follows.
Read the introduction to Illustrated Key to West North American
Pelecmod Genera, by Keen and Frizzell, and identify several specimens
by it. This excellent publication should be owned if possible and consulted extensively.
Each student should sample also Veymouth, Oldroya, and Packard.

KEY TO TBE MORE COUON PEUCYPODA

1. Two valves distinctly different due to attachment by one valve;

one or both valves markedly rough and irregular (fig. 129,A,B)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
...............
.............................
........................
......................
.......
........
............

1. Two valves approximately alike, not especially irregular; at2

tached by byssus or not at all
2. Brown to black; valves elongated, with umbo at or very near
end
3
2. Light colored; valves not especially elongated or if so not with
umbo near end
8
3. Rock borers; narrow
4
5
3. Attached to rocks or piles, or half buried in mud
4. Smooth externally
"The Smooth Pea-Pod Borer," Botula californiensis
4. With fine, transverse ridges over most of shell; color uniform
"!he Rough Pea-Pod Borer," Botula Jalcata
4. With coarser, oblique ridges at posterior end; color obscured
over part of shell; shell wider than in Botula but with posteLithophaga pluwla
rior end markedly pointed
6
5. Hinge and beaks at extreme tip NYti lus )
7
5. Hinge and beaks at side or subterminal (ddodiolus)

..........
............
.......

IUORBTORY AND FIELD

152

TEXT IN INVERTEBRBTE ZOOLOGY

6. Larger; narrower with a few longitudinal r i b s ; growth l i n e s

prominent; brown t o black-brown
"The California Sea h s s e l , " Mytilus californianus
"The Bag Mussel," Hytilus edulis
6. Smaller; broader; smooth; black

.......

7. Large; without r i b s ; width nearly uniform; buried portion dark

brown, polished; f r e e portion yellowish, hairy
Modi olus (= Volsella) rectus
7. Smaller; with prominent radiating, longitudinal r i b s ; hinge end
narrowed
Jfodiolus demissus

..................
...................

8. Both valves swollen a n t e r i o r l y and sculptured f o r boring rocks

( f i g . 129,C)
8. Not so (except f o r Petricola pholadiformis which i s sculptured
anteriorly)

. . . . . . . . . . . . . . . . . . . . . . . . . 32
......................... 9
9. With radiating r i b s . . . . . . . . . . . . . . . . . . . . . . 10
9. Without r i b s but often s t r i a t e d . . . . . . . . . . . . . . . . 14
10. Each valve with f l a t expansions i n front of and behind umbo;
monomyarian . . . . . . . . . . . . . . . . . . . . . . . Pecten
10. No such expansions; dimyarian . . . . . . . . . . . . . . . . . 11
11. Ribs of approximately same type and s i z e throughout; s h e l l
suborbicular o r subquadrat e . . . . . . . . . . . . . . . . . 12
11. Tmith a few l a r g e a n t e r i o r r i b s and numerous narrow r i b s on
Petricola pholadiformis
remainder of s h e l l ; s h e l l elongated
12. Very small, s h e l l subquadrate
Glans carpenteri
12. S h e l l l a r g e r , suborbicular
.13
13. Ribs broad and prominent; growth l i n e s not marked; margin
scalloped; i n t e g r i p a l l i a t e
"The Basket Cockle," Clinocardium nuttallii (= ~ardium)
13. Ribs narrow; growth l i n e s prominent; margin not scalloped but
often distorted; s i n u p a l l i a t e
."The Rock Cockle," Protothaca (= Venerupis = ~ a ~ h i astaminea
)
14. Concentric growth l i n e s prominent, often protruding; shape
15
irregular, often d i s t o r t e d owing t o i t s being a n e s t l e r
14. Not so
17
15. Growth l i n e s i n form of t h i n prominently projecting flanges,
Irus (= Penempis) lamellifera
15. Not so
16
16, & d i m cardinal tooth ( f i g 127 ) prominent, plate-like; growth
l i n e s close s e t and r e l a t i v e l y regular; s h e l l without
periostracum; t i p s of siphons bright purple
."The Rock Dweller," Petricola carditoides
16. Without cardinal t e e t h ; growth l i n e s i r r e g u l a r i n shape and
distribution; s h e l l chalky, covered with a brown periostracum,
some remnants of which a r e present i n nearly a l l old s h e l l s ;
siphon t i p s red
Saxicava pholadis
17. I n colony of compound ascidians; higher than long
hfytilimepianuttallii
18
17. Not associated with ascidians

..

. .. .. .. .. .. .. .. .. .. .. . . . . . .

....
.

. . . . . . . . . . . . . . . . . . . . . . . . . .. .. ..
. . . .. .. .. .. .. .. .. .. .. .. .. .. .. .. . . . . . . . . . . . . . .
.
...........

...............

.....................
.................

. . . . . . . . . . . .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..

19
18. Small, less than an inch long
18. Not so
24
19.kine . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 0
dlusculium
19. In freshwater; teeth minute or obsolete; suborbicular
21
20. 5 mm. long or less
23
20. More than 5 mm. long when adult
21. Shell rosy; with transverse chondrophore in both valves;
nestling in barnacle beds
Unidentified
22
21. Shell white; without transverse chondrophore
22. Sinus deep, dorsally directed; latera1 teeth not conspicuous
Gemmagemma
22. Sinua not noticeable or anteriorly directed; lateral teeth
conspicuous, ridge-like
Psephidia

......................
................
.............
.........

..........................
...............

23. Nestler in pholad holes; teeth prominent; shell short, oval,

smooth with gleaming green periostracum
Kellia (= Chironia) laperousii
23. Often in kelp holdfasts; almost three times as long as high;
teeth inconspicuous
Lyonsi a californi ca
24. Long-oval, much elongated; with a prominent ridge running downward from the teeth on the inside of each valve
l1 The Razor Clams," S
i1i qua
24.Notso . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 5
25. Both valves bent to the right at the posterior end
"The Bent-Nosed Clam," Macoma nasuta
25. Not so
26
26. Without chondrophore
27
26. With chondrophore (fig. 127) or pseudochondrophore
29
27. Thick; subquadrate; gaping widely "'The Geoduck," Panope generosa
28
27. Thin, ends rounded or pointed
28. Larger, up to 4 inches long; posterior end somewhat. pointed;
without conspicuous periostracum
Macoma secta
28. Smaller, up to 2 inches long; both ends rounded; with conspicuMacoma irus (= inquinata)
ous brown periostracum
29. A vertical chondrophore on each valve; a brown tent-shaped
cardinal tooth on left valve; gaping widely behind
."me Giant Clam,"Schiaothaerus
29. A conspicuous horizontal chondrophore on one or both valves . 30
31
30. Umbo at or near center
30. Umbo near posterior end; both ends truncate
Platyodon cancellatus
31. Posterior end truncate, with faint external striations; commensal with Callianassa
Cryptomya californica
31. Posterior end somewhat pointed, without external striations
Mya arenaria

.................
............

...................

..............
............................
.....................

......
. . .. . . . . . . . . . . . . . .
.........
.......

................
.
....................
.....................
.........
..........................

154

LCLBORAMRY AND FIELD

TEXT I N

llmm?l"rBRATE ZOOLOGY

32. Sculpture fine to moderate, confined to a triangular section

of anterior half of shell, consisting of fine concentric
ridges undulated by fine radial ridges; with one or more
conspicuous accessory plates along dorsal margin (fig. 128) 33
32. Sculpture coarse, over anterior half of shell, consisting of
coarse concentric ridges prolonged into spines at regular
intervals which are therefore in radiating rows; without
Zirfaea
conspicuous accessory plates

.............

Fig. 128.. Rock borers (aFter Keen and ~ r i c z e l l ) :A-C, m ~ h o l a s

( ~ o n r a d ) ;D-F, Pholaoenita Conrad.

33. Dorsal margin doubled and reflected both anterior and posterior
to beaks; posterior half of shell divided approximately in
half by a conspicuous radiating angle which marks change in
direction of concentric lines; a conspicuous posterior extension of brown periostracum; siphon tips pale lavender
Paraphol as cal iforni ca (fig 128,A-C )

.............

Fig. 129. A , Pododesmus macroschidma (Deshayes); B, H i m i t e s multirwosue ( s e e

Gale) (s " giganteus Gray")8 C, Pholadidea penita Conrad. ( ~ l from
l
Guberlet )

rnCYP0DA
33. Dorsal margin doubled only anterior to beaks; without conspicuous radiating angle; no posterior extension of periostracum;
siphon tips white with faint brownish tinge
Pholadidea penita (figs. 128,E-F, 129,C)

............

34. Lower valve with hole or deep notch near hinge

. . . . . . . . . . . "The Rock Jungle," Pododesms (fig. 129,A)

. . . . . . . . . . . . . . . . . . . . . . 35

34. Both valves entire

35. Iarge, approximately circular, with distinct wings on either

side of umbo; purplish within; with radiating ribs; left valve

and younger portions of right valve distorted
"The Rock Scallop," Hinnites (fig. 129,B)
35. Small, with pointed beak; elongate to circular; surface of upper
valve laminated or irregularly plaited
."The Vestern Oyster," Ostrea lurida

...........

..............

LIST OF PELECYPODA
Order Filibranchiata
*I Botula californi ensis Philippi
*M Botula fa1 cata Gould
D *M Hinnites m l t irugosus (Gale) (= "if. giganteus Grayff)
M Lithophaga plumula Fianley
*B dlodiolus (= Volsella) demissus Dillwgn
*I Modiolus rectus Conrad
D *Id Myt ilus californianus Conrad
*B Mytilus edulis Linnaeus
D *M Pododesms macroschismus Deshayes
Septifer b ij'urcatus (Conrad)
Order EulamRllibranchiata

TB Macoma secta Conrad

M Milneria minima (Dall)
*I?
Musculium
*B Id Mya
arenaria Linnaeus
D M Mytilimeria nuttallii Conrad
B Ostrea g igas Thunberg, Japanese Oyster
*B M Ostrea lurida Carpenter
TB Panope generosa Gould, The Geoduck
M Parapholas californi ca (Conrad)
*M Petricola carditoides Conrad
B Petricola pholadifonis Lamarck

156

LABORATORY AND FIELD TEgT I N INVERTEBRA'%E ZOOLOGY

Order Eulamellibmchiata (Contiwed)

D M Pholad idea parva Tryon
D *M fioladidea penita (Conrad)
M Platyodon cancellatus Conrad
*B D *M Protothaca (variously Paphia or Venerupi s ) staminea
(Conrad)
D M Psephidia lordi Baird
TB Sangu i no1 ar i a nut ta11i i (Conrad)
D *M Smi cava pholad is (Linnaeus)
TB Saxidonus nuttallii Conrad
B TB Schizothaems nuttall i i (Conrad)
B Siliqua lucida Conrad
B Sil iqua patula (Dixon)
TB Tell i na bodege~si s Hinds
*B Teredo navalis Linnaeus
M Zirfaea pilsbryi Lowe (= 2. gabbi)
PHYLUM ENTOPROCTA (Kamptozoa, Calysmzoa)

This group has usually been included with the Bryozoa, but differs
from it fundamentally in structure. !The term entoproct refers to the
inclusion of the anus within the circle of tentacles (the anus is outside in the true or ectoproct Bryozoa). Externally, the simplest way
to distinguish the two is to note whether the tentacles are folded into
the center when withdrawn (as in a hydroid), in which case it is an
entoproct, or whether the whole anterior end of the body is introverted
into the basal prt, with the tentacles still pointing forward, in
which case it is an ectoproct . The nodding movement of individual zooids
is also characteristic of the Entoprocta. The Entoprocta are much less
common than the Ectoprocta, but will probably be collected as they are
often found forming small, moss-like patches on coralline algae.
PHYLUM BRYOZOA (Ectoprocta)

Under the low power of the microscope see that the colony or zoarium
is composed of numerous individuals. W e yourself familiar with the
terms defined below which commonly occur in keys and descriptions. Each
individual consists of a zooecium (pl. zooecia) and a polsnide. The
zooecium is a chamber or sack, the proximal portion of the individual,
and is covered externally by a secreted wall of calcareous, chitinous,
or rarely, gelqtinous, or membranous nature. The polypide is the distal
half of the individual, when extended, including the 1oDhoDhow (which
bears the 3entacle) and the alimentary tract. The polypide is retractile
within the zooecium. The opening at the end of the zooecium when the
polypide is withdrawn is the orifice (Plate XVI,B). The front wall of
the zooecium is often called the awrture. Do not confuse this with
the orifice.
Avicularia (sing. avicularium) are birdts-head-likeappendages of
the zooecia of certain species. Vibracularia (vibracula) are long setalike a pendages of the zooecia of certain species. !Pb ooecium (pl.
ooeciaj or ovicell is an expanded chamber, above a zooecium, in which
the embryos develop.

BRYOZOA

157.

Zoaria may be creening, erect, suberect, encrusting, flexible,

or stiff. They may be articulated (divided into internodes by joints

or nodes) or n o n a r t i c u m e XVI ).
See expanded living individuals. Seek to identify common species
by meane of the key which follows.
TO T.EE BRYOZOA AND ENTOPROCTA
A PFU?,LU6INARY
OF TBE MOSS BEACH REGION
1. Individuals composed of distinct stalk and body regions and
connected by stolons; individuals separately moveable. EN!NPROCTA (base of stalk bulbous)
Pedicellina (Barentsia) gracilis (Plate XV1,A)
1. Colony (zoarium) composed of individuals (zooecia) not readily
divisible into stalk and body regions; individuals usually not
movable. ECTOPROCTA
2
2. Colony gelatinous; zooecia without calcareous skeletone CTENOS T O M A T A . . . . . . . . . . . . . . . . . . . . . .
3
2. Zooecia with calcareous skeletons
4
3. Zoarium brownish, covered with branched reddish, spine-shaped
pmjections which arise from between the zooecia; zooecia
hardly discernible; colony resembling seaweed

.........

......................

......

................

. . . . . . . . . . . . . . . . . . . . .Alcyonidium cervicornis
.......................
................
...........................

3. Zoarium gray or colorless, without spines; zooecia with distinct

outlines
A.myti1i
4. Zooecia tube-like, with orifice at one end (Plate XVI,C,D);
colony branching. CYCLOSTOMILTA
5
4. Zooecia box-like, orifice on "frontn (Plate XVI,B,E). CHEILOSMUTA
6
5. Zooecia uniserial, with jointa between each; joints black,
giving characteristic speckled appearance to older parts of
Crisidia franciscana
colony (Plate XVI,C )
5 . Zooecia biserial; joints few, not present between each zooecium
(Plate XV1,D)
Crisia pacifica
7
6. Zoarium encrusting rocks, algae, shells, other Bryozoa, etc.
6. Zoarium not encrusting, but forming a branching mass
.17
7. Front almost entirely occupied by large orifice
8
7. Orifice covers not more than half of the front
9
8. Avicularia sessile, in spaces between zooecia
Callopora horrida
8. Avicularia stalked, among spines surrounding orifice (Plate XVI,F)

...........
..................

..
.....
.........
.........
..

. . . . . . . . . . . . . . . . . . . . . .Cauloramphus spinifer
9. Front wall calcareous, crosaed by somewhat irregular rows of
large pores . . . . . . . . . . . . . . . . . . . Cri bril ina sp.
9. Front wall calcareous, not traversed by rows of pores, but
sometimes with scattered small ones . . . . . . . . . . . . . 10
9. Front wall membranous, with small semicircular orifice; zooecium
oblong, margins slightly raised

. . . . . . . . . . . . . . Membranipora membranacea

(fig. 130,A)

I
I

PLATE XVI

r+**.*
.-,,

p"""".

-.Y
\A

an

bur

"
------

--

C
"-r i n i d i n

pacifica

franoisc~na

B
Sohsmatio aooeoium

Microporella oiliata

- ...--

Muoronella sp.

Bryozoa and Entoproota (figures partially after ~obertson)

BRYOZOA
10. Zooecium possesses a special semicircular pore, with a sieveplate i n it, below the o r i f i c e ; o r i f i c e semicircular, with
straight lower margin (Plate W I , E )
10. Zooecium without special pore

. . . . . . . . . . . . . 11

. . . . . . . . . . . . . . . . . 12

Fig. 130. Bryoaoan colonies (from ~ u b e r 1 e t ) rA , $fembrani~oramembranacea ( ~ i n n a e u s ) ;

B, R e t e m r a ~ a o i f i c aRobertson: C , Flustra lichenoides Robertson.

i
i
I

Ii

j
1

11. Zooecium bears spines above the o r i f i c e

Yicroporellaciliata (PlateXVI,E)
11. Zooecium does not bear spines
hf. californica
12. Orifice simple, usually contracted near the lower margin, covered
with brown operculum; color of zoarium i n l i f e old rose
.Eurystomella bilabiata
12.Notasabove
13
13. Orifice simple, with narrow sinus on lower margin; zooecium
with transverse ridges (Plate WI,G)
Hippothoa hyal
- ina
13. Secondary o r i f i c e developed by growth-of peristome -(plate
WI,H)
14
14. Orifice with deep sinus on lower margin; zooecia erect or suberect, heaped together
Costazzia incrassata
14. Orifice without sinus on lower margin; zooecia not erect
15
15. Peristome elevated, forming secondary o r i f i c e channeled on
lower margin (Plate W1,B); primary o r i f i c e dentate
16
15. Peristome elevated, forming secondary o r i f i c e , which has a
mucro o r tooth on its lower margin (Plate XVI,H)
Micronella sp.

..............

...........

....................
.........................
....
...........................
..........

...
.....

.........................

16. Zooeciumwith two spines on upper margin of o r i f i c e and two or

three large blunt protuberances below the o r i f i c e
Smittina collifera
16. Zooecium spineless, with no large protuberances
Smittina reticulata

.......................
......................

160

LBBORBTORY AND FIELD TEXT IN IXVER'I'EBRATE ZOOLOGY

17. Branching regular; one or more main trunks, from which periodic
whorls of branches are given off ; colony has the appearance
Bugula californioa
of having been twisted into a spiral
17. Branching much less regular than above; branches do not come
off in whorls
18
l8. Zooecia have only lateral avicularia; joints occur usually beNenipea occidentali s
tween every three zooecia
18. Zooecia have frontal avicularia as well as lateral ones. bre
than three zooecia between each joint
Scrupocellaria californica

....
........................
.........
...................
LIST OF ENTOPROCTA AND BRYOZOA
Barentsia (= Pedicel1ina) gracilis

Order Phslactolaemata
Plumatella (fresh water)
Order Gsmnolaemata
Suborder Ctenostomata
D *M Alcyonidium cervicornis Robertson
Alcyonidium mytili Dalyell
Suborder Cyclostomata
Crisia pacifica Robertson
Crisidia Qranciscam Robertson
Suborder Cheilostomata
D *M Bugula californica Robertson
Callopora horrida Hincks
Caulormphus spinifer Johnston
Costaaaia irrcrassata h r c k
Cribrilina sp.
%rystomella bilabiata Hincks
Hippothoa hyal ina Linn.
Menipea occidentalis k s k
B TB D Membranipora membranacea Linn.
Hicroporella californica Wlsk
Microporella ciliata
Mucronella sp.
Scrupocellaria californica Trask
Sinittina collifera Robertson
Sinittina ret iculata Macgillivray

PHYLUM PHORONIDEA
This is a very small phylum with but two genera ( ~ h o r o n i sand
Phoronopsis ) and r e l a t i v e l y few species. Under favorable conditions
they occur i n great numbers as i n Elkhorn Slough, Bolinas Bay, and
Tomales Bay. In t h e upper end of Tomales Bay students have estimated
t h a t over a b i l l i o n individuals were living i n one acre. Six species
have been reported from the Pacific Coast, two from British Columbia,
one from Humboldt Bay and m e t Sound, and three from southern California. Just which species (one, two, o r more) occur i n central
California is not lmown. The special paper of T. H. Bullock represents the only attempt t o identify these animals i n t h i s vicinity.
Study preserved specimens. Keep i n mind t h a t while they a r e
elongated worm-like animals the mouth and anus open close together
and t h a t terms of orientation such as anterior and posterior must be
used with caution. Note the lophophore, the great double semicircular
whorl of tentacles with incoiled ventral edges, the c i l i a on which
serve t o s e t up currents which bring plankton t o the mouth.
I f time permits, study sections t o get some idea of the i n t e r e s t ing anatomy of t h i s type. Consult Borradaile and Potts o r Cambrike
Natural Historg f o r figures

PHYLUM BRACHIOPODA
A very few small individuals of t h i s small phylum a r e found a t
the lowest t i d e levels i n t h i s region. They belong t o a species, Terebratal i a transversa (Sowerby ), found i n large numbers i n deeper waters
or i n shallow waters f a r t h e r north.
The two-valved s h e l l suggests a pelecypod mollusk but they a r e
e n t i r e l y different s t r u c t u r a l types. Superficially they a r e e a s i l y
differentiated by the unequal valves of the s h e l l which a r e dorsal and
ventral instead of r i g h t and l e f t .

PHYLUM ECHINODERMATA
The present -day echinoderms f a l l into f i v e d i s t i n c t classes, the
Asteroidea or sea stars (starfishes), the Ophiuroidea or b r i t t l e s t a r s ,
the Echinoidea o r sea urchins, the Holothuroidea or sea cucumbers, and
the Crinoidea o r crinoids (sea l i l i e s ) . A l l save the last c l a s s have
representatives i n the l o c a l fauna. W e certain that you are able t o
recognize and distinguish the members of these classes. Since only the
c l a s s Asteroidea is represented by a number of species, we s h a l l conf i n e our laboratory study t o t h e sea stars.
o r arms. the o r a l surOn a sea star make out the radiating
face with the central m u t h , the ambulacral m o v e , i n the o r a l surface
of each arm containing the tube f e e t (podia o r -en&es).
On the aboral s f e locate the anus and the madre~oric late and note
the absence of t e feet. Note the
t h e i r -,
and
arrangement a r e of importance i n classification. Using a lens seek

ti?

a;

a,

162

UBORAMRY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

minute pincer-like organs, t h. e- e~

These a r e more s t r i k i n g l y
developed on the sea urchin. Examine the l i v i n g demonstration specimen
under the microscope.

buccal p l a t e
jaw
mouthmouth papillae

arm spines

--&m
..;. - -.. .-. .
.- ... :9

1.:.

opening of g e n i t a l bursa

tentacle (moified tube f o o t )

tooth papillae
buccal plate (mouth parts)

tentacle s c a l e

F i g . 131. Mouth parts of b r i t t l e stars: A , O~hioderma; B, b h i o ~ t e r i s .

Scrape away the outer body l a y e r s and the spines t o expose the
p l a t e s of the skeleton of one e n t i r e arm and the area around mouth and
anus.
Using P r a t t when available, attempt t o determine t h e family of
one species. Meanwhile, i d e n t i f y your specimen, using the key below.
On a b r i t t l e s t a r locate mouth with " jawsw (fig. 131) and o r a l
papillae, buccal plates, g e n i t a l bursae, t e n t a c l e s (modified tube f e e t ) ,
disk scales, r a d i a l plates, dorsal arm plates.
Identify your specimen using the key below.

ECHINOD?BuuIATA

KEY TO TEE ECHINODERW

Holothuroidea (Sea Cucumbers)
1. Pale (usually white) i n color; lacking tube f e e t ; small i n size
Leptosynapta albicans
2
1. Colored; tube f e e t present
2. Tentacles 20 or more, f a i r l y short, each somewhat resembling a
flower; size of animal large
Stichopus cal ifornicus
2. Tentacles about 10; tentacles longer, dendrit i c (tree-like i n
.3
branching)
3. Dorsal and ventral surfaces of body markedly distinguished from
27tyonepsolus nutriens
each other; small; red i n color
3. No distinction of dorsal and ventral surfaces; some species
large, some small
Cucumuria Spp.

. . . . . . . . . . . . . .. .. .. .. .. .. .. .. .. . . . . . . . . . .

......
.........................
......
................
Echinoidea (Sea Urchins)

1. Body very flattened; spines short; below i n t e r t i d a l zone

"The Sand Dollar: Dendraster excentricus
1. Body not greatly flattened; spines long. "Sea Urchins"
2
2. Small (2 inches o r l e s s i n diameter); purple; spines r e l a t i v e l y
short
Strongylocentrotus purpuratus
2. Large; red o r purple; spines longer

............

.....

...............
. . . . . . . . . . . . . . . . . Strongylocentrotus franciscanus

( B r i t t l e Stars, w l

ong mobile arms)

.........,........

2
1. Diak with overlapping scales
1. Disk with granules o r granule-like spines; r a d i a l plates covered; lowermost spines claw-like near t i p of arm
Ophiophol is aculeata var. kennerlyi (Clark, p. 132)
2. A single dorsal arm plate t o each segment
3
2. litmy dorsal arm plates of varying size; four mouth papillae, f i r s t
twice a s long as each of the others
Ophioplocus esmarki
3. Smaller, disk usually l e s s than 5 mm. wide; r a d i a l plates i n
contact save a t proximal t i p , three pairs of o r a l papillae,
first a s long a s next two together; dorsal arm plates not i n
contact, l e s s than twice a s wide a s long
Amphipholis pugetana
3. Larger, disk usually about 10 mm. wide; r a d i a l plates separated
by smaller disk scales save a t bases; a single pair of o r a l
papillae; dorsal arm plates broadly i n contact, twice a s wide
aslong.
AmpSLibdiaoccidentalis

.......

............
.....
.

................
Mteroidea (Sea S t a r s )

1. Leathery outer covering

. . . . . . . . . . . ."The Leather Star," Dermsterias imbricata

......................2

1. Spiny outer covering

LABORATORY AND FIELD

164

TMT I N INVERTEBRATE ZOOLOGY

2. About twenty arms

"The 20-rayed o r Sunflower Star," Pyonopodia he1 ianthcides
3
2. Not more than s i x arms
4
3. Small, $ t o 28 inches i n diameter
6
3. Large, with f i v e arms
4. Five arms; spines of aboral surface i n elongated groups (pseudopaxillae)
young of Henricia leviuscula
5
4. With s i x arms; spines a s usual
5. Small, usually l e s s than one inch i n diameter; arms slender;
spinulation sparse; common i n t i d e pools of mid-tide horizon
Leptasterias pusilla
5. Larger, reaching a diameter of 2$ inches; arms s t o u t e r , somewhat swollen a t base; spinulation close-set; common i n t i d e
Leptasterias aequal i s ( f i g . 132,A)
pools of low-tide horizon

...

.....................
................
......................
.............
.................

......................

..

F i g . 132. Sea stars (from ~ u b e r l e t ) :A , JleDtasteriasr B. Henrioia leviusouler

(Stimpson).

. . Patiria ( ~ s t e r i w )
. . . . . . . . . . . . .7

6. Disk very broad; arms short and t r i a n g u l a r

6. Disk smaller; arms long, subcylindrical

7. Arms slender; spines very t i n y , i n groups (pseudopaxillae); color

an even orange red
Henricia leviuscula (fig. 132,B)
7. Spines more conspicuous; body and arms s t o u t e r
8

......

.........

8. Spines conspicuous, forming a network

"The Ochre Starfish," Pisaster ochraceus
8. Very large; spines conspicuous but not forming a network

............

.......................
...............

Pisastergiganteus

8. Spines very short and inconspicuous, forming a network; pale

pink i n color
Pisaster brevispinus

i"
"x

ENTERoPNElJsTEl
LIST OF ECHINODEFM

M Cucumria Spp
M Leptosynapta a1b icans (Selenha)
M Stichopus californicus (Stimpson)
M Thyonepsolus nut riens Clark
ECHINOIDEA
excentricus (Eschscholtz)
*MM Dendraster
Strongylocentrotus franci scanus (A@;assiz)
*M Strongylocentrotus purpuratus (Stimpson)
OPHIUROIDEB
M Amphiodia occidentalis ( L m )
M Amphipholis pugetana (Lyman)
M Ophiopholis aculeata kennerlyi (-1
M Ophioplocus esnrarki Lyman

Dermasterias imbricata (Grube)

Henricia leviuscula (Stimpson)
Leptasterias aequalis (~timpson
)
Leptasterias pusilla Fisher
Patiria ntiniata (Brandt)
Pisaster brevispinus (Stim son)
Pisaster ochraceus (Brandt
Pisaster giganteus (Stimpson)
Pycnopod ia he1ianthoides (Brandt)

PHYLUM CHORDATA

SUBPHPLIJb! HEMICHORDA
Class ENTEROPNEUSTA
One species of this group (Mesoglossus intermedia) has been found
in small numbers at Moss Beach. The Fntsropneusta are worm-like animals
(our species small, generally not more than an inch long) which can be
distinguished from other worm-like creatures by the division of the
body into three parts; proboscis, collar, and trunk. The species which
has been found at Moss Ekach is small and is taken on the under sides
of rocks lying in coarse sand in protected tidal channels or lagoons.
SUBPHPLWd UROCHORDA ( TUN1CATA
There are three classes in this subphylum, the Larvacea, Thaliacea,
ia. The first consists of small pelagic forms which will not
red in this course. The Thaliacea are also pelagic, but are
common. Some may occasionally be seen floating in

166

LABORATORY AND FIELD TEXT I N INVERTEBRATE ZOOLOGY

the water or washed ashore. They are s o f t , jelly-like, transparent,

colorless except f o r a r e l a t i v e l y small "nucleus." Salpa is the genus
commonly encountered. The Ascidia a r e attached t o rocks o r p i l i n g and
form a conspicuous element i n our l i t t o r a l fauna. Borne of them a r e
s o l i t a r y ; they a r e sac-like and possess two o r i f i c e s a t o r near the
d i s t a l end: one, the incurrent o r branchial o r i f i c e , the other the excurrent or a t r i a l o r i f i c e . The majority of species, however, a r e colon i a l . !The individuals, called zooids, may be merely united by t h e i r
outer layer ( t e s t ) o r t h e i r c a v i t i e s may be variously united. One common arrangement i s the fusion of several a t r i a l o r i f i c e s i n t o a common
cloaca1 o r i f i c e , the branchial o r i f i c e s remaining separate, arranged i n
a c i r c l e around the cloaca. The s t i m a t a referred t o i n t h e key a r e
i n t e r n a l g i l l openings between the branchial and a t r i a l c a v i t i e s . See
Borradaile and P o t t s o r another t e x t f o r other i n t e r n a l characters
mentioned i n the key.
Many d i f f e r e n t species of compound ascidians a r e known. b4my of
them form smooth, lobed colonies and a r e often spoken of as sea pork
( f i g . 135).

A KET TO THE MORE COMMON LITTORAL ASCIDIlWS OF

By Robert L. Fernald

TBE ME8 BEACH AREA

.......
..............
.........
...................

1. Individuals fixed and s o l i t a r y . ASCIDIAE SIMPLICES

2
1. k n y individuals more o r l e s s completely embedded i n a common
cellulose mass. ASGIDIAE COMPOSIW
4
2. Long, club-shaped; peduncle or s t a l k a t l e a s t a s long a s body;
prevailing color dark red
Styela rnontereyensis
3
2. Without peduncle; sac-like
3. Test translucent and without color; p l a t e s surrounding the a t r i a l
and b r a c h i a l o r i f i c e s
Chelyosona producturn
3. Test tough and coated with foreign material; basic red-brown
Halocynthia Sp.
color with bright-red siphon t i p s
4.

..........
........
Zooids with basal portions only embedded i n common t e s t ; connected by stolons. "Social Ascidians" . . . . . . . . . . . . . 5

4. Zooids e n t i r e l y embedded i n common cellulose mass

. . . . . . . .6

5. Individuals mound-like, 2 t o 4 mm. i n diameter; colored various

shades of red
Metandrocarpa michaelseni
5. Individuals ovoid, 2 mm. by 3 mm.; pale greenish-yellow; t e s t
transparent; attached t o rocks, c o r a l l i n e algae, etc.
Perophora annectens
5. Individuals elongate, club-shaped, 2 t o 4 cm. high; t e s t transparent; pink i n t e r n a l organs c l e a r l y v i s i b l e . Clavelina huntsmani

.............

.......................

6. Colony broken up i n t o few o r many club-shaped masses each with a

peduncle about twice a s long as the more o r l e s s rounded head 7
6.Colonyotherwise . . . . . . . . . . . . . . . . . . . . . . . . 9
7. Sand-encrusted
8
7. Test f r e e of sand, transparent; yellow-orange zooids (common)
Macroclinumpellucidum

.........................
.....................

rnICATES
8. Common cloacal orifice in small heads; zooids with 16 seriee of
stigmata; atrial orifice with flat, truncate, languet; stomach
cylindrical with seam on left surface but free from foldings
(fig. 133,A)
Macrocl inum par-fust is
8. No common cloacal orifices; zooids characterized by 8 series of
stigmata; atrial and branchial siphons of about equal length;
wall of stomach with 6 or 7 longitudinal folds (fig. 133,B)
Amuroucium aequali-siphonis

..............

...................

.........
........................

10
9. Numerous calcareous spicules present in test
9. Spicules absent
11
10. Colony 1 t o 4 mm. thick; white or flesh-pink in color; zooids
with 4 series of stigmata; spicules with blunt rays (fig.
134.8)
Di demnun c a r m l entum

..................

--

Fig. 134.-A,
spicule of Didemuurn oarnulentturn (enlarged)$ B , spicule of Trididemuum
della v a l l e i (enlarged). (Both a f t e r Ritter and Forsyth.)

168

LABORATORY AND FIELD TEXT IN lXVEBTEBMTE ZOOLOGY

11. Zooids divided into three body regions-thorax, abdomen, and

post-abdomen (fig. 136,A); atrial orifice without lobes but
with long, tapering languet; 8 to 14 series of stigmata; stomach cylindrical with longitudinal folds sometimes branching or
discontinuous (fig. 136,A); abundant
Amaroucium californicum (fig. 135)
12
11. Zooids divided into tmo body regions-thorax and abdomen

...............

Fig. 135.-Amarouoium

...

c a l i f ornioum (from ~ u b e r l e t )

12. Atrial siphon as long as, or longer than the branchial siphon;
13
3 to 5 series of stigmata
12. Atrial siphon not well developed; four series of stigmata
15
13. Colony hard, much sand embedded in the test; brown to claret
in color; zooids in systems; atrial siphon longer than
branchial; three series of stigmata; abundant

.................

..

. . . . . . . . . . . . . . . . . . . . . . . fidistoma psamnion
..............

13. Colony not hard, relatively free of sand; lighter thanE.

14
psammion, zooids not in systems
14. Colony thick and encrusting, pedunculate or lobate; color
opaque white through yellow-brown to light vermillion;
three series of stigmata
Eudistoma diaphanes
14. Colony thick and encrusting or lobate; five series of stigmat8
D i s t o m lobata
15. Colony soft and encrusting, pedunculate-orcapitate; wide
color variation; zooids may or may not be in systems; atrial
languet well developed; stomach ovate, wall not folded but
with a network of ridges on inner surface (fig. 136,B); very
abundant
Distaplia occidentalis
15. Colony soft, thin, and encrusting; systems if present, irregular; atrial orifice large with simple opening; stomach smooth
walled
Diplosoma sp.

..........

......................

................

......................

S~eciesaccounts of the above mav be found in Ritter and Forssth.

- * 1917, Acidians of the ~ittoral"Zone of Southern California.
Univ. Cal. Publ. Zool., 16: 439-512.

VERTEBRATES

A
Fig. 136.-A,
zooid of Amaroucium californicum (enlarged); B , zooid of D i s t a p l i a
occidentalis (enlarged). ( ~ f t e rR i t t e r and Forsyth.)

SUBPHYLUM CRANIATA or VERTEBRATA

We have designated our field as Invertebrate Zoology. The vertebrates, however, have very important relations to the invertebrates
and as predators and in other ways powerfully influence their lives
and the conditions under which they live them. For that reason we give
brief consideration to them here.
Class PISCES

LABORATORY AND FIELD TEXT I N INVERTEBRATE ZOOLOGY

170

gives them again the normal conditions of submergence under which they
a r e equipped t o l i v e . During the f a l l i n g of the t i d e s , also, many of
the animals important i n the l i f e of the seashore swim out with the
t i d e s . The picture gained from low-tide studies is, therefore, very
p a r t i a l f o r both these reasons and every attempt should be made t o
visualize the t r u e conditions of l i f e during high t i d e .
As a i d s t o t h i s we have recourse t o the t i d e pools, o r under
fortunate conditions t o the glass-bottomed boat o r the diving b e l l .
The l a s t two give us, a s does the t i d e pool t o much l e s s e r extent,
some idea of the importance of the f i s h e s i n the i n t e r r e l a t e d commun i t i e s of the i n t e r t i d a l zone. They f i g u r e largely a s predators, putt i n g a premium on numbers, o r swiftness, o r defensive weapons, o r
protective devices and behaviors of many kinds.
We s h a l l not devote any time t o the study of the fishes. The ones
commonly encountered by us w i l l be the small spotted sculpins of the
t ide-pools (01 igocottus ma~u10sus) sometimes called rock-pool johnnies.
This and other common species may be i d e n t i f i e d by the key which f o l lows. For f u r t h e r d e t a i l s o r f o r other species see Keys t o the Fishes
of Washimton. Ore~on. and Closely Adjoinina Regions, by L. P. Schultz,
University of Washington Publications i n Biology, 2: 103-228.
FIELD KEY TO m'FISHES COMMONLY SEEN OR TAREN AT MOSS BEACH
(Figure references a r e t o Walford, Fish Bulletin 28, 1930)

......................
............................

1. Elongated, eel-like
1.Notso.

2
4

2. No paired f i n s ; median f i n s v e s t i g i a l ; g i l l opening a small

hole; head b i l a t e r a l l y f l a t t e n e d
"Green Moray, " Gymnothorm m o r d m ('Jalf ord f i g )
2. Pectoral f i n s present; median f i n s f i n - l i k e ; g i l l s opening a s i n
other f i s h e s ; head dorsoventrally f l a t t e n e d
"Blenny Eels," "Eels," 3

.........

.....................

3. Median f i n s prominent; t a i l f i n d i s t i n c t ; pectoral f i n s longer

Cebidichthys violaceus (Walford, f i g . 110)
than eye

.......

3. Median and t a i l f i n s narrow, inconspicuous; pectoral f i n s very

Xiphister mucosus (Walford, f i g . 109)
short

...........

4. Body very deep and t h i n (Surf Fishes, "Perch"); commonest species

with longitudinal s t r i p e s of d u l l omnge and blue
"The Striped Perch," Taeniotoca lateralis ~Walford, f i g . 81)
5
4. Body not deep and t h i n
6
5. With a v e n t r a l sucker
5. Without sucker
7
6. Tadpole-like rock-pool f i s h , with large f l a t head, and a single
dorsal f i n
"Cling Fish," Caularchus meandricus (Jordan, f i g . 651)
6. Head smaller, normal i n shape; two dorsal f i n s
"Pretty Goby," Lepidogobius 1 epidus

...

.....................
......................
.........................

......
.? . . . . . . . . . . . . .

BIRDS

........................

7. Two dorsal f i n s
7. A single continuous dorsal f i n from head t o t a i l ; small spotted

tide-pool f i s h

.......

"Spotted Kelp-Fish o r Senorita," G i bbonsia elegans

. . .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..
.....................

8. Without s c a l e s
9
8.Withscales....
12
9. Small rock-pool f i s h
10
9. Large, dark i n color with extreme v a r i a t i o n i n color and patt e r n "Cabezone," Scorpaenichthys marmoratus (Walford, f i g . 102)
10. With abundant c i r r i , giving woolly appearance
"Woolly Sculpin, l1 Cl i nocottus anal is
10.Withfewifanycirri.
11
11. With ugly depressed head bearing a strong perpendicular spine
with two o r t h r e e hooks; greenish above, abruptly white below;
pectorals and caudal banded with black
"Smooth Sculpin," Leptocottus armatus
11. Commonest rock-pool f i s h ; head normal, spine very short; with
many color shades and patterns
"Rock-Pool Johnny," 01 igocottus maculosus (Jordan, fig. 566)
12. Three large projecting spines i n a n a l and 12 o r 13 i n a n t e r i o r
dorsal; a n a l with few rays; a single l a t e r a l l i n e
."Rock Cods," Sebastodes spp. (\Talford, f i g s . 90-101)
(Many species, usually separated with d i f f i c u l t y )
12. No projecting spines; a n a l with about 20 rays, dorsal with about
21; f i v e l a t e r a l l i n e s ; males and females d i f f e r e n t i n shape
and color (see Walford)
"California Sea Trout, l1 Hexagrammos decagranimus

..............
....................

..............

..

......

.........

Class
I n contrast t o t h e f i s h e s , birds a r e most a c t i v e i n the i n t e r t i d a l zone during low t i d e s when they pick up great numbers of invertebrates. Most important, no doubt, a r e the g u l l s which i n addition t o
being scavengers e a t great numbers of small animals, notably sand crabs
( h e r i ta ) and the mud crab (Hemigrapsus oregonensis ) and a r e , therefore,
the terminal animals of a number of food chains. Several g u l l s occur
along our shores a s seen from the list below, but only one, the Western
Gull, is a summer resident.
Other birds feeding l a r g e l y on shore animals a r e the curlews, the
sandpipers, the oyster catchers, and some ducks such a s the scoters.
The curlews and sandpipers feed l a r g e l y upon dwellers i n the mud
and sand and the list of animals involved i s probably a large one, consisting chiefly of species of Crustacea and Polychaeta.
One animal which occurs i n enormous numbers at times and serves a s
a most important key industry animal, since i t is fed upon by most of
the birds ment 'oned, i s the sand crab (Emerita analoga). Only a l i t t l e
o~servation*
show t h e g u l l s t o be regular hunters f o r these crustaceans. Likewise even t h e curlews and smaller shore birds can be seen
joining the f e a s t , shearing off t h e l e g s and eating them when the animal i t s e l f is too l a r g e f o r them.

LABORATORY AND FIELD TEXT IN ImTEBRATE ZOOLOGY

172

As for the scoters they put on a truly amazing performance in

their search for these sand-dwelling plankton feeders. lashing up with
a wave they follow the backwash of it down the sandy beach for a short
distance, digging madly for fierita, then waddle clumsily down to
repeat the process on another wave.
Besides these birds which feed directly on the invertebrates there
are several which feed on the fish which themselves feed on the invertebrates. Important among these are the pelicans and several species of
cormorants (see list below).
Finally we might mention birds of entirely different food habits,
the plant feeders, which affect the invertebrates only indirectly. An
interesting bird of this type to be seen at Dillon Beach and at A.30
Nuevo Island is the black brant, which seems to feed entirely on extensiva local growths of an Ulva-like green algae.
A list of the common and scientific names of the commoner species
of birds of the types mentioned here is given below. For distinctions
between the species see Hoffmann, Guide to the Birds of the Pacific
States.
Gulls:
Glaucous-winged Gull (Larus glaucescens Naumann)
California Gull ( L a w s cal ifornicus Lawrence )
Western Gull (Larus occidental is Audubon)
Ring-billed Gull (Larus del auarens i s Ord )
Herring Gull ( ~ a r u sargentatus smi thsonianus Coues)
Bonaparte Gull ( ~ a w phi
s 1adelphia [Ord] )

Terns :

Caspian Tern (Hydroprogne caspia imperator [Coues] )

Forster Tern (Sterna forsteri Nuttall)
Oyster catcher ( ~ a e m aopus
t
bachmani Audubon )
Curlews:
Long-billed Curlew (Numenius ameri canus Bechstein)
Hudsonian Curlew (Phaeopus hudsoni cus [Latham] )
Scoters:
White-winged Scoter (Nelanitta deglande [Bonaparte] )
Surf Scoter (Melanitta perspicillata [Linnaeus] )
Pelicans:
Brown Pelican (Pelicanus occidentalis Linnaeus )
Cormorants:
Double-crested (Farallon) Cormorant (Phalacrocorm auritus [Lesson])
Brandt 's Cormorant (Phalacrocoracc penicillatus [Brandt] )
Bairdls Cormorant (Phalacrocorm pelagicus Pallas)
i

Brant :

lacks st (Branta nigri cans [Lawrence] )

MAmALIA

Class MAMMALIA
Important in the pelagic communities are the whales, which we may
see blowing offshore. They are feeders on invertebrates, consuming
vast quantities of them. The smaller porpoises are predators on fishes.
Other fish feeders commonly noted on our trips are the Steller Sea
Lion (Eunetopias jubata) and the Harbor Seal ( ~ h o c arichardii).
Finally and most important, so far as the intertidal fauna is concerned, is man. His activities as clam digger, abalone hunter, and
collector of mussels, marine snails, and sea urchins have sadly impoverished the faunas. One has only to visit a biological preserve such as
Afio Nuevo Island, south of Pescadero, to realize the effect of man's
activities on the large gastropods. There the red abalone (Haliotis
rufescens ) is abundant and the black abalone (Haliot is cracherodi i )
forms solid layers overlapping under especially favorable ledges, even
well above the low-tide level, while family groups of the giant limpet
(Lottia) are seen on every favorable rock surface.

FIELD STUDIES
These studies are designed to give the student a firsthand howledge of
living invertebrate animals in their natural habitats, of their behavior and
interrelations, their adaptations as correlated to the various factors of the
environment, and the influence of physical conditions upon their local distribution.
Naturally such ,studiesare largely in the nature of field trips with an
all-too-brief period in the field during which those animals needed for further
study are collected and as complete observations and records as possible are
made of the ecological settings characteristic of the different animals seen, of
their distribution within the ecologically varying environment, and of the major
differences in this environment which seem to be responsible for the different
distributions.
The approach and spirit of these studies is, therefore, essentially ecological whereas the method can rarely, if ever, be the quantitative and experimental one characteristic of modern ecolo@;y. This is due in part to limitations
of training and equipment but essentially to lack of time.
It might be said that we make ecological surveys or reconnaisances and
seek to set up preliminary hypotheses as to the role of different factors in
determining the distribution of the animals, hypotheses based only on readily
observable and relatively obvious facts with regard to such features for example as degree of exposure as determined by vertical level of occurrence, by
the nature and arrangement of the substrate, by position with regard to wave
impact, wind, rain, sun, etc.
These field trips are interspersed with the laboratory studies according
to a logic of necessity. Groups are studied in the laboratory not according to
any taxonomic order but as best to prepare for the field trips as they occur.
The order of the field trips in turn is determined by various factors chiefly
those of season, weather and tide. Hence the schedule of events will differ
not only as between the spring semester and the intersession but also for each
from year to year. Examples of such schedules will be found in Appendix A.
The conditions of study vary greatly as between the spring semester when
the student is carrying other courses and the intersession when for six weeks
nothing else intervenes and only time and tides must be considered. These
courses (Zoology 112, and S112 and S119) will be considered separately therefore. First, however, come certain general instructions for field work.

GENERaL DIRECTIONS FOR FIELD W O E

nt needed: Geologist's pick (if possible); forceps; containers of
variA-ably
screw-topped); hard-backed notebook with automatic
pencil; if possible a magnifying glass (the magnifying glass and, if possible,
the forceps to be attached in some way); and some sort of bag or basket for
carrying equipment and specimens.
General Swtirestions: Plan your program beforehand, especially if at the
seashore, as every minute of low tide must be made to count. Go out at once to
the tide line. Follow the tide out. Collect back as the tide drives you in.
Take peat care not to be caught by the tide.
Methods of Collectiw: Do not collect unless you plan to use the specimens
for (1) study or identification, or (2) for your class or personal collection.
Do not collect great numbers of the same species.
Destroy as little as possible either of animal life or of the natural
conditions of the environment. Replace overturned stones to their original
position, since the fauna living on the under side or under the stone cannot
survive when exposed to air and wave action.

FIELD STUDIES

175

Field Notes: If specimens are taken merely for immediate study or practice
in identification, field notes are perhaps not necessary. Any specimen to go
into a permanent collection, however, should be accompanied by certain information to make it of value to any future student. Experience has shown that most
collections ultimately come to be used for study whether that was the original
intention of the collector or not. Many troublesome errors and difficulties
have arisen because of the lack of information with regard to specimens in such
collections as well as from accidents and carelessnesses in making and preseming field records of collections designed for study.
collection should be
made, therefore, which lacks the essential information with regard to the specimens in it, least of all a collection made in connection with a study of
natural history.
It is an exceedingly difficult thing to insure the correlation of specimens
and notes in general collecting where time is greatly limited by the tides and
organisms occur in profusion. There are two common ways of keeping these notes.
One, perhaps the safest, but by no means the easiest, or the most commonly used,
is to write the information in pencil and drop it into the container with the
specimen, or wrap it with the specimen if several are in one container. When
the collection is put into permanent form later the information thus obtained
must be copied on the label in (or on) the bottle and on the museum or collection record, whether card or journal. The second method is to drop into the
container or wrap with or attach to the specimen a number written at the time
or, better yet, already prepared, and, after the same number to give the desired
information in a field notebook or on cards used for that purpose. The last
method, while easier, involves the danger of loss of the notes, in which case
the number means nothing, and also the danger of mixing numbers and notes due
to haste in the field must be guarded against. Experience will dictate your
choice of methods. Your problem is to obtain essential information at the time
of collection and keep it in such a way as to insure that it will be correlated
with the particular specimen concerned and ultimately preserved as a permanent
record and in part, at least, placed with the specimen itself as a museum label.
There is almost no limit to the amount of available information which might
prove valuable to the student of distribution variation, or ecology if connected
with a specimen. Points commonly noted are (1) exact locality, including county,
(2) date, including hour if of any significance, (3) exact location (as regards
tide level, or altitude for example), (4)environmental situation (underside
of rock), (5) any organisms or organic remnant apparently associated with it
("with polychaete worms,no. 63, in hole of boring clam," for example), (6) anything unusual in the behavior or stage of the animal, and (7) the collector's
name. The absolute minimum must include locality, date, location and environmental situation, and name of collector.
The difficulty of collecting and keeping proper notes under the conditions
of wet hands, lack of place to sit, presence of rain or spray, lack of time,
difficulty of keeping specimens separate, must again be emphasized to make clear
the necessity of careful planning and discipline of the will to accomplish the
required end. Such notes will be required with the collections which are to be
turned in as a part of the required work of the course and for the various
ecological and observational studies. They can only be obtained by limiting
the objective, making it sharply definite, and refusing to be deflected from
it. For that reason the earlier trips will usually have more general objectives
and records will be confined to those having to do with ecologic features

A shell bag or fishing creel is convenient for carrying vials, bottles,

etc. A wise provision against loss by breakage on the rocks is to use tins to
hold vials and even bottles. Another danger is the losing of corks and the
consequent loss or mixing of specimens. See that corks are in tidtly or,
better yet, use screw-top vials and jars in so far as possible.
The most rewarding places for collecting are, (1)deep shaded tide pools
near low-tide level, (2) on and in seaweed, particularly in its holdfasts,

I
1

F,
,
k
i

176

LaBORATORY AND FIEID TEXT' I N INVWTEBRATE ZOOLOGY

titularly i f the
(3) i n the crevices and cavities of rock and within rock,
rock be soft, porous, o r covered with corallj rs algae, (4 under rocks, and
(5) i n sand and mud i f they are not subject t o constant movement by the tides.
Small rocks, especially favorable pieces from larger rocks, and holdfasts
of kelp or Laminariahfrom the outer low-tide level m y be brought i n and the
animala removed and cared f o r at leisure. This requires but the one s e t of
notes i n the f i e l d f o r many specimens. Also numerous specimens from the same
exact environment (a low-level t i d e pool, f o r example) may be placed i n a single
container with the proper notes and be sorted l a t e r . Care must be taken, however, t o segregate certain types of organims. Most worms, for example, secrete
mucue and tend t o form a tangled, often inextricable, mass with other organisms. This is particularly true of nemertean worn, which should be kept separately. Certain other animals, particularly crabs, tend t o t e a r others. Voracioua animals, such as large isopods, often devour smaller forma and the
smaller,
spines of echinoderms form a place of escape and entanglement f o r
mobile RnimAls. Intelligent care w i l l be necesaarg, therefore, i f satisfactory
collections are t o be obtained by putting numerous specimens i n the same container i n the field.

P"

ERVATION AND S.TUDY

On return to the laboratory the f i r s t move should be t o get material segregated and e i t h e r into fixative o r into dishes of clear seawater, taking care
always t o maintain the correlation between specimens and f i e l d notes. Ba a
routine it is worth while t o observe under the microscope, while still living,
any small animal, or developmental stage, not already studied, as this w i l l
often bring out much of beauty and interest otherwise lost. I f time permits
and there is any probability that your specimens are to be used f o r future
systematic studies, careful color notes w i l l be of value.
Well-known animals may be preserved at once or steps taken i n that direction. Others should be preserved when they have been studied and the proper
notation6 made. For methods of preservation see Billeting 39-42 of the United
States National Museum (bound together )
'

THE SPRING SEMESTER

(Zoology 112)
You w i l l be given a tentative schedule of exercises f o r the semester.
Fasten t h i s i n your syllabus and note on it a t the time such changes as may be
announced from time t o time.
You w i l l note that t h e exercises consist of a number of f i e l d t r i p s interspersed with laboratory exercises designed to give you a knowledge of the animals t o be studied i n the field. The f i e l d t r i p s constitute the heart of the
course. The dates of a number of these t r i p s w i l l be determined by t i d e s and
hence no regular schedule, as regards days of the week, is possible. These irregularities are unavoidable and have been cut t o a minimum. Please note them
a t once and make provision f o r them i n your program.
The f i e l d t r i p s usually undertaken a r e l i s t e d below i n the order i n which
they are considered i n the pages which follow. It w i l l be seen that they sample
three major environments, marine ( i n t e r t i d a l ) , f reahwater, and t e r r e s t r i a l , and
one, the estuarine, intermediate between the f i r s t two. This order while a
logical one cannot be followed chronologically because of seasonal requirements
and the tides. The schedule of exercises should be consulted f o r the chronological sequence.
These studies should be cunrulative and comparat ive Each new situation

FIELD S'IUDIFS

177

should be compared to the others and differences and similarities should be

noted as to environmental conditions, faunas, and their interrelationships.
FIELR TRIPS OF TBE SPRTNG SEMFSTER

A. Point Richmond in San Francisco Bay. Tide flats and a rocky beach in an en-

closed bay.

B. Halosaccion Flats, Moss Beach, San kteo County. A flat rock surface on the
open sea, exposed at intewdiate tides.
C. Dillon Beach, W i n County. A three-day trip permitting comparison of the
faunas of (1)an exposed rocky shore (Tomales Point ), (2) partially proteoted rocky shores (Bodega &y near Dillon Beach) and (3) protected tide
flats in an enclosed bay (head of Tomales Bay).
Estuarine

D. Iake Merritt, Oakland. A brackish-water, estuarine lake subject to great

annual shifts in salinity but not usually subject to tidal shifts of level.

E. Fruitvale Bridge, Alameda Estuary, Oakland. Fauna of under-bridge piling in

a tidal estuary.
Freshwater

F. Fauna of a freshwater pond or lake.

G. Fauna of some seasonal pond or pool.

Terrestrial

B. Strawberry Canyon, Berkeley Hills. Terrestrial habitats and faunas of a

wooded canyon.
DIRECTIONS FOR TBE FIELD TRIPS OF THE SPRING COURSE AND TEE LABOMTORY
EXERCISB I W I B E L Y PREPBRBTORY TO THEM

A. The Animals of Point Richmond. a Rocks Shore with Tidal Flats

in an Enclosed E ! q

..
ellrmnars Iaboratom Study.-This

study is a necessary preliminary to the

first field trip. A liat of the animals commonly encountered will be posted.
A study collection of these animals will be on display to enable you to get a
preliminary familiarity with their appearance and names. Your work in the field
will be to find and observe as many as possible of these animals in their natural habitats and associations. Several species of decapods form a conspicuous
part of this fauna. Specimens of these will be distributed through the class
and they should be identified by means of the accompanying key. Do the same
for such other specimens as are available. Keys follow to the decapods, isopods, amphipods, limpets (Acmaea) and annelids. These keys cover only the spec i e ~reported from Point Richmond; consult the index for more complete keys
for these animal groups found in the preceding section.
Field Trin to hint Richmond
h w a ~ :(1) To learn the various animals of this region as they exist
under natural conditions, i.e., each in its normal ecologic setting ("nichen);
(2) to determine their distribution with regard to physical and biotic factors;

178

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

(3) to learn their groupings by zones or other environmental situations; and

(4)to obtain as clear an idea as possible of their interrelationships in such
groupings, especially their food relations.
Procedure: (1)W e field notes with diagrams or sketches. (2) Collect
such animals as are necessarg to identify species using the keys which follow
and where needed those in the preceding section.
A second trip will probably be necessary to complete these studies for
those who have no previous knowledge of the fauna.
Re~ort:The report, if any, will be in the form of a synthetic vertical
section of the shore at Point Richmond showing the relative location of all
important habitats or associations, accompanied by lists of the species found
in each of these different habitats or associations. A discussion of the
reasons for the distribution of the different types is optional.
If this trip follows the one to Halosaccion Falts (see below) it should
raise many questions as to the great differences in the faunas of the two regions.

A Sam~leSet of Questions on the Point Richmond Fauna

'

(1) Name three major factors which probably account for the sparse nature
of the fauna. (2) Which large and important phylum of animals was not found
represented at all? (3) What conditions probably account for their absence?
(4)Where were the oysters largest and most abundant? (5) What differences in
physical conditions were correlated with this abundance? (6) PFhat predator was
confined to the area of the abundance of oysters? (7) What could be shown to
be the usual explanation of the occurrence of shells of gastropods at intertidal levels where the living species were not found? (8)What is the most
important predator (or scavenger of freshly killed animals) in each of the
following situations: (a) under stones fitting closely against the substrate?
(b) in the small, high, rock-bottomed tide pools during low tide? (c) over the
higher rock surfaces during high tide? (9)List the animals found in the zone of
pebbly beach. (10) Which if any of these occurred in the sandy beach-a few
inches away? (11)How do you explain this great difference? (12) Name a different amphipod characteristic of each of the following locations: (a) hightide pools in rock; (b) under surface of smooth stones resting on substrate,
low in tidal zone; (c) felted mass of algae over surface of rock outcrop in
mussel zone; (d) algal debris at or near high-tide line. (13) What biotic
reason might you give for the absence of relatively large predators such as
the larger sea anemones and starfishes both of which are present at the yacht
harbor? (14)What two physical conditions may also be concerned? (15) Which
two species are diagnostic of the higher splash zone? (16) Which two of the
lower splash zone or the highest tide line? (17) What three vicissitudes must
be met by animals living in small high-tide pools? (18)What is the present
status in the mussel-bed succession? (19)Judging from the situations in which
you found them name as many as you can of the more than a dozen species whose
numbers would probably be greatly increased if the mussel beds were fully
developed. (20) On a line, representing the beach line from the middle of the
pebbly beach (small stones between mud flats and sand beach above) to the fixed
boulders at the point, indicate by brackets where you would expect to find each
of the following: (a) Hemigrapsus oregonensis; (b) Hemigrapsus nudus; \C) Thais
lamellosa; (d) chitons. (21) Choose four species whose niches seem best defined
by our study and attempt to state the factors involved. (22) Which one species
had the widest intertidal range? What makes possible this wide range? (23) Which
species common here are absent at Halosaccion Flats? Attempt to explain the absence of each species.

FIELD STUDIES

179

A KEX TO TIBE DECAPODA OF POINT RICDIOND

1. Antennae long (generally a t l e a s t a s long a s carapace); body elongated,
2
not flattened
1. Antennae short (much l e s s than half the length of carapace); body short
5
and broad, g r e a t l y f l a t t e n e d (True Crabs)
2. Abdomen twisted, carried i n a mollusk s h e l l
Hermit Crabs, Pagurus spp. ( f i g s . 97, 98)
2. Abdomen symmetrical, exposed
3
3. Abdomen not narrowed behind; one cheliped of male very large; l i v i n g i n
Ghost Shrimp, Call ianassa ( f i g . 96)
burrows i n mud o r muddy sand
3. Abdomen narrowed behind a t base of t a i l fan; chelipeds a l l slender; not
True Shrimps 4
burrowing
4. Small ( l e s s than one and one-half inches long); rostrum long, slender
Spirontocaris paludicola ( f i g s . 85, 87, D)
4. Larger (one and one-half t o two inches); rostrum short, dorsally f l a t tened; a c i r c u l a r black area on e i t h e r side of 6th abdominal segCrago nigromaculata
ment
(Other species of Spi$ontocaris and Crago may possibly
be found. See Key, p. 104.)
6
5. Body nearly square
8
5. Body more or l e s s oval, much broader than long

.............................
...............

................

......................

....

........................
................
........................

...........................
.............

6. Back with numerous transverse l i n e s

Striped Shore Crab, Pachygrapsus crassipes
6. No l i n e s on back
7
7. Tips of legs without hairs; body usually purple o r dark red
Purple Shore Crab, Hemigrapsus w d u s
7. Tips of legs hairy; body usually yellowish-green
h d Crab, Hemigrapsus oregonensis

...............
............................
..................

....................
.............
.................

8. Margin between eyes (the front ) extending forward (produced), with f i v e

nearly equal t e e t h
Red Crab, Cancer productus
9
8. Fmnt not produced, f i v e t e e t h unequal
9. Hairy; with red spots on ventral surface; antennae conspicuous, a s long
Rock Crab, Cancer antennarius
a s width of front
9. Smooth; not spotted below; antennae very short
Edible Crab, Cancermagister

.............

......................

A KEY TO TEE ISOPODA OF POINT RICHJilOND

1. Uropods terminal ( f i g . 61); largely above high-tide mark

...........................
..
.
...............
..........................
......................
......................
..............................
...........................

Ligia occidentalis
2
1. Uropods l a t e r a l ( f i g . 58); or ventral ( f i g . 54); not above high t i d e
2. Uropods valve -like, inf lexed ( f i g 54); pleopods largely branchial;
.Pent idotea (fig. 56)
color green o r brown
2. Uropods forming with terminal segment of abdomen a caudal fan ( f i g . 58);
3
color gray o r white
3. Abdomen usually of 6 segments
5
3. Abdomen usually of 2 segments
4
4. Smaller; caudal fan spreading ( f i g . 58,A); surface not densely granular;
fiosphaeroma
4. larger; caudal fan not noticeably spreading; surface densely granular
.Sphaeroma pentodon

180

LABOMTORY AIVD FIELD 'JEXT I N INVEFtTEBFLATE ZOOLOGY

5. Size moderately large; color speckled or dark gray; not boring i n

wood
Ci rolana (fig. 58,C-E)
5. Size minute; body white, eyes black; boring i n wood
~imnoria

......................
A

.......

TO !lTB AdhPHIPODA OF POINT RICHMIND

P" . . . . . . . .. .. .. .. .. .. .. .

1. Body l a t e r a l l y compressed as usual i n am ipods; prominent side

plates covering bases of legs (fig. 63
2
1. Body not l a t e r a l l y compressed (fig. 70); no side plates
.10
2. Antennule (fig. 63) very short, only a s long as f i r s t two segments of
peduncle of antenna; habitat usually above high-tide mark; large
Orchestiatraskiana
2. Antennule at l e a s t a s long a s e n t i r e peduncle of antenna; habitat
intertidalorlower.....................
3
3. Third uropod (fig. 63) extending posteriorly beyond second uropod
,
4
(study under microscope); antennule with an accessory flagellum
3. Third uropod not extending beyond second; antennule without accessory
flagellum..............................
6
4. m e round o r irregularly round; outer ramus of the t h i r d uropod seve r a l times a s long a s the tiny inner ramus
Me1 ita sp.
4. Eye bean-shaped; outer ramus of t h i r d uropod not more than three
times a s long a s inner ramus
5
5. Inner ramus of third uropod at l e a s t two-thirds as long a s outer ramus
Ganu~alvspri bilofensis
5. Inner ramus of t h i r d uropod l e s s than one-half as long as outer ramus
Gbmarus confervicolus

..........................

....

. .

.........
....................
.........................
.........................

6. !Third uropod with two prominent upward-pointing h o o k a t the t i p of

the outer m s ; color i n l i f e green o r gray stippled with small
brownspots.
Ampithoevalida
6. No such hooks on t h i r d uropod
7

. . . . . . . .. .. .. ............. .. .. .. .. .. . . . . . . .
7. Antennule longer than antenna . . . . . . . . . . Neopleustes pugettensis
7.Antennule shorterthanantenna . . . . . . . . . . . . . . . . . . . . 8

8. Antenna with dense t u f t s of b r i s t l e s ; antennule two-thinb t o threeAllorohestes plumulosus

fourths a s long a s antenna
8. Bristles of antenna fewer, not i n t u f t s ; antennule l i t t l e more than
half the length of antenna
9
9. Color black and gray, sometimes with transverse white s t r i p e s ; conHyccle pugettensis
spicuous i n high-tide pools
9. Color pale; size smll; not common
Allorchestes angustus
10. Body somewhat flattened (fig. 70); living i n tubes
Corophium insidiosum
10. Body very slender, abdomen reduced t o stump; appearance very unlike
Caprella sp. (Plate XI, no. 5 )
that of other amphipods

..........

.....................
. . . .. .. .. .. .. .. .. .. . .

.........

A KEP TO THE SPECIES OF ACdWFA OF POINT RICEWND

1. Apex near one end of s h e l l ( r e f e r t o Plates XII-XV)
2
1. Apex a t least one-third of length from end
4
2. With well-developed r i b s (radiating ridges on s h e l l ) and irregular mrgin
3
2. With fine s t r i a t i o n s and e n t i r e margin
The Speckled Limpet (rare ), A. personu

. . . . . .. .. .. .. .. .. .. .. .. ..
.................................
.................

FIELD STUDIES

181

3. Ribs close together, rough; margin semte; inner, central area por-

.........
......................
. . . . . . .. .. .. .. .. . . . . . . . . . . . . .
.......................
..................

The Rough Limpet, A. scabra

celaneous, white or mottled
3. Ribs well separated, rounded; margin crenate; inner area shiny, with
one central brown spot; apex usually overhanging
The kkLimpet,A. digitalis
4. Elevated; profile with straight sides
5
The Plate Limpet, A. scutum
4. Verg low; profile convex
5. Shell heavy; inside with a considerable white area
The Shield Limpet, A. pel ta
5 . Shell thinner; black externally and brown internally
The Owl Limpet (rare), A. fenestrata
A m T O m ~ D A O F m m R I ~ N D
(hfer to plates, figures, and explanations of terms under
Phylum belida in preceding section)

1. Non-segmented; tough-skinned, finely tuberculated; with an introvert

terminated by dentritic branchiae (fig. 13,E); in rock crevices.
SIPUNCULOIDEA
Physcosoma
1. Segmented worms, with parapodia (figs. 13.8 and 14) more or less
developed and variously modified; often with accessory head and
body appendages. POLYCHAFTB
2
2. Body dorsally covered with overlapping scales (fig. 17,B). POLYNOIDAE
Pol$"noids
2. Body without scales
3
3. Head completely covered by feathery tentacles (fig. 16); body in a
caloareous tube. SERPULIDAE
Serpulida
3. Head completely covered by a conspicuous spiny operculum (fig. 1 9 ) ~
Sabellarids
body in a tube which is not calcareous. ,WlEIUUIDBE
4
3. Head otherwise; body generally not in a tube
4. Anterior end of body with long filamentous outgrowths, usually extending
Cirratulids
forward over head. CIRRA'IITLIDAE (fig. 17,H)
4. Filamentous outgrowths, if present, not concentrated near anterior end
5
5. Prostomium bearing tentacles or palpi and often accompanied by peristomial cirri (fig. 15)
6
5. Prostomium plain, without tentacles or palpi, and not accompanied by
peristomial cirri
.10
7
6. Prostomiurn with conspicuous tentacles or palpi
6. Prostomium with minute, inconspicuous tentacles (fig. 17,F); proboscis
actively everted when disturbed
9
7. Prostomium with tentacles, but without palpi; small, very active,
Eteone (fig. 28,~)
mud-flat dwellers. -DOCIN
(fig. 17,E)
7. Prostomium with a pair of long, prehensile, tentacular palpi (fig. 17,s)

........................

......................
................................
...........................

.................
. . . . . .. .. .. .. . . . . .
.........
.

..........................
..........................
.............
....................
....
SPIONIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . .Boccardia
. . . . . . . .8

7. Prostomiurn with a pair of thick, fleshy, palpi (fig. 15)

8. Parapodia distinctly biramous: prostomium with only paired antennae:
Nereids
dorsal cirri never articulated. NEREIDLE
8. Parapodia uniramous; prostomium with a median antenna in addition to
paired antennae; dorsal cirri often articulated (fig. 17,D); mostly
minute. SYLLIDAE
Syllids
9. Prostomium angular (fi 27,A): recurved cirri between branches of
parapodia (fig. 27,Bj; pearl-gray in life; in sand. IEFVITIDBB N e ~ ~ h t y s
9. Prostomium pointed,
parapodia tiny, without recurved cirri.
GLYCERIM
Glycerids (fig. 17,F)

............

........................

......

............

182

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

10. Most of dorsum coversd by interlacing, cirriform branchiae; prostomium shovel-like. ORBINIIDAE
.
. Nainereis (fig. 25,A)
10. Without branchiae; prostomium pointed or rounded
. .
11
11. Parapodia reduced; bluish-red in life; in mud flats. CAPITELLIDAE
...... .. ...... .
... .
Capitellids
11, Parapodia more or less normal; orange to reddish-metallic in life; in
Lumbrinereids (fig. 17,A)
sands or sandy mud. LUbBRINEREIDAE

. .... .. .
... .. ....
. ....
.. ..
.....

B. The Animals of Balosaccion Flats. a Horizontal Rock bosure at About

Mid-Tide Level. on the O ~ e nCoast. bbss Beach. San bteo Counts
Location: The name Halosaccion Flats is our own, given because of the
presence of the peculiar, finger-like, gas-inflated green abae in great numbers near the center of the rocky flat which lies (see Map I) just south of
Moss Beach Point and immediately north of the lagoon-like cnve we have designated Seal Cove.
Ph~sicalConditions: These have been largely stated: A rocky substrate of
fairly hard slaty shale, with relatively few crannies but cut at intervals by
narrow crevices which are tidal channels at intermediate tides but contain
long, narrow tide pools at low tides, often containing accumulations of sandy
gravel. The precipitous seaward face of the rock, rising about two feet above
low-tide level, receives the full force of the waves and is cut at intervals
by tidal channels, most of which do not penetrate far. Shoreward at the northern end the reef rises to near high-tide level. Shoreward also are protected
channels and tide pools with a sloping rocky or sandy beach rising to the
sheer cliff.
Physical habitats of diverse types are found, therefore. The factor of exposure, varying primarily with vertical height with regard to the low-tide
line, is variously affected by the nature and arrangement of the substrate,
Thus we have tide pools; shallow, gravel-filled crevice pools; deep protected
tidal channels; outer exposed tidal channels; ledges; exposed rock surfaces
with varying relations to wave impact, etc.
Soecial Biotic Features: (1) Living seaweed forming a close growth over
most of the horizontal rock surface, (2) close-set aggregates of a sea anemone
(Bunodactis elegantissima) which cover themselves with pieces of shell and
gravel, (3) a Nytilus-Mitella zone on the high northern shoulder.
Pur~osesof Study: Familiarity with members of fauna and their names;
experience in the observation of animals in the field in terms of the effects
of ecological factors on the distribution and success of species and on the
faunal associations.
Procedures: If this is the first field trip it will be preceded by a
labomtory study of the animals most commonly to be encountered. Go out to
the sea-front to take advantage of the low tide. Study the animals at the
lowest tidal exposure first Look under ledges. Break off overhanging ledges
to get a more intimate knowledge of the dwellers on the surfaoe and in the
crannies and burrows. Distinguish between borers, nestlers, and cranny dwellers. Work back across the horizontal surface, noting differences of faunas,
if any, (1) between the seaward and landward sides; (2) correlated with different algal ty-pes; 3) of the narrow crevice pools; (4)of the gravel-bottomed
high-crevice pools; 5) of protected inner tidal channels and tide pools; (6) of
the higher northern shoulder; (7) of the rocks between the reef and the beach
slope; (8)of the beach slope; (9) of the crevices at the foot of the cliff.
If this follows the Point Richmond trip, differences between the two
faunas should be noted and an attempt made to explain these. Some of the differences are those to be expected between bay and open shore faunas. Others
are due to special conditions in the two situations. Attempt to distinguish
between these two types of differences.

7 he ~ a t Tub
h

Pacific Ocean

Iididaea c h a n n e l

Map 11. Mass Beach shore area (loaal names)

FIELD STUDIES

185

Sample questions:
1. What phylum well represented here was absent at Point Richmond?
2. Differences in what two important ecologic factors might be thought to explain this difference?
3. Which two phyla were represented most extensively in the fauna of the horizontal algae-covered flat?
4. Can you give any reason for the absence of large individuals and species
here?
5 . Contrast the assemblages found in crevice pools and surface growth. What
explains the differences?
6. Compare the limpets (Acmuea) of the flat and of the inner shore rocks and
high shoulder. How would you explain this situation?
7. Attempt to explain the occurrence of the patches of Bunodnctis. Why not more
extensive?
8. What seems to restrict the occurrence of Anthopleura in this area?
9. Describe the home habits of Lophopanopeus in the larger, more shoreward
crevice pools.
C. Com~arisonof Faunas of b o s e d Rocks Shore (Tomales Point 1,
Partially Protected Rocb Shore (Bode~aBay) and Protected
Tidal Mud and Sand Flats (Tomales'Bay

Location: The class will have its laboratories and living quarters at
Dillon Beach, &rin County, California (see Map IV) on the shores of Bodega
Bay near the mouth of Tomales Bay.
Pumoses: This will be by far the most important field trip of the course.
It will usually come relatively late in the semester and enable the student to
focus the knowledge gained from the preceding field trips and laboratory studies
in an intensive study of three faunal assemblages not far apart spatially but
ecologically distinct. The opportunity given by continuous study for the three
collected by all, thus giving
days makes it possible to identify the &owns
a relatively complete picture of these rich faunas, and to compare notes and
impressions as to ecological features and relationships.
Procedure: One morning the class will visit an area of tumbled rocks whose
bases are exposed at low tide at a point known as "The Second Sled Road" (see
Nap V) along the shore of Bodega Bag north of Dillon Beach. Here a relatively
rich and diversified fauna will be found in tide pools, on rock surfaces, among
algae, in gravel and sand about roots of eelgrass, etc. This will suggest in
some ways the fauna of Helosaccion Flats. Differences from it should be noted
and explanations for these differences sought. Again it should be compared with
that to be seen at Tomales Point a more exposed region where rocky formations
abound with deep tidal channels, tide pools, etc.
Another morning will be devoted to the fauna of the tidal channels, tide
pools, and rocks of Tomales Point. Among other things make certain that you
investigate fully the association of animals in a well-developed mussel bed
such as will be found here on the higher rocks. Be able to explain the role of
each type of animal in the association or what advantages it receives by the
association.
Finally one morning will be given to the faunas of animals to be found at
various levels of the tidal flats of muddy sand just inside the mouth of Tomales
Bay (map V). Here the numbers of species are not so great as in the faunas so
far studied but there is a wealth of individuals. Particularly important here
are the giant clam, Schiaothaerus, the bent-nosed clam, Macoma nasuta, Phoronopsis sp., whose pale-green lophophores practically cover the ground for several acres, polychaete worms of several interesting types especially Clymenella,
and capitellids. At certain levels in a restricted area the mud shrimp, Upogebia, will be found, while Callianassa occurs in great numbers at somewhat

F R E N C H M A N ' S

N
0

60

100

150

200

FROM BASE OF CLIFF

-0

H
H

SECrlON 6-6' (DIP _

.-

::

tl

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

188

higher levels. In the time available get as clear a picture as possible of

these major types in their natural setting, and the other species which will
be encountered, and of the varying ecologic ranges of the different species,
seeking to assign environmental causes for the limitations of these ranges.
In identifying species use the keys to the various groups as listed in
the index. Report any species not listed in the syllabus and turn these in with
information as to place of collection.
D. lake krritt: A Brackish Water Estuarine Iake
Lake Merritt has been formed from a branch of Alameda Estuary. Its level
is controlled by tide gates and hence the factor of the tides is eliminated.
During the rainy months the salinity is very low. During the summer and fall
it reaches a concentration 30 parts per 1000) not far below that normal for
seawater (34 parts per 1000 An abundant algal growth, chiefly green algae
and diatoms, furnishes a rich food supply. Aa a result a few species, adjusted
to low and varying salinity and not adapted to withtand exposure or wave action, flourish and occur in great numbers except during periods of high salinity.
Such species are the tube worm Mercierella enigmatica, the associated
amphipods Corophium bonellii and G'umrus confervicolus, the polychaete, Neanthes
succinea, the crab, Rhithropanopeus harrisii, all of which live among the
masses of tubes of Mercierella, the endoproct bryozoan, Barentsia (= Pedicellina), common as a mossy growth over the tube-worm masses, especially hear the
exit of the lake, and the horse mussel, Modiolus demissus.
Several other species, while typically marine, invade the lake at times
of high salinity but maintain a precarious existence during the remainder of
the year. Such-arethe soft-shell clam, Mya, the barnacles, the encmsting
bryozoan, Membranipora, the mud snail, Ilyanassa obsoleta, the oyster drill,
Urosalpinx, the tunicate, Ciona intestinalis, and various other species.
Procedure: Study the fauna to be found on the sides of the pier of the
Municipal Boathouse. See a boat lifted in the shops near by and observe the
enormous numbers of gammarids leaving it as it leaves the water and the thick
encrustation of Mercierella tubes. Observe Modiolus in life in the mud near
the inner end of the pier and the young Mya just w e ~ tof the boathouse. Can
you distinguish as to niche, between Nereis and Neanthes.
Tows will be made which should be studied on return to the laboratory.
These will probably contain very numerous larvae of Mercierella in various stages
of development, trochophores, polytrochs, and young worms. Also veliger larvae,
probably of Nya, possibly also of Modiolus, m y be found in great numbers. In late
spring or summer entomostraca, especially copepods, are present in great numbers,
as also mtifers.
Mud from the bottom will show great numbers of worms, chiefly of the genus

\.

Streblospio.

Break apart one of the tube-worm masses and study the relations between
the different animals in this association.
Be responsible for a howledge of this fauna, the food habits of all its
elements, and the chief physical and biotic features responsible for the
abundance of each.
Keys for the identification of amphipods and polychaete~are given below.
Identifications of other groups must be made by the general keys.

I
I

A KEX TO TEE AMPEIIPODS OF LggE MERRIW BM) OAKLAND ESTUARY

............
. . . . . . . . . . . . . . . . . . . . . .. .. .

2
1. Side plates typical (fig. 63), body compressed
1. Side plates 2-4 fused; a small, broad species common among hydroids in
Met opa
Oakland Estuary
1. Side plates reduced; dorsoventrally flattened; tube-builders
Corophium

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I,ABORATORY AND FIELD !nXT IN INVWTEBRBltf& ZOOLOGY

2. Outer branch of uropod 3 broadened and much longer than that of uropods
1 and 2 (Family Gammaridae)
.3
2. Outer branch of uropod 3 not broadened and not longer than that of uropods 1 and 2; in moist but not in wet places (Family Talitridae)
Hyallela
3. Antenna 1 at least one-half as long as body; eyes not crescent-ahaped;
dorsal groups of spinules only on fifth pleon segment; Lake MemittMelita
3. Antenna 1 not one-half as long as body; eyes crescent-shaped; pleon
segments 4-6 with dorsal groups of spinules; Lake Merritt
G a m r u s con.fervicolus

.....................

................................

.................................
.........................

1. In closely packed, calcareous tubes; head covered by feathery tentacles

Merci erella enigmat ica
(fig. 16); operculum spiny
1.Notso
2
2. Larger; head with a pair of fleshy palpi, 2 small anterior antennae and 4
pairs of peristomial cirri (fig. 15)
3
4
2. Smaller; head with 2 long, prehensile palpi or without palpi
3. First few segments provided dorsolaterally with oblique wrinkles; posterior notopodia with only pointed setae
Neanthes succinea
3. First few segments mooth dorsally, without oblique wrinkles; posterior
Nereis vexillosa
parapodia with some blunt setae
4. Anterior end of prostomium with 4 small antennae; palpi absent (figs.

. . . . . . . . . . . .. .. .. .. .. .. .. .. .. .. .. . . . . . . . . . . .
. . . . . . . . . . . .. .. .. .. .. ..
........
............

% , B * C ) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

4. Anterior end of prostomium without antennae; posterior margin provided

5
with conspicuous prehensile palpi (fig. 17,J)
5. Prehensile palpi accompanied by a pair of foliaceous appendages; a dorsal collar on segment 3; in muddy tubes
Streblospio lutincola (fig. 34,A)
5. Prehensile palpi not accompanied by foliaceous appendages; without dorsalcollar
6
6. Pale white; minute; constructing erect tubes which project from crevices; dorsal branchiae not present anterior to segment seven
Polydora
6. Salmon to red in life; inhabiting burrows constructed in clay banks or
in crevices; dorsal branchiae present on segments 3 to 5 as well as
Boccardia (17,J )
more posteriorly
7. Slender, thread-like; yellowish or somewhat green; fLrst segment wellEteone californica
developed dorsally
7. Depressed, short; pale or white; first segment dorsally reduced

.............

....................
..............................
.

...................
.................
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Eteone lighti
E. Pile Communities and Zonation a8 Seen at Fruitvale &i&e

laboratory Prewration: A group of invertebrates which is an important

part of the Fruitvale Bridge fauna but which has not been previously encountered in any abundance in this course is the hydroids. The Hydrozoa should
therefore be studied in the laboratory not long before the field trip.
Field Procedure: Collectione and notes must be made from boats. The animals taken will be studied and identified in the laboratory on the same day.
hkterials needed are several screw-top jars, several vials for individual specimens, forceps, scalpel.
Observe and record the vertical and horizontal distribution of the animals

FIELD STUDIES

1
!

191

found. What factor usually determines vertical distribution? You will find
some types here at higher levels than would ordinarily be expected. k t are
these types? Do you find any difference in this regard between central and
marginal piles? W a t is the factor thus modifying the action of the factor
which usually determines vertical distribution?
Note especially the hydroids and the anemones. Attempt to distinguish the
different genera of hydroids and the different species of anemones in the field.
Collect specimens to enable you to check your field determinations. Be able to
place each in its characteristic zone. barn to see the highly modified amphipods (Caprellidea) which are abundant on the hydroids. Study these in the laboratory. Are the caprellids different on different genera of hydroids? Were do
you find nudibranchs and their coiled egg masses?
Some of the piles will be covered with muddy debris. This is made up of
tubes of great numbers of small amphipods. Idmy species of animals seek shelter
in and about the mud tubes. Collect some for later study. Does the location of
these piles give you any clue as to why they, and not others, bear this association?
Select a pile which shows zonation especially clearly. Working in teams of
three or four, study the zonation carefully, designating the various zones,
measuring their extent and relation to low tide level. Keep notes on the animals found to be characteristic of each zone. Seek to determine the factors, physical and biotic, influencing .the zonation.
As time allows, stop at other neighboring piles. Compare them with your
first pile. Study -the piles on the other side of the estuary. Account for differences in the fauna of the different piles.

F. A Permanent Freshwater Pond or Iake

The laboratory preparation for this trip requires introductory studies of
some of the invertebrate groups whose members are largely inhabitants of fresh
water. Such are the Rotifera, the Cladocera, the Ostracoda, and the Copepoda.
Pumoses: To get a comprehensive picture of the fauna, food chains, and
food cycle of a freshwater pond or lake.
.Procedure: Collect as completely as poesible from various habitats in the
pond. Especially important are (1) floating plants; (2) the surface of floating
wood, etc. ; 3 ) under the bark of floating branches and logs; (4)the fine roots
of trees; (5 plants growing in the water at various levels; (6) the bottom
mud. Tows will be made near the surface, some inches below, and near the bottom. Mud will be taken from the bottom for making cultures.
Identify your collections, using the keys in the preceding section. Sample
the fauna of the tow and the bottom mud and identify the animals found there.
Insects need not be identified beyond the family or order groups, thus,
belostomid bugs, dytiscid beetles, damsel-fly larvae, dragon-fly larvae, chironomid larvae, phantom larvae (Corethra). Use Needhem in Ward and Whipple, or
A Guide to the Study of Freshwater Bioloa (1930) by Needham and Needhem, and
works dealing with individual groups which are listed in the bibliography.
The common, small, somewhat transparent leeches belong to the genus Glossiphonia and are predators on the snails rather than bloodsuckers as are typical leeches exemplified by the larger species (Genus ~ i r u d o ) .
Consider carefully the food habits of each species or group found. Fit
them into food chains and these into a food cycle for the pond. Put the food
cycle in the form of a diagram. This is to be turned in for criticism and
evaluation.

Some Animals of a Permanent Pond Listed According to Food Habits

Plankton Feeders: Protozoa, sponges, rotifers, Cladocera, Bryozoa, Musculium
Copepoda.

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LABORATORY AND FIELD TEXT IN IJWERTEBRBTE ZOOLOGY

Predators: Hydra, hrgesia (= planaria), leeches, water mites, spiders, dragonfly larvae, water bugs (back-swimmers, water boatmen, ~elostomatidae),
water-striders (Gerridae), beetles and their larvae (Dytiscidae, Hydrophilidae), ghost larvae (Corethra), fishes, Amphibia, reptiles, birds.
Plant Feeders: Isopoda, snails, aphids, amphibian larvae, some fishes.
Plant Scavewers (including detritus feeders): nematodes, oligochaetes, Ostracoda, Isopoda, Amphipoda, may-fly nymphs bloodwonna (larval Chironomidae),
aosquito larvae, gnat l a m e (~inruliidae).
Animal Scavewers: Ostracoda, Isopoda, Amphipoda, water-striders (Gerridae),
Gyrinidae.

Life in all small bodies of water is subject to relatively rapid seasonal

changes due to changes in temperature and other climatic factors. The life of
a permanent pond or lake undergoes also a vastly slower series of changes,
known as geoglogical succession owing to the fact that physical conditions
change slowly, each body of water tending to become more and more sedimented
and ultimately passing through a marsh or a bog stage to become a meadow.
The seasonal pond undergoes each year profound changes in its life since
it is not only subject to the influence of climatic factors but also a change
from dry land to pond and from that back to dry land through marshy stages.
Only those organisms will be found in it which have a resistant stage to carry
them over the dry period or such as may be introduced either by visiting animals or by wind-borne resting stages.
In such ponds the shifts of population are very rapid. First a short
period of rapid development of plankton which furnishes the food basis for a
rapidly pgramiding population of plankton feeders (Protozoa, phyllopods,
Cladocera, copepods, rotifers) which develop in enormous numbers followed somewhat later by a horde of predators and scavengers. The predators are largely
insects, waterbugs, and water beetles, larvae and adults. Important aa ecavengers at this stage are the ostracods, which oftentimes occur .in very great
numbers in such ponds.
To get the true picture of the seasonal mcce~sionof life in such a pond
would require a number of visits beginning early in the spring. Ve shall designate the pond to be studied, suggest the time for two earlier visits, and
set aside a day for a visit by the class when the population can be thoroughly
sampled.

H. Terrestrial Habitats of Strawberry Canyon

1
I

The laboratory preparation for this trip will consist of a study of invertebrate groups which are primarily terrestrial, chiefly arthropods: centipedes, millipedes, arachnids, etc. Many insects will be found. They should be
placed to the order and their ecologic position with respect to other invertebrates should be noted wherever possible.
In the field investigate all habitats, espeoiallg (1) under logs, (2)
under dead bark, (3) in decaying wood, (4) in damp surface debris, (5) under
stones, (6) in debris and under loose stones in th creek bed. The aquatic
habitats of the stream should be noted and compared with other freshwater
habitats studied.
Collect animals of whose systematic position you are uncertain. Keep records of others. Keep in mind the food relations of the various species seen.
Identify the slugs and isopods using the keys listed in the index.
No report will be called for but you will be held responsible for the
knowledge thus gained.

FIELD S!lUDIES

193

!JIB SPECIAL PROBLEM

Each student will choose a special problem for careful study and report.
This problem will deal with the classification, the anatomy, the development,
the behavior, the distribution, or the ecology of some local animal or group of
animals, or with the fauna of a certain circumscribed area.
The problem should be chosen and work initiated early in the course. A
certain part of the laboratory time will be allotted to it. It is to be carried
out and reported on as a research problem. &erg. effort should be made to make
it as scientific as possible. All statements should be supported by the author's
accurately presented findings or by definite reference to authority. Reference
should be made to sources found to be useful such as previous reports on the
same subject, general treatises and especially articles in scientific journals.
Care should be taken to limit the scope of the problem so as to make this possible.
The following is suggested as a standard form for these reports:
Detailed presentation of results
Brief table of contents
Discussion of results (if needed)
Statement of problem
Summary
Brief surveyof literatureandprevious reports
Bibliography
Materials and methods
General findings, including lists of
species, keys, etc.
Illustrations ahould be placed in the body of the paper where they will
be of most value. Both Zoolo~icalRecord and J3iolonical Abstracts may be used
in making up the bibliography. The standard bibli~gra~hihal
forms should be
used, as in the example below, and references made by name and year, thus
Oldroyd (1924):
Oldroyd, I. S.
1924. The marine shells of the west coast of North America.
Stanford Univ. Publ. Geol. Sci., 1: 1-247, pls. 1-47.
A rough draft, or at least an outline, should be turned in for criticism
and suggestions as early as possible, not later than two weeks before the last
laboratory period during the regular session or one week during the summer session.
The finished reports will be held for use of future students in the
course. Duplicates should be made if the student desires to retain a copy.
Carefully labeled examples of the species dealt with are to be turned in with
each report.

THE SUMMER COURSE

(Zoology S112, S119 )
In alternate years the course is given at the seashore during the six
weeks of the Intersession, usually from near the middle of k y to near the end
of June. S112 is given at this time for undergraduates or graduates seeking a
teaching certificate in zoology or biology and for other graduates who, for
good reason, do not wish to take S119.
Candidates for the Ph.D. degree in the department of Zoology of the University of California at Berkeley may take S119 in fulfillment of the requirement in k i n e Zoology. The subject matter will be much the same as that in
S112 but greater proficiency will be expected of these students, their examinations will be fewer and separate and they will do much more intensive problem work.
The summer course has been given at two different stations, Moss Beach or
Montara, San Mateo County (Map I), and Dillon Beach, k i n County (Map IV).

194

LABORATORY AND FIELD TEXT IN INVXFtTEBFUTE ZOOLOGY

Each location has its advantages. Facilities for board, room, and laboratory
usually decide which shall be chosen in any given year. Directions are given
for each, therefore.
Board and room for the six weeks usually ranges from a m i n i m of from
55.00 where a number of students share a room to a m a x i m of from
75.00 where a single room is desired. This includes the use of the
laboratory and at Dillon Beach transportation across the bay on two or three
trips and to the tide flats on two or three trips.
As noted above, undergraduates and those graduates not taking the course
to fulfil the marine zoology requirements for the doctorate yill take 5112.
This has a unit value of 3 but the amount and nature of the work done during
the six weeks is the equivalent of a five-unit course and students desiring the
additional two units may enroll for two units of 5119 (without additional laboratory fee).
Those desiring six units or those who have taken Zoology 112 may enroll
for problem work (S199 or S224), the number of units to be arranged with the
instructor, who is enabled to act for the registrar and the comptroller.
The course giyen at the beaches, has the same general aims as the regular
course. h c h of the general discussion in the early part of the syllabus, as
well as the laboratory directions, will be applicable to it. The program of the
summer course will naturally be very different because of the short period of
time involved.
The detailed program will vary from year to year with the incidence of the
tides, but the projects will remain much the same.
The first necessity is to learn the fauna, its distribution, in general,
and the major physical features which determine this distrivtion. Following
this, problems of a more definite nature will be attempted. Since advantage
mst be taken of all low tides and since the lowest tides usually occur during
the first or the second week of the courae, these will necessarily be especially strenuous, the very early morning being devoted to field study and collecting and the remainder of the day to the identification and study of the
animals collected on those field trips. Those exercises which are to be carried on independently should be undertaken early in the course and plans made
for 'their completion.
During the remainder of the course, in addition to continuing the study
of the fauna as a whole, more emphasis will be placed on problems of a general
or specific nature.
GENERAL PURPOSES OF S112 and S119
The gaining as rapidly as possible of the ability to identify from the specimen or the name the common invertebrates of the region, chiefly those of
the marine littoral. This is a necessary preliminary to other types of
study. To facilitate the gaining of this knowledge preliminary lists will
be posted and named collections put on demonstration. Relatively early in
the course an identification test will be given.
The study of those characters used in the identification of the various
groups of invertebrates.
Experience in the identification of animals by means of keys.
A knowledge of the ecologic niches (the characteristic environmental settings) of all the common animals seen.
A knowledge of the grouping of the different species in faunas of specific
areas and habitats.
A knowledge of the grouping of species in associations and their interrelationships in associations, most obviously their relations in food chains
and cycles.
A knowledge of the distribution and density of population of certain species
and of groups throughout the region as a whole with a view to understanding
some of the factors which limit their distribution.
A knowledge of the habits and behavior of certain species.

FIELD STUDIB

[
[
a

Read those portions of the introduction dealing with the use of scientific
names and systems of classification, and with field and laboratory methods.
Read Flattely and Walton, The Bioloas of the Seashore; and Ricketts and
Calvin, Between Pacific Tides; papers by MacGinitie on feeding habits and natural history of a number of m i n e animals; also kcGinitie on Littoral Marine
Animal Communities; Shelford et al., Some k i n e Biotic Communities of the
cific Coast of North America; Hewatt, Ecolo~icalStudies on Selected Marine
gtertidal Communities of Monterey
Newconibe, Certain Environmental Factors of a Sand Beach; and other articles listed in the bibliography as
dealing with ecology and natural history.
Collections should be made on all field trips of animals not already
Imown or not in the demonstration sets. Those belonging to the Mollusca and
decapod Crustacea should be identified at once and those at all common learned
in preparation for the identification tests. Others should be preserved for
future identification in connection with scheduled exercises. Polychaetes
should be killed by adding formalin, a few drops at a time, and preserved.

w;

1. AS GIVEN AT MOSS BEACH

lkercises 1 to 16 deal with the marine littoral fauna of Moss Beach and
San hteo County. Exercises 13, 14, 15, and 16 should be begun at the earliest
possible oportunity. Read instructions under individual projects, p. 204,
study the exercises and organize your program accordingly.
The diversity of fauna of b s s Beach and vicinity is due to the wide
variety of physical conditions within a relatively narrow area, especially to
the diversity in nature and arrangement of the substrate. Maps I and I1 give
the local names which it will be necessary to learn in order to discuss the
different faunaa.
The following points should be identified at the outset in preparation
for work in the field: North Point, Sand Rocks, Sunshine Creek, the Reefs,
Kelp Cove, Moss Beach Flats, San Vicente Creek, Moss Beach Point, Nye's Channel, Nye's Rocks, Egregia Flats, Balosaccion Flats, Seal Cove, k i n e View
Point, b i n e View Rocks, Frenchman's Reef, the Lagoon.
Read the account by Charles Reed of the geology of the Moss Beach Region.
General Pur~osesof These Ezercises (1-161
1. Knowledge of all marine littoral invertebrate animals of the region.
2. Knowledge of the fauna of various specific habitats.
3. An appreciation of the factors responsible for major differences in faunas,
as between Moss Beach Flats and Frenchman's Reef for example.
4. A study of the distribution of certain species or groups with a view to
determining the limiting factors.
5. The behavior of certain groups or species.
These have been separated out into exercises in an effort to make them
specific. They actually overlap, forming part of a continuous program necessarily
altered by the nature of the tide and the weather and other considerations. These
exercises should be read over to gain a comprehensive picture and to allow for
greater efficiency in their prosecution.
Exercise 1. A Preliminam Survey of Moss Beach and
the Fauna of Moss Beach Flats
If the time before low tide permits there will be a brief survey of the important local landmarks.
After this follow up the receding tide across Moss Beach Flats to Nye's
Rocks and Channel and Egregia Flats, collecting any animals not known to you

196

LaBORATORY BM) FIELD TEXT IN INVERTEEW"I' ZOOLOGY

and keeping notes on those destined for your collection, together with notes
on the faunas of the different habitats.
barn the names of as many animals in the field as possible.
%en the tide has driven you from profitable collecting grounds take your
collections to the laboratory for identification, preservation, or study under
the microscope.
Do not bring in specimens of species known to you and not needed for your
collection or sea urchina and sea stars (except the small individuals of
Leptasterias and Henricia.
Exercise 2. Halosaccion Flats
Ecological distribution of a meager fauna in an area of little diversification.
Exercise 1 should have given you some general idea of the animals to be
encountered, their range of size, and ordinary distribution in a region of
diversified environmental conditions.
In this exercise we shall confine our efforts to a circumscribed area of
the flat central portion of Halosaccion Flats (hps I and 11). Study the environmental conditions, both physical and biotic. Consider such factors as
intertidal range, nature of substrate, position and disposition of the substrate, degree of exposure to sun, wind, wave impact, predators, etc.
Distinguish all the different habitats represented, determine the specific
assemblages in each, seek to understand presences and absences in terms of
definite physical or biotic conditions.
Is the fauna as a whole characterized by any conspicuous absences? If so,
how is this explained in the light of the ecological features? Is it conspicuous as a whole for largeness or smallness of the species involved or the individuals of the species? If so, set up an hypothesis to explain this feature.
Exercise 3. Fauna of Sand Rocks and Vicinitv
Location: Sand Rocks just beyond the Reefs which form the northern boundary of Kelp Cove (see M p 11).
Physical Conditions: Au area of varied physical conditions involving several very distinct habitats (and their intergrades) such as (1) high tide 01s
(in top of Sand bcks); (2) sheer, soft rock surfaces (sides of Sand Rock$
exposed to tidal and wave action; (3) a rock surface sloping upward toward the
shore compased of granite interspersed with softer areas and ending in a sheer
or overhanging shoreward face; (4) tiny tide pools contained in the surface of
this rock layer; (5) a shaly low low-tide bottom exposure; (6) the inner ends
of nearly upright reefs of hard rock (The Reefs ) extending out some distance;
(7) a region of rocks and small boulders near low tide line; and (8)a sandy
inner beach of coarse sand.
Biotic Conditions: These are correspondingly varied, from areas practically demoid of life to those with heavy growths of large seaweeds.
Pur~ose:Experience in thinking in terms of ecologic factors as determining habitats and ecologic niches. We are using the term ecologic niche to
refer to the sum total of environmental conditions, physical and biotic, to
which a particular species is adapted and which are necessary to its continued
existence. A habitat on the other hand is a
of place, a place or region
characterized by certain major environmental features. A habitat will possess
a characteristic fauna, each species of which has its own niche. Thus, for
example, a tide pool will contain a number of different species each with its
own niche. Since the distribution of ecologic conditions within a habitat may
spatially limit an ecologic niche we may, for convenience, connect certain
niches with places within the habitat, but this is not true of many/ niches and

FIEU) STUDIES

197

not really correct for any niche. To completely define a niche would involve
an immense amount of quantitative and experimental investigation of the physical and biotic ecological factors not yet begun for the species we study. However,' we are usually able to discern one or more obvious features of each
niche which give it objective reality and allow us to give it a preliminary
definition and to use it in discussion.
Jkcedure: Study each of these habitats and its characteristic fauna. Be
able to discuss the distribution of the different groups and species in this
area in tern of the ecologio factors involved. Be able to make a field key
to the species of Acmea, and to answer such questions as the following: Why
are the gastropods (aside from Acmaea) absent from the Sand Rocks? Which genus
is dominant on the sides of Sand Rocke? What is the dominant species in the tide
pools in the top of Sand Rocks? Where did you find zonation?
There will be no report but a written test will be given covering the information gained in Exercises 1-3.
You have now seen three different habitats or locales each with its own
characteristics and variations. You should begin to form a picture of the ecologic ranges of certain species, such as those of the genus Acmaea (see Exercise No. 12). the hemit crabs (see the key, p.113 and the special paper by
Harris), and the flat-topped crabs of the genus Petrolisthes(see special paper
.
by Fields and Benton).
Sam~leQuestions on Sand Rock and Vicinitg
What species not found or rare at Sand Rock and vicinity were found on
Halosaccion Flats proper?
Name species found abundantly in the vicinity of Sand Rock that were rare
or not found on Halosaccion Flats proper.
Suggest one general reason for the fact that the latter list is longer
than the former.
What were the most important plant f,eeders;scavengers; predators; plankton feeders, in the Sand Rock vicinity?
Name animals characteristic of Sand Rock proper that were not found among
smaller rocks around its base.
List differences in fauna between loose rocks on sandy substratum closer
in and farther out.
Which of the three major factors seems most important here? Illustrate.
On the basis of today's observations how could you distinguish
the niches
of the three shore-crabs?
Nhat evidences did you note of vertical zonation of the species of Acmaea?
Which was the most abundant polychaete?
What two features of the situation probably account for its abundance?
What are the advantages and disadvantages for A. scabra of sandrock as
against granite rock?
Why SO few barnacles and no mussels on sandrocks?
IA4y so few littorines?
What structural character of Cirriformia lmriosa not found in C. spirabrancha is correlated with its habit of life?
Name tL-ee genera of isopods encountered. Characterize the very distinct
ecologic niche of each. Which two may be found together? Where and why?
Vith what other animal did the presence of Acanthina seem to correlate?
Vhy were borers not abundant? Nestlers?
What terebellid worm was abundant? Give some features of its niche.
Exercise 4. Frenchman's Reef
Location: Between Marine View and Pillar Point (see k p s I and 11).
Physical Features: A rocky reef, with a relatively flat ,top, extending
into the open sea, exposed on its high side to direct wave action and cut by
numerous, deep, narrow, tidal channels (see Map 111) with steep, often overhanging walls; numerous high tide pools; ledges under exposed horizontal rock

198

LABORBTORY ANIl FIELD TEXT IN INVERTEBRATE ZOOLOGY

strata; a protected, shallow lagoon between shoreward end and shore on north
side.
Biotic Features: Very great abundance of algae except on high central
shoreward portion; beds of narrow eelgwss and Laminaria, etc., on protected
side; sea urchin beds and mussel bed near outer end.
Procedure: Study first the deep tidal channels and the fauna of the seaweeds and eelgrass of the protected sides. On the return study and attempt
to explain shoreward changes in fauna on the reef surface. On the way out or
back study the under-rock fauna of the lagoon, comparing it with that of Iridaea
Channel and &rine View Rocks. Bgdroids should be collected wherever found and
taken back to the laboratory for study and identification (at least ten species should be found).
A colony of the sea anemone, Corynactis, occurs under one of the high
ledges about halfway out. Be certain to see it.
Attention should be paid to three species of limpets each confined to P
single plant species: A. paleacea on the eelgrass (Phyllospadix), A. insessa
on Egregia,and A. instabilis on the stalks of Laminaria; all found on the north
side of the reef.
Some bestions on Frenchman's Reef
1. List nine or more habitats or subhabitats each with an essentially distinct
fauna. Give one to three characteristic species for each.
2. Name five extremely stenotopic species of sessile gastropods.
3. PVhat advantages probably explain the abundance of Plumularia on the bases
of the brown algae near the laminarian zone?
4. At rmty points a fauna characteristic of the lowest littoral or the sublittoral was encountered at relatively high levels. How could you explain this in terms of physical factors?
5. b t biotic factor (associated with a physical feature of the subhabitat)
operates with regard to the hydroids?
6. What factors do you conceive to operate to determine the distribution of
Strongylocentrotus purpuratus?

7. f i t major difference with regard to S. j'ranciscanus?

8. Name four types belonging to as many hyla which are found competing for
space in certain favorable spots? &at makes these spots favorable?
9. Name five genera of hydroids encountered. Which was most abundant?
10. What arthropods seem adapted for life on this particular hydroid?
11. What highly stenotopic pelecypod was encountered?
12. Why the relative scarcity of limpets biotic factor)? Of barnacles? Of
Tegula except near inner end of ree ?

Ekercise 5. Iridaea Channel

Location: Iridaea Channel (see Map 11), really lagoon-like, is a lower
area in Moss Beach Flats which drains at low tides through Bye's Channel and
Egregia Flats.
Physical Features: Protection from strong tidal or wave action and from
exposure except at extreme low tide. Numerous rocks resting in gravelly sand
containing m c h organic detritus.
Procedure: Learn the fauna and attempt to explain it on the basis of physical features of the environment. Be able to name another location where the
same general conditions prevail. What two factors seem most significant in
determining the nature of this fauna?
Some Questions on Iridaea Ch&
1, Pmat other spots have furnished a somewhat similar fauna? Which one especially, at Frenchman's Reef?
2. List five physical features and two biotic features which conspire to bring
together the characteristic fauna.

[
FIELD STUDIES

(1) the in-rock dwellers, as borers, nestlers or cranny

the on-rock dwellers both those of exposed and of under
the under-rock dwellers; and (4)the dwellers in the
under-rock substratum.
4. Have in mind the food habits of each and, in so far as possible, the
reasons for their presence there.
5. For each of the following name one or more species which are conspicuous
in, or characteristic of, the fauna:
Larger snail-like gas- Surface-dwelling
Brachgvrous anomurans
pelecypods
Anomurous anomurans
tropbs
Cranny-dwelling gasBrachyurans
Asteroids
Chitons
tropods
Limpets
Errant polychaetes
Barnacles
Rock-dwelling brachyurom anomram
Sedentary polychaetes
Isopods
Aberrant annelids
Rock-boring pelecypods Ophiuroids
Nestling pelecypods
Holothuroids
Small snail-like gastropoas
6. b t two types of prochordates (chordate invertebrates) are found?
7. What was the largest animal? W a t correlation between its size and its
location?
8. What important predators were not seen by us?
9. Why were barnacles less numerous than on Nyels Rock?
10. What species of Acmaea is found only here? b t does the nature of its
shell and the places where it is found suggest as to the factors limiting
its distribution?
Exercise 6. brine View Rocks and Seal Cove
Location and Ecoloaic Features: Most important controlling physical condition is the presence of a broken line of outer rocks and reefs which more or
less completely protects (1) the area of rocks of various sizes lying in gravel
or sand which is Marine View Rocks (Map I); and (2) Seal Cove just north of it
and between it and Halosaccion Flats already visited. These outer reefs support
a heavy growth of many types of seaweed which are frequently torn away and
carried in over the Marine View Rocks, and along the shoreward side of Seal
Cove, where they break down and decay, furnishing food for many animals.
Note similarities hetween conditions and fauna at bhrine View Rocks and
other areas studied. One rock rises high enough to have a sampie of a formation different from that found elsewhere in the vicinity. What gastropod marks
this formation and why does it occur at that particular spot?
Seal Cove is particularly interesting for the occurrence there in numbers
of two species which occur rarely or not at all in the other localities visited.
These are Olivella biplicata, to be found in the sand at the end of surface
trails, made presumably at night, and a large cirratuled worm, Cirriformia s p i rabrancha, whose slender, greenish tentacles resemble nemerteans.
Locate these two and attempt to explain their abundance here and their
absence or rarity elsewhere.
Exercise 7. The Fauna of a High Rock ExDosed on Its
Outer Face to Violent Wave Action
Location: NyelsRock (see &p I1 ), separated from the high outer front
of Mosa Beach Flats by a deep narrow tide channel, Nyels Channel.
Physical Features: Rock sufficiently hard to withstand wave action but
still penetrable by borers and sea urchins; base below low low tide, hence in
Laminaria zone; high enough on seaward face to be exposed except at high ;hter; with two lower shoreward shelves.
Biotic Features: A relatively rich growth of seaweeds of various types;
mussel beds at certain areas and levels.

200

LABORAMRY AND FIELD !LECT IN INVERTEBUTE ZOOMGY

Habitats: A great number of habitats, some protected from wave action,

others exposed to its greatest violence or to the scouring effect of tidal
channels, tide pools, crannies, mussel beds, close set algal growths, Iaminaria, etc.
Procedure: W e as complete as possible a study of the fauna. Note differences between exposed and protected sides. Seek an explanation for the location
of the,different species or associations, especially for the mussel and Mitella
beds and the sea urchins.
Exercise 8. Fauna of a Tidal Cave
Drake's Cave, really a tunnel, furnishes excellent opportunities for a
study of the influence of light or its absence. Attempt to segregate the influences of reduced light from those of the peculiar water flow resulting from
the configuration of the cave and its varying diameter.
Before the field trip read the special problem report by Gmody and Villee
on "The Biology of Drake's Cave Check your findings egainst theirs as to the
fauna and its distribution. Am the differences real or simply differences in
sampling? If real, have you any explanation for them?

."

Exercise 9. Under-Rock Faunas

Purpose: To visualize the importance of the factor of protection.
Procedure: Choose a rock resting on or partly embedded in ?and or gravel,
one small enough to be turned over, but yet of considerable size and so located
as to be relatively stable.
Note carefully the physical location of the rock with regard to tide
level, protection from wave action, etc., also whether embedded in or resting
on the substrate, or with space between it and the substrate.
Overturn the rock and make immediate notes of the active population which
almost at once seeks protection elsewhere.
Carefully list and evaluate the remaining fauna of the lower surface of
the rock and of its crevices and crannies if any.
Attempt to arrange the total fauna thus listed on various bases: (1) as
to the reasons for their presence there, (2) their food relations, etc.
Exercise 10. Rock Dwellers
At a low tide choose a smll rock from Fgregia Flats or break off a small
piece from the shoreward bank of Nye's Channel.
Collect and make careful record of the surface-dwelling animals.
Break it open and identify and list (1) the crevice and cranny dwellers,
(2) the rock borers and (3) the nestlers.
Attempt to explain the presence of each species in its particular location.
Exercise 11. Fauna of a Seaweed EIoldfast
By means of a geologist's pick separate from the rock a holdfast of one
of the larger algae (Laminaria, Egregia, etc. ) found at low tide level along the
front of Moss Beach Flats, Nye's Channel, or on Egregia Flats.
In the laboratory break it open and list the different species present.
Attempt to explain the presence of the different species.

FIELD S'IIIDIES
Exercise 12. The Distribution and Habits of the Limpets of the Genus Acmaea
Furuose: The gentm Acriiea offera an unusual opportunity for the study of
numerous species of the same genus living in the same general habitat. Fourteen species occur in this area, of which ten should have been encountered by
all. The purpose of this study is to consider the structural differences of
these species, their interrelationships within the genus, their habitats and
habits of life insofar as time allows with a view to discovering what correlation, if any, is to be found between these structural characters and the habitats and habits of life of the different species. The following ten species
will be used for the study, all of them common except Acmuea persona and
A. instabilis which will be pointed out to you:
1. Acmaea asmi
6 . Acmaea paleacea
2. Acmaea digitalis
7 . Aomasa persona
3. Acmaea insessa
4. Acmaea mitra
5. Acmaea pelta

8. Acmaea scabra
9. Acmaea scutum

10. Acmaea instabilis

(Other species occurring but not so favorable for such studies are:
Acmaea genestrata, A. 1imatula, A. ochracea, and A. triangularis. )
Procedure: Make careful notes as to the characteristic location of these
species a8 seen by you during low tides. Look for characters of general shape,
color, and marginal ornamentation which seem to have an adaptive relation to
the particular niche which each occupies.
Read introductory portions of thesis of Dr. Avery Grant and papers on
homing behavior by Hewatt, and by Villee and Groody.
If time of tides allows (as during the summer) observe their behavior on
a nocturnal low tide. Again look for correlations between their feeding behavior, their "homen locations, and their structural characteristics. To what extent is there competition between species?
Exercise 13, A Collection of bbllusks. Crustacea and Pol~chaetes

correlating these notes with the specimens, in preparing the specimens, in

their identification, and in labelling them properly. The work should be conducted at all times as if the materials were to be used for purposes of scientific research as, indeed, may well be the case.
The specimens turned in should, in general, be only those collected by
the student involved. If specimens are obtained from others the collector
should be cited on the label and it should be clear who is responsible for
the ecologic data. Auy other procedure or attitude vitiates the whole situaJuveniles or worn or damaged specimens should not be used except under
unusual conditions.
Collections will be evaluated on the basis of completeness, correctness of
identification, excellency of the preparations, neatness in organization and
labelling, accuracy and completeness of data furnished and especially on the
general and specific indications of scientific attitude and method.
Read carefully the general directions for collecting and for the keeping
of field notes, and also the section on the use of scientific names.
Identifications should not be made lightly. Read the section on the identification of invertebrate animals. Where the species is uncertain the specimen should be labelled with the generic name followed by sp.? Thus: Tegula
sp.? If it is believed to be a certain species but the identification is in

LABORATORY AND FIELD !l!EXT IN lTlEWEBRATE ZOOLOGY

202

doubt, a question mmk in parentheses should follow the author's nam, thus:
Tegula finebra1 is Adams (? ).

Chitons: Unimum, 5 species. For permanent preservation chitons should be

placed in a mixture of equal parts of water, alcohol and glycerine. For our
purposes fixation in 5% fomlin after flattening on a glass slide or other
flat surface will be satisfactory. Specimens thug fixed may be left in formlin. Peleosoods: Minimum, 9 genera. Remove soft m e , using hot water if necessary. Gmtropods: Minimum, 30 species in at least 15 genera. Bodies are
best removed after leaving for some days in 30% alcohol. This is too expensive
and time consuming for our purposes and it m y be accomplished by placing them
in hot (not boiling) water. This may slightly alter the color and surface of
the shell hence making them unfit for a permanent collection.

Minimum, 15 species in 10 genera. For permanent preservation specimene

should be anaesthetized, fixed in special fixatives and preserved in alcohol.
For our purposes they may be preserved in formalin and identified in connection
with the laboratory study of this group.

1aounts of at least 5 species, includMinimum requirements: Co~e~oda,

, least 8 species.
ing at least one from each of the common orders. h ~ h i ~ o d aat
Isowda, at least 6 species. Macrura, at least 3 species. Anomura, at least 6
species in at least 4 genera. Brachsura, at least 10 species in at least 6
genera.
Ornanization of collections. Specimens should be arranged in a species
list by phyla and classes and Ey collection numbers. When turned in they must
be so arranged as to be easi.ly inspected and checked. Iabels should be made
with India ink. See page 175 for instruction as to contents of label. FuZl
scientific names should be used.
Exercise 14. Collection of a Sin~leGrow and a Studs of
Its Ecolo~icalDistribution

This should be begun as early as possible.in the course to allow for

studies at low tide. Choice of group should be approved by the instructor. The
degree of emphasis on the taxonomic or the ecological phase of the exercise
will be determined in some cases by the size of the group involved, in others
by the interest of the student.
Groups suggested are: the hydroids, the Bryozoa, the sea anemones, the
seastars, the true crabs, the shrimps, the Anomura, the species of Petrolisthes,
the species of Pagurus, the Acmaea, the small gastropods, the species of Crepidula , the amphipods, the isopods, harpacticoid copepods, the polychaetes in
general or any group within them such a9 the Nereidae, the Phyllodocidae, the
Spionidae, the Cirratulidae, etc.
Exeroise 15. Fauna of Small Tide-Pools with lhphasis on
Behavior and Feediw Habits
Location: A small, permanent, relatively high tide pool. A particular
pool will be assigned for this study.

FIELD S'lWDIES

203

Procedure: Visit the pool at various times and under different conditions
as regards tides, etc. Do not disturb unnecessarily the inhabitants or remove
any of them at any time during the study. Station yourself so as not to cast
a shadow on the pool.
W e a careful survey and census of the population keeping a faunal list
with approximate numbers of individuals. W e a diagram to show distribution
of the inhabitants. Species not sufficiently deen to allow for diagnosis mag
be described and identifications will be supplied by the staff.
Note and record any activities observed during a period of 15-30 minutes
after'the original disturbance, if any, has quieted down. Attempt to explain
these movements.
Devise experiments to determine reactions of different animals to various
types of stimuli.
Introduce a small piece of mussel previously prepared so as to avoid disturbances. By this method seek (1) to determine the numbers and locations of
cahivorous animals, (2) to understand and record their feeding behavior, (3)
to determine whether sight, chemical sense or touch enables them to find their
food.
By varying this devioe seek to determine which are facultative carnivores
and which obligatory. By using tiny fragments of dead seaweed detect plant scavengers if any. By using whole tiny organisms seek to determine which, if any, of
these carnivores are predators. By using different animal bodies seek to determine whether they have favorite foods. Interpret negative results with caution.
Consider also relations with other species present only during high tide.
Attempt thus to build the food chains involved. Do they form a cycle? Visit the
pool numerous times on different days, preferably on sunny days and after the
water has become warm. Before leaving the pool remove any debris or food particles which you might have added. Leave the pool as undisturbed as when you
found it.
W e an effort to determine (1) to what degree the fauna changes from day
to day, (2) to what extent the animals concerned have permanent locations.
Turn in a report covering all the points studied.
Please make this an entirely individual study and report avoiding discussion or conference.
W e the study and report an exercise in the scientific method.

Exercise 16. The Fauna of the Hi& !t!ideDools between Sand Rocks and
North Point and the Problem of the Distribution
of the Red Cope~ods(Tiqriopus californicus)
These tide pools vary in size from a few inches in depth and diameter to
deep pothole pools on the top of sand rocks and elsewhere. The substratum also
varies from soft shaley sandstone through conglomerate to granite. Other physical factors possibly concerned are elevation, exposure to sunlight, etc.
The dominant animals of these pools are three, Pachygrapsus crassipes,
Boccardia probiscidea, whose waving tentacles are readily seen on close observation, and the red _copepod, present in varying numbers or absent. Some of the
pools are inhabited by the cirratulid worm, Cirrij'ormia luxuriosa, whose dark
tentacles form conspicuous patches.
Study the distribution of the copepods with a view to determining why they
are absent or nearly so in some pools and present in others. Important among
the factors to be considered are predatory animals and food requirements.
Previous studies have shown the copepod adapted to withstand extreme ranges of
temperatures and salinity as would be expected of an animal living in small
high tide pools.
On the basis of early observations propose an hypothesis. Make observations
at later times, organize your evidence, and criticize your hypothesis in the
light of the evidence. Incorporate this in a report to be turned in not later
than the end of the fourth week.

204

LABORATORY AND FIELD TEXT IN INVXRTEBRATE ZOOMGP

Bercise 17. The Identification of Pobchaeta

Identify and label all the polychaete worms collected using the
key on page 34. Refer to Hartman (thesis and various papers), and Fauvel,
Errantia (1923) and Sedentaria (1927) in Fame de France.
Co-operate with the staff (1) in accumulating a correctly labelled series
of well-preserved reference specimens, and (2) in gathering all animals seemingly not in our lists or new to the region to be sent to Dr. Hartman for
identification.
Exercise 18. Fauna of Tidal Flats in an Enclosed Bay (Tomales
Bay near Dillon Beach. k ~ IV
s and V)
For directions see kercises 28, 30, 31, and 32.
Read: kcGinitie, "Ecological Aspects of a California Marine Estuary
(Elkhorn Slough)" and "Littoral hrine Communities"; Shelford and Towler,
"Animal Communities of San Juan Channel and Adjacent Waters1'; Shelford
et al. "Some Marine Biotic Communities of the Pacific Coast of North America";
Alice Mulford, report of special problem on "Distribution of Callianassa."
Individual Proiects
All members of the 8112 class will do the following as individual projects:
1. Either Exercise 13, 14, or 15, with collection (or report if 15).
2. An intensive study of the behavior of some one common species.
A special problem (see suggested subjects below) may be done either in
addition to the above or, with the instructorlspermission, in place of 1 or 2.
If additional credit is desired for special problem work the student should
enroll for extra units early in the course.

Each member of the S119 group will carry out at least one special problem. The number of possible problems is almost endless. A careful study of any
species from any one of several points of view would produce results of interest and value. In addition there are many problems of various types dealing
with groups and local faunal assemblages.
Swsestions for S~ecialProblem8

Distribution and behavior of the two littorines, especially factors differentially affecting the two species. A collection of nemerteana with notes
on their behavior and distribution for Dr. Coe. Further studies on Tegula
finebralis (see paper by Gorbman). The distribution of Tegula brunnea, limiting factors and an analysis of population. Behavior and habits of the large
sand flea, Orchestoidea corniculata. Orohestoidea cal ifornioa, its behavior
and distribution (Half b o n Bay) and limiting factors. Is it a separate species? Fauna of Montara tidal caves. Habits and behavior of Olivella biplioata;
food. Anatomy of Olivella with a view to its availability for use as type for
the study of streptoneury, asymmetrg. and extreme torsion. Claasification and
ecologic distribution of any common family of polychaete worms such as (1)
Lumbrineridae and Glyceridae, (2) Opheliidae and Orbiniidae, (3) Nereidae,
(4)Phyllodocidae and Nephthyidae, (5) Syllidae, (6)Terebellidae, (7) Polynoidae, (8)Sabellidae, etc. The anemones, with a key for their identification.
Structural and ecologic differences between the two large anemones, Anthopleum
xanthogramanica and Bunodactis. A key to the hydroids of Frenchman's Reef with

FIELD s m 1 m
notes on their distribution. !be fauna of the inner lagoon of Frenchman's Reef.
Taxonomy and biology of the common flatworm. Fauna of a sandy beach (bntara
Beach). A practical key to the common Bryozoa. The harpacticoids. Stages in the
development of the isopod, parasitic in barnacles at Marine View Rocks. AllonisCUS, the high beach isopod, its habits and ecology. Marine and beach dwelling
insects, their behavior and ecology. Authentically named demonstration collection of Polychaeta. Botula falcata and B. californica; distribution; borers or
nestlers? If borers, how accomplished? Anatomy of pholad compared with that of
typical clam and an attempt to determine adaptive features of anatomy. Acmaea
limatula; distribution; limiting factors. Same for A. fenestrata. Ehlosaccion
Flats: (1) Distribution of polychaetes; (2) distribution of decapods, especially
hermits. Feeding habits of Acanthina. Further studies on ecologic segregation
of the species of Pagurus (see paper by Morgan Harris). Further study on homing
of Acmea scabra (see papers by Hewatt, and by Villee and Groody).
2. AS GIVEN

AT DILLON BEACH AND VICINITY

Dillon Beach lies on the shore of Bodega Bay not far north of the mouth of
Tomales Bay (Maps IV and V) and within reach of numerous diverse faunas.
The bold rocky outer coast of Tomales Point affords excellent opportunities
for the study of animals of tide pools, tide channels, and exposed rocks rising
from the low tide line. Just off this shore lies Bird Island, which supports a
fauna less reduced by human collecting than that of the shore. Tomales Bay has
m y tidal mud- and sand flats supporting rich faunas. Within reach, also, are
numerous other interesting marine faunas. Such are those of Bodega Head to the
north ( k p IV), Duxbury Reef, Bolinas Bay and Bolinas Iagoon to the south
(&p IV), and Moss Beach south of San Francisco (hps I, 11).
Tomales Bay, as its long, narrow, straight shape would indicate, is a
fault bay. Bolinas Bay and the creek which runa into it from the north are continuations of the same fault line, the San Andreas Rift, which traverses San
Francisco.
The fault line marked by the bay separates the Point Reyes block from the
mainland. This block is supposed to have been an island during much of Tertiam
times, perhaps connected with the Farallons. Its mesozoic granites are exposed
along the western shore of the Point, where we shall collect.
The eastern shore of Tomales Bay is a part of the bed of the ancient Merced
Sea (Pleiocene). Along most of this shore the Merced deposits have been eroded,
exposing the older Franciscan deposits (Jurassic?) like those about Berkeley.
In the northeastern portion, however, and along the shores of Bodega Bay, the
Merced deposits remain. The conspicuous rocky outcrops east of Dillon Beach
formed part of the shores of the Lrced Sea, and the rocks which we studg at
Perch Rock Point and Second Sled Road must be from the Merced or from Franciscan deposits.
The sand dune area south of Dillon Beach partly closing up the mouth of
Tomales Bay consists of much younger (Pleistocene and Recent ) alluvial deposits.
Consult Dickerson (1922, Proc. Calif. Acad. Sciences) for a detailed account of the geology of the region.
Read the general statements with regard to thb summer course (p. 193) and
individual projects (p. 214). Begin as soon as possible the reading of appropriate chapters in Flattely and Walton and in Ricketts and Calvin, especially
those dealing with the tides. C o d t also the schedules given you and the list
of low tides and begin to plan your program. Study k p s IV and V and make yourself familiar with the local names in preparation for field trip.
A list follows of field studies which may be made with Dillon Beach as
headquarters. Not all these are possible in any one six-week period and during
the tidee available. A tentative schedule will be issued each year listing
those field trips to be attempted and the dates as well as the laboratorg exercises to be interspersed among them. Keep this schedule and amend it from time
to time on the basis of announcements to be made.

206

IABORATORY AND FIELD TE!P IN INFTERTEBRllTE ZOOLOGY

FIELD S W I E S FROM DILLON BEACH

Exercise 19. The Distribution of Polychaete Worms

Exercise 20. The Ecologic Distribution of the Limpets of the Genus Acmaea
Exercise 21. Field and Iaboratory Observations on the Adaptations in Behavior
and Structure of Callianassa, Upogebia and herita
Exercise 22. A Study of the Fauna in an Area of Rocks Resting in Sand (Perch
Rock Point, the first field trip)
Exercise 23. The Fauna of Iarge Bcposed Boulders of Hard Rock (Second Sled
Road Point, second field trip)
Exercise 24. The Fauna of an Outer Exposed Rocky Shore (Tomales Point)
Exercise 25. The Fauna of a Rocky, Exposed Islet (Bird Rock)
Exercise 26. The Fauna of a Long, Warrow Rocky Reef Extending into the Open
Sea and Cut by Deep, Narrow Tidal Channels (Frenchman's Reef,
Moss Beach)
Exercise 27. Boring, Nestling and Cranny-Dwelling Animals
Exercise 28. A General Survey of the Faunas of Various Areas of the Tidal Flats
in Upper Tomales Bay
Exercise 29. The Fauna of the Tidal Mud Flats of Bolinals Lagoon
Exercise 30. The Fauna of the Eelgrass Flats of Upper Tomales Bay
Exercise 31. The Phoronopsi s aod the Maooma-Phorbnopsi s Aasocietl
Exercise 32. Ecological Succession and Distribution on the Sandy Mudflats with
Bnphasis on the Fauna of the High Flats
Exercise 33. Estuarine Faunas of Piles and k d Flats (Inverness on the Southwestern Shore of Tomales Bay)
Exercise 34. Oyster Breeding and the Fauna of a Commercial Oyster Bed (Bivalve
or Ahshall on the Eagt Side of Tomales Bay)
Exercise 35. A Study of the Minute Plants and Animals of the Sea, the Plankton,
mich Constitute the Food of
Seashore AnimaLe
Exercise 36. An Individual Study of a Small High Tide Pool
Exercise 37. An Individual Study of the Fauna of a Limited Area of Rock Surface
in the Intertidal Zone
Exercise 38. An Individual Stddy of the Fauna and Food Cycle of a Fresh Plater
Pond (Rolland Pond or another)
Exercise 39. The Fa- of a Small Coastal Stream ( S b n Creek) with an Attempt
to Visualize Adaptations to Flow and Some of Its Food Chains
Exercise 40. Terrestrial Faunas

Exercise 19. E C O ~ O RDistribution

~C
of the Polvchaete Norms

A large number of species of polychaetes belonging to many genera and families are to be found in this diversified area. See Dr. Bartman's thesis for
a list of species and something of the ecologic distribution of some of them.
Polychaetes should be collected whenever found and, after microscopic observation and identification, properly preserved and labelled. If it is not possible
to identify them at the time preserve them for future identification. About the
end of the second week a period will be devoted to the identification of polychaetes.
Please co-operate with the staff in its efforts to accumulate a wellpreserved, correctly-identified, study collection and also in the accumulation
of rare forms or those new to this locality. Since the summer course has been
given at Dillon Beach 0114once previous to 1941, much remaine to be known of
the polychaete fauna and its distribution.
There is a distinct value in identifying and becoming familiar with a wide
range of types in a group. It gives a picture of the range of structural diversity, the results of a long evolution. Here, in the realm of taxonomy, a
rare species is just arr important as a common one, or more so. To the student

FIELD SllJDIES

1
I-

of ecology, however, the species which occur in considerable numbers are the
important ones since they have a proportional effect upon the communities of
which they f o m parts.
M e certain, therefore, that you collect and identify all of the species
of polychaetes which occur in any oonsiderable numbers, keeping as complete a
mcord as possible of this occurrence and ecologic distribution. Be prepared to
report on the ecologic distribution of at leLt a dozen such common polychaetes.
kercise 20. The Ecoloaic Distribution of the Lim~etsof the Genus Acmaea
Read introductory portions of the thesis of Dr. Avery Grant and papers on
homing behavior by Hewatt, and by Villee and Groody. Identify in the field or on
return from it all species of the genus Acmea. Place them ecolo ically with
regard to (1)vertical location. (2) nature of substrate, and (37 food.
Keep this in d n d on all field trips and be able to combine your total
findings into a chart or diagram.

. Distribution
. .
oise 21. The Ecoloaic
of Ca
' a . and Bwritg
C o n a i w in the liaht of Their A d a p t a t ~ B e h a v i o r
(An-)

Read papers by AkcGinitie on the natural history of these three decapods

with highly specialized structure and habits.

Pumos~: (1) To understand the correlations between special structure, behavior, and environmental location, that is, the structural adaptations, of
these three a n o n decapods adapted to a life in the substrate, and (2) to
see how differently the fundamental decapod type of structure has been modified
in the thres species.
WcedurQ: Bemove and arrange in parallel series all the appendages of the
right side of the three species. Care n w t be taken in rearoving the peraeopods
to obtain the gills with them. Each peraeopod is provided with a gill attached
near its proxhl articulation.
W e a table for these appendages, with separate columns for the three
species in the order Calliaassa, Upogebia, mrita, to bring out all differences. Attempt to correlate differences of structure with differences in behavior and these with life-habit differences. Is Upogebia more like Callianassa?
If so why? Which of the two is most like Bhrerita? Do you think this is to be
explained in terms of relationship? If not, on what basis do you explain it?

B. Field Studies

The sandy beaches from Sand Point up to Second Sled Road Point and beyond
abound in sand crabs, the adults scattered but numerous, the young in countless thousands. These should be observed from the first field trip. Be able to
describe their method of swimming, walking and digging in, the type of trail
they make and the part played in these different activities by the different
thoracic appendages. M: kcGinitie, nMavements and kting Habits of the
sma clgb."
See if you can discern by your own observations an animal which preys on
them. This will best be made out when the tide is coming in. Be able to present
your evidenoe in convincing form.
W e yourself familiar with the holes and burrows of the ghost shrimp in
the upper end of Tomales Bay. See the various associated organisma especially
the gobies, the pinnotherid crabs and Crypto~ya.P r e l m reading: Mulford,
Distribution of Call ianassa californiensis Dana at Sand Paint; IkoGinitie, "The
Natural Hietory of Callianassa." Run a transeot from the inner eelgrass associes

11t

208

LABORATORY AND FIELD TEXT IN INVERTEBME ZOOLOGY

I/

to the sand dunes noting the change of fauna enroute. Be able to draw a diagrammatic section showing this change in fauna in relation to increasing height of
surface of subs'grate
In running the above transect individuals of Upogebia may have been encountered at the lower levels. If not, make a search at or very near the lowtide level along the inner channels to locate these mud-sbrimps. Attempt to
state how spatially and ecologically they are isolated from Callianassa.
In this connection read EkcGinitie, "Natural History of the kd-Shrimp."

Exercise 22. Fauna of a SandB Beach and Scattered Elevated Rocb in Sand
Location: Perch Rock Point just at northern edge of the Dillon Beach
settlement.
Pbsical conditions: Facing the open sea to the west, hence subjected to
wave action, violent at times of storm. Rockt~relatively hard with few if any
crevices or crannies. Beach relatively long with gentle even slope from low
tide level to the rocb cliff. Rocb irregular in shape but generally sloping
to a narrow top, bottoms buried for unknown depth in sand. Sand washed out
about bases of most rocks, leaving temporary tide pools. Two major habitats,
rocks and sand.
Biotic conditions: Living seaweed small in quantity: very little on rocks,
a few scattered clumps of Xgregia in sand. A certain amount of dead seaweed is
present. That at upper level should be investigated for plant scavengers. Algal
film seemingly lacking on upper rocks. Two striking rock communities, one dominated by Mytilus californianus, the California Mussel, the other almost pure
sea anemones.
The following animals are among those present:
Rock Fauna
Anemones, large and small

Acmaea 1 imatula Carpenter (rare)

Myt ilus cal ifornianus Conrad
Ligia (The Rock Louse)
Katharina
Mopalia ci1 iata Sowerby
Balanus glandula, the common acorn

Tegula finebral e Adams

T. canal iculata Duclos
m a i s emurginata Deshayes
Littorinu scutulata Gould
Littorina planaxis Philippi
Acmaea digitalis Eschscholtz
Acmuea pelta Eschscholtz
Acmaea scabra Gould
Acmuea scutum Esch~choltz

barnacle
B. oariosus, the thatched barnacle
Chthamlus, the tiny barnacle
Mitella, the leaf barnacle

In kssel Bed
17rais canuliculata Duclos
Petricola (rare)
Nerei s vexillosa Grube
Petrolisthes, the flat-topped crabs

Pachycheles, the big clawed crabs

Pogurus
Cirolana

Amphipods

Sand Fauna
Nainereis
Nephtys caeca Fabricius
Nere is vexillosa Grube

Emerita analoga, the sand crab

Orchestoidea, the sand flea

Others
Pisaster ochraceus, the ochre seastar
Pachygrapsus, the lined shore crab
Hemigrapsus nudus, the purple shore crab

Epialtus ~roductus,the kelp crab

Cancer antennarius, the rock crab

FIED STUDIES

209

@r~ose:To gain familiarity with the members of the fauna, their names,
their ecological niches, their interrelationshipe.
Procedum: Identify as many of the animals as possible in the field, use
their names again and again until they become familiar. Bring those not identified to the laboratory for identification. Keep careful notes as to distribution and associations and of any biotic or physical factors which seem to explain them.
Be able to answer such questiona as the following:
b t is the dominant animal in the most oonapicuous animal association of
the area? R m t other animals are present in this association? What is their
relation to it? b t other sedentary animal is present in considerable concentration? Does it seem to compete with the other association? W e a field key
to the species of Acmaea. What evidence did you note of differences in niche of
these species? What is the most important sand dwelling organism? %at plant
scavengers were noted? %ere are the different polychaetes to be found?
Do you find any clams? Reasons?
How does the method of progression of Emerita differ from that of most
crabs? Can you differentiate their trails?

Exercise 23. Hard Rocks in Intertidal Zone (Second Sled Road Point 1
Recognize as mang animals as possible in the field. Collect and identify
those not thus recognized. Get as clear a picture as possible of the distribution of the animals of this locality in relation to the diversified physical
and biotic factors involved.
Attempt to classify the general area into more or less definite habitats
each marked by a characteristic faunal assemblage. Such, for example, would be
the beds of narrow-leaved eelgrass ( ~ h ~ l l o s ~ a d1i z
Find the outcrop of softer rock and collect the borers, nestlers, and
cranny dwellers in it for study in connection with Reroise 26.
Exercise 24. An Outer Emosed Rocks Shore (Towles Point 1
This area offers the richest fauna of any habitat to be studied in the
Dillon Beach locality and approaches that of Frenchman's Reef. Especially to
be studied are (1)the tide pools, (2) the tidal channels, (3) under rock assemblages, (4)the mussel beds, here at climax.

Exercise 25. The Fauna of a Rockv Island (Bird Island). with Tide
Pools. Tidal Caves and Tidal Channels
Get as complete a picture of the fauna, its distribution and the ecologic
features involved as is possible in the brief time available. Biotic features
require attention. Such are the luxuriant growth of seaweed and the negative
one of the relative absence of the disturbing presence of man. Be able to explain various features of the fauna in the light of these factors as well as
the physical conditions.
The tide pools are on the flat rock at the southwestern end of the islands,
the tidal cave at the northern end, the tidal channels along the western shore.
On top of the high flat-topped plateau which form the eastern half of
the island is a colony of nesting cormorants and the nests of gulls, etc. These
are interesting as forming end organisms of some of our food chains.

LBBORATORY AND FIELD TEXT IN INVER!l!EBRA!CE

210

ZOOLOGY

&
J
d i $m into
and Cut by Deen, Narrow Tidal Channels
(Frenchman's Reef, Sen Mateo County, &ps I and 111)

Sea
&

This region is &iscussed elsewhere, Field Trip C, p. 183 and Exercise 4,

p. 197.

Attempt to segregate in your mind some eight or ten very different types
of habitats found here and their characteristic assemblqes. Were elsewhere
have you found the same or similar faunal assemblages? Have they been correlated with similar environmental conditions? If so, you can begin to fornorlate
general statements as to the animals to be found under given sets of environmental conditions.
W e a special effort to see and to collect for identification as w
hydroids as possible as well as certain species of A o m a not colamon elsewhere
(A. paleacea, A. instabilis, and A. insessa).

A number of pelecypods are able to bore into relatively hard substrata or

occur in cavities in substrata so hard as to seem to indicate that they have
bored into it. It is not known just how this boring is accomplished. Some have
thought it was purely mechanical abrasion. others that it was due to the dissolving action of some secreted acid and, mom recently it is believed that
carbonic acid may be involved.
As a result of the action of the borers, including the purple sea urchin,
many of the softer intertidal rocks are full of holes and crannies in addition
to those which might normally arise from fracturing and weathering. These holes
and crannies furnish protected sites for a large array of species of varioua
groups. Some of them live within the ahells left by preceding species. Such have
been termed nestlers in distinction to the cranny dwellers whioh live in the
cavities of one kind or another in the rock. Sometimes we n q have a series of
successive nestlers. For example, a tiny young Kellia in a Saxicava Bhell, itself in a Protothaca shell which lies within the shell of Pholadidea, the
original borer.
The conditions concerned vary all the way from deep, almost entirely shut
off cavities to shallow ledges or surface cavities. We shall encounter conditions favorable for such studies in several localities. Some of the rocks in
the vicinity of the Second Sled Road are soft enough to permit of their being
occupied by borers. The same is true of many of the rocks at Frenchman18Reef
and especially of many near the lagoon. At Duxbury Reef also will be found situations for the study of these borers, neetlers and cranny dwellers.
W p notes concerning them as encountered at different times and make
lists of all the species by phyla which fall under these different categories.
Attempt to determine the food habits of all of them. Refer to varioue papers
by hcGinitie.
s

Exercise 28. Ecoloaical Distribution as Seen on the Sand Plats s&&&

e
b!ha Flats on t

Just inside Tomales Bay on the northeastern shores are extensive areas of
sandy mud flats already sampled in Exercise 21. Here wave action is large4
eliminated and sedimentation active. Here tidal currents while not violent,
exaept in certain relatively limited channels, are prolonged bringing an immense amount of water over a given area of bottom.
This area supports a fauna whioh seem to belong to the Macona fomation
as defined by Shelford and Tower (1925) with general and special modifications
due to local conditions. We shall'sample this fauna in general on this trip,
compare it with another tidal flat (Rerciae 29 on Bolinae Lagoon) and study

FIELD STUDIES

211

more intensively two of its subdivisions (Exercises 29 and 30), and finally
consider it in terms of ecologic variation and succession in Exercise 31.
In preparation for these exercises and during their prosecution make yourself familiar with the ideas and terminology in the article by Shelford and
Towler and read MacGinitielstlLittoralkrine Communities" and his "Ecological Aspects of a California &trine Estuary (Elkhorn Slough)."
The three important associes, studied separately, are at different levels
as regards the tidee. The eelgrass associes or dlcrcome-Schizothaerus associes
(perhaps same as the Macoma-Synapta associes of Shelford) is lowest, being
mostly a few inches under water at low low tide due in part to the very slow
run-off in this area. The Phoronopsis or Macoma-Phoronopsis associes is just a
few inches above low low tide level, while the Call ianassa or Macoma-Call ianassa associes, already studied, is much higher. These should be studied with
the idea of an ecological succession in mind whose climiix may be the sand dune
formation.
In narrow zones are other associes. Among these should be studied the
Clymenella associes which form narrow zones between the others. More broadly
distributed than the limits of these associes but less widely than Macoma is
the abundant horse clam Schiaothaerus nuttallii and the much less common geoduck, Panope generosa.
Study the distribution of the three types of clams, and of Phoronopsis,
Clynenslla and other common polychaetes. Turn in a map showing their distribution.
Exercise 29. Bolinas h o o n
This may be combined with a study of the fauna of Duxbury Reef (Exercise
27) since the tides are much later in the lagoon. It should be sampled at
least at two points, one very near the mouth where a Phoronopsis-Urechis-Macoma
association should be studied, the other high up the lagoon where the mudcrab is the dominant animal as is the case at numerous points along the lower
reaches of Tomales Bay.
Get a clear picture of the other members of the associes there and seek
to place them in terms of food chains or a food cycle.
Exercise 30. The Eekrass Flats
These outer flats near the mouth of Tomales Bay, separated from the shore
flats by a tidal channel are subject to fluctuation in level and cohsequently
in fauna and flora. Whether this is a seasonal variation or associated with the
extensive shifts in location of sand deposits which are caused by differences
in wind and tide is not hown. In 1935 they supported a luxuriant growth of
eelgrass (Zostera) and various seaweeds and were never completely exposed
even at the lowest tides. In the spring of 1940 they were a few inches above
low-tide level and the eelgrass and large algae were largely lacking.
Whatever their condition they should be studied and their fauna vvell
known. If these outer flats are lacking in extensive plant growth this may be
found and should be studied in lower areas near the shore, bordering on the
Macoma-Phoronopsi s association.
Go prepared for wading in several inches of mud and water. ,
Collect and make notes on the ecologic distribution of the rich but
monotonous fauna of this habitat. There are really three important habitats or
zones here, (1) the mud below, (2) the water, and (3) the luxuriant growth of
eelgrass and large algae ( ~ l v a ,etc. ). Do not neglect the fauna of the mud.
hterial will be collected and brought to the laboratory to allow for the study
of the sub-macroscopic fauna of the water and the algae masses and the eelgrass.
Be on the lookout for holothurians and Edwardsiella (an 8-tentacled anemone). Note those species present here only because the large seaweeds give them
sites above the muddy bottom.

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

212

Exercise 31. Phorono~sisand the Macoma-Phorono~sis Associes

laborators Studg: Phoronopsis represents a small group of uncertain systematic position, the Phoronidea. The U-shaped intestine and lophophore suggest
bryozoan affinities but the well-developed blood vascular system and the entirely different embryology render this relationship unlikely. We shall consider it a phylum as does Borradaile and Potts.
Study whole specimens dissecting to get an idea of the structure of the
lophophore, etc. Study serial sections to see the body cavity, the circulatory
system, the finer structure of the lophophore, etc.
On living individuals see the ciliary action of the lophophore.
Field Studies: Where the substrate has been built up (by ordinary deposition or ruud transportation) to the level of low tides the plant growths are
inhibited and here are to be found luxuriant clans of Aoronopsis, with Macomu
sacta less abundant.
The class will attempt a preliminary population study in this area. Working ih teams of two, two areas six inches square will be chosen by each team,
half of these so as to avoid any clam holes which perhaps crowd out a certain
number of Phoronopsis and certainly obscure their openings. Six-inch square,
open-ended metal boxes will be pushed in surrounding these areas and a careful
count made and checked, of the holes visible at the surface. The earth will
then be removed on all sides of the box and the number of individuals within
the box carefully counted. This should give the correction necessary to make
surface counts accurate. How do you explain the discrepancy, if any, both where
clams are present and where not? To what extent, if any, do the clams seem to
reduce the Phoron6ps is population?
Half of the teams will now measure off areas of a yard square and on the
basis of a count of the clam openings present, estimate the population of
clams, remembering that Macoma has two separate siphons, one smaller than the
other.
The other half will measure off areas of six inches square uaing the
metal boxes and after attempting to determine the number of clams involved
from surface counts carefully excavate to determine definitely the number aotually present.
Exploratory transects will be run to determine the limits of the partioular Phoronopsis clan and an estimate made of the number per acre (43,560
square feet)

Exercise 32. Eoolonic Succession

Study again the relation of the shore tide flats from the lowest algae
grown portion to the sand dunes with the possibility in mind that the sand
encroaching on the upper end of the bay is responsible for the change, locally
at least, (1) from Zostsra egsocies to the Macoma-PRoronopsis amocies, (2)
from this to the Clynenella aarrocies, (3) from it to the Macoma-Callianussa
associes, (4)from this to the Staphylinid-beetle aesocies and (5) from it
in turn to the sand dunes. Or is the succession in the other direction with
wind and current resulting in the encroachment of the marine on the sand dunes?
See again the special paper by Alice Mulford on the distribution of Call ianassa.

Exercise 33. Estuarine Faunas on Piles and Mud Flats

Estuaries or deep estuarine bays have characteristic faunas which are
rich in individuals but low in number of epecies. Here food is abundant and
wave action is negligible but inhibiting factors are (1) lowered salinity and
(2) great amounts of sediment and often of polluting materials in the water.
The study will be made at Inverness on the west side of the inner end of
Tomales Bay. Here pile^ offer surface for sessile animals and broad mud flats
afford favorable substrate for others which live within the substrata.

FIELD STUDIES

213

W e certain that you know all species involved and are able to discuss
their distribution in terms of ecological factors.
Fixercise 34. Commercial Oyster Beds at Bivalve on Tomales Biz
These beds will he visited to see the methods of commercial oyster culture, the ecological conditions favorable to it and the associated fauna.
Especially interesting are the introduced species.
Exercise 35. The Plankton

A surprising number of littoral marine invertebrates are plankton feeders.

No true understanding of the food relationships, food chains, and food cycles
is possible from the low tide investigations to which we are largely confined.
We shall therefore devote at least one period to a study of the floating population, the organisms concerned, and something of their relation to the animals
already studied.
Identify common forms to the group and seek to understand their place in
the food cycle. Supplementary aids to this end will be supplied. Refer also to
Johnstone, Scott and Chadwick, The k i n e Plankton.
Be on the lookout for larval forms which should be brought to the attention of the instructor for demonstration to the class.
Exercise 36- Individual Studs of a Small Tide Pool
See Exercise 15 for directions.
Exercise 37. Individual Studv of the Fauna of a Limited Area
of Rock Surface in the Intertidal Zone
As an alternative individual exercise if the tide pools are not available,
as is frequently the case due to movements of sand, study a limited area of one
of the rocks at Perch Rock Point or Second Sled Rock, or a limited especially
interesting area of beach or tide flat.
Exercise 38. The Fauna and Food Cycle of a
Freshwater Pond (Rolland Pond 1

A anall segregated body of water lends itself to this purpose much better
than the marine littoral with its tidal changes. Even here, however, there are
interrelations with nonaquatic or amphibious organisms which should be brought
into the food cycle so far as ascertainable.
Begin this study early in the course and continue it from time to time.
Read Exercise P on page 191.
T m in your food cycle as finally conceived.
Exercise 39. The Fauna of a Coastal Stream
(Salmon Creek or Uouer Stemole Creek1
This is to be an entirely individual problem.
The individuals should scatter out along the creek, each working alow.
W e a collection of macroscopic and microscopic life in connection with
notes on ecological features.
In your report place each species found in its individual niche in a food
cycle and attempt to explain the special adaptations of each species to its
niche. In Salmon Creek note especially the peculiar shrimps, Sycnaris (see
special paper by Lee).

214

LABORATORY AND FIELD TEXT IN I

~ ZOOLOGY
T
E

Exercise 40. Terrestrial Fauna

Terrestrial faunas in general are less rich in numbers of species and of
individuals than are those of the seashore. Also their constituent species
are less easily observed since the necessities of food-getting and protection
require either great speed or retiring life habits.
Conditions such as those furnished by a wooded canyon, where moisture
abounds and many types of cover are available, make for rel~tivelyrioh and
diversified fauna. Study such a fauna referring to page 192 for suggestions
as to the types to be expected.
We shall take advantage also of the near-by sand dunes to study terrestrial
life under much more severe conditions. Tnese conditions result in limitation
of the number of species to a few highly specialized types. The detection of
these would be a baffling problem indeed were it not that many of them leave
trails to indicate their activities. AP it is, we shall find it an intriguing
problem to determine the animals of this community, especially so, since I
know of no investigations of this fauna. For studies of sand dune communities
elsewhere, see chapter 19 of Chapman's Animal E c o l o ~or Pierce, Dwight and
Pool (1938. 1939 1.
h e kmero;s visits to the dunes, keeping a record of all trails and
animals observed. Some visits should be made at early morning and evening
hours and at least one at night, using flashlights. Excavate wherever trails
seem to end and elsewhere to determine the in-ground fauna in general. Gain
as complete as possible a picture of the conrmunity, its constituents and
interrelationships.
Individual Projects
Instructions as regards individual projects and special problems for S112
and S119 will be found on pp. 193 and 204.
There is great need for systematic, faunistic. and distributional studies
in the Dillon Beach area, the results of which will be of value to future
classes. The summer session course has been held here only once, in 1935, and
very much remains to be known of the fauna and its ecologic distribution. Following is a list of a few of the many problems which suggest themselves.

The pelecypods
The amphipods
The harpacticoid copepods
Other benthonic copepods
The isopods
The sponges

The tunicates
The Bryozoa
Phoronopsis: Are there two species?
Their distinguishing characters?
The terebellid polychaetes
The Nereidae
Faunistic

The sand-dwelling polychaetes

Polychaetes of the eelgrass flats
Polychaetes of the shore tidal flats
A survey of the shore tide flats eastward around the eastern shoulder of
Tomales Bay
The fauna of the sand flats in front of Dillon Beach
The fauna of the sand flats at Perch Rock Point
The fauna of the sandy beach frpm Dillon Beach to Sand Point
The fauna of Stemple Creek Lagoon
The fauna of the narrow-leaved eelgrass beds (Second Sled Road and elsewhere)

FIELD ST[TDIES

215

Ecolo~iaDistribution and Other Ecolonic Studies

The Cly~enellaassocies, its constituent species and distribution
The degree of intergradation between the Clymenella aseocies and other associes
and an attempt to explain the segregation of these associes
The relative abundance and distribution of Panope as compared to Schiaothaem
Schizothaerus ae a dominant animal and its influence on other members of the
association
Distribution of Roronopsi s and factors limiting it
Evidence as to the degree of correlation between color and pattern of Acmaea
digitalis and Acnaea scabra and that of their backgrounds (Balanus, Mytilus,
Mitella, etc. )
The smaller gregarious sea anemones (Bunodactis elegantissimu) of the Perch
Rock Point rockc Bre they different structurally from the solitary anemones
(~ntho~leura
xantho rammica)? b t are the ecologic features characteristic
of the sites of eac ? h t differences as to behavior? As to food?
Succession of animals settling on glass slides exposed below low-tide level at
Sand hint

APPENDIX A
SAMPLE SCHEDULES
ZOOLOGY 112: SCHEDULE. 1940
January
26. F.
27. S.
February

.....
Decapoda
..... Amphipoda

..... Pelecypoda
Animals at Point Richmond
.....
....
Field Trip to Point Richmond 2:00 p.m. at beach
..... Field
Trip to Point Richmond 2:00 p.m. at beach
F. ..... Free
S. ..... First Mid-Term including tests on identification and ecology
F.

S.
Su.
M

and behavior of animals of Point Richmond

.....
Polychaeta
.....
..... ~oeienterata
Field Trip to Fruitvale Bridge (be at bridge at 2:00 p.m. )
W. ..... Same as above at 2:30 p.m.
F. ..... Free
S. ..... Free
F. ..... Copepoda
S. ..... Entomostraca completed
F. ..... Rotifera
S. ..... Free
F. ..... Free
S. ..... Gastropoda
F. ..... Spring Recess
S. ..... Spring Recess
F. ..... Terrestrial Arthropoda
S. ..... Field Trip to Strawberry Canyon
F. ..... Second Mid-Term
S. ..... Field Trip to Lake bkrritt
F. ..... Free
S. ..... Field Trip to Freshwater Pond
F. ..... k i n e Gastro ods
S. ..... Continued: ~05chaeta
} . Field Trip to Dillon Beach for three tide8
S. ..... Free
F. ..... Free
S. ..... Property clearance
F.
S.
T.

uL-

1.
2.
8.
9.
15.
16.
22.
23.
29.
30.
April
5.
6.
12.

S112 AND 8119. SCElEDlJIE. 1939

&Y

...........

15. M.
Adjustment of room.,etc. 7:30 p.m. Preliminary meeting.
16. T. 8:30 a.m. S119 (alone) for orientation
9:00 a.m. S112 and S119. General introduction, Laboratory study
of Decapoda
2:00 p.m. Gastropoda
17. W. 8:30 a.m. Pelecypoda
2:00 p.m. Polychaeta

LABORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOC;JI

b y (Continued)

18. Th. (-0.5, 4:07 a.m. ) Trip on S.S. Scripps and study of plank-

nel
(-0.7, 8:06 a.m.) Special problems
Lecture by Prof. E. T. Schenck, Palaeontol2:00 p.m.
ogist, Stanford University
24. W.
(-0.4, 9:00 a.m.) S ecial problems
2:00 p.m.
ciitons
25. Th. (0.0, 9:40 a.m. ) Special problems
Lecture by Dr. Robert C. Miller, Director of
2:00 p.m.
the Museum and Aquarium of the California
Academy of Sciences on "The Pacific Ocean
a Balanced Aquarium"
26. F. 8:30 a.m.
Gestropoda completed
2:00 p.m.
Amphipoda, Isopoda
Nothifig scheduled
Nothing scheduled
29. M. 10:OO a.m.
Mid-Term
2:30 p.m.
Lecture
8:30 a.m.
Decapoda completed
30. T.
Lecture by Dr. Avery R. Grant on speciation
2:00 p.m.
in the genus Acmaea
31. W. (-0.6, 3:40 a.m. ) Special problems
23. T.

........
........
........

........
........
........
........
.......
.......
.......
.......

1. Th.
2. F.

-0.7, 4:25 a.m.

1-0.8, 5:00 a.m.
2:00 p.m.
3. S.
-0.8, 5:40 s.m.
4. Su. 1-0.6, 6:15 a.m.
-0.4, 7:00 a.m.
[-0.1, 7:40 a.m.
7. 1. 8:30 a.m.
2:00 p.m.
8. Th. 8:30 a.m.
2:00 p.m.
9. F.
10. S.
11. Su.....................
12. M. 8:30 a.m.
2:00 p.m.
13. T. 8:30 a.m.
2:00 p.m.

Special problems
]
Marine Vier Rocks and Seal Core
........ Polychaeta
Polychaeta of Moss Beach Flats
11 Unscheduled
Special problem

Special problem
........
Freshwater Ecology, Entomostraca
........ AOstracoda,
Cladocera
........
Freshwater Pond
........ Rotifera
....................
Special problems

.................... Free
Special problems
........
Terrestrial Arthropoda
........ Field
trip in Sunshine Creek Valley

........
Bryozoa
........ Porifera

23. F.

2:00 p.m.
8:30 a.m.

.......
checked in and packed
....... Property
Final Emmination

S112 AM) S119

TENTATIVE S~EEDULF:. 1941

19.
20.

21.
22.
23.
24.
25.

......................
Housing arrangements
7:15 p.m. .......... F i r s t meeting f o r general discussion
8:30 a.m. .......... S119 alone f o r orientation
T.
9:00 a.m. .......... S112. Introductory
9:30 a.m. .......... S112, S119. Decapoda
2:00 p.m. .......... Decapoda
W.
8:30 a.m. .......... Isopoda; 2:00 .m. Amphipoda
Th. 8:30 a.m. .......... Polychaeta; 2:{0 p.m. Polychaeta
F.
8:30 a.m. .......... Gastropods; 2: 00 p.m. Pelecypoda
S.
8:30 a.m. .......... Chitons
Su. (-0.3, 5:10 a.m.)
M.

Field t r i p t o Perch Rock Point. Ex. 21

Ex. 21 completed; 3:00 p.m. t e s t on fauna
of Perch Rock Point
6:50 a.m. + 1 ) Field t r i p t o t i d e f l a t s , Ex. 27
7:30 a.m. )
Field t r i p t o Second Sled Hoad, Ex. 22
a t 8:10 a.m.
Individual projects
a t 9:00 a.m.
Individual projects
5:40 a.m.
6:15 a.m.

-0.5,
(-0.5,

28.
29.
30.
31.

W.
Th.
F.
S.

(-0.6,
-0.6,
-0.4
-0.3

1.
2.
3.
4.

Su.

W.

5.

Th.

6.

F.

a t 9:50 a.m. )
a.m.
Mid-Term; 2:30 p.m. Bryozoa
a.m.
Sponges; 2:00 p.m. Pelecypoda
a.m.
Copepoda; 2:00 p.m. Cladocera and other
Branchiopoda
8:30 a.m.
Laboratory portion of Ex. 20 on Call ianassa, Upo ebia, and h s r i t a ; 2:00 p.m.
~oelenterafe'es
8:30 a.m.
Coelenterates; 2:00 p.m. Gastropoda
-0.7 a t 3:35 a.m. ) Individual projects
-1.2 a t 5:15 a.m.
Field t r i p t o t i d e f l a t s , Ex. 29
4:28 + 501)
F i e l d t r i p t o T o m a l e s P o i n t , E x . 23
Field t r i p t o Bird Island, Ex. 24
Second f i e l d t r i p t o Tomales Point

June
M.
T.

(-0.1
9:30
8:30
8:30

..........
..........
..........
..........
..........

........

13.
14.
15.
16.
17.
18.
19.
20.
21.
22.

F.
S.

Su.

................ Field
tri t o Frenchman's Reef,
~ndividua! projects

45).
t-0.7 a t 8:25 a.m.
-0.2at9:15a.m.!

Individualprojects

..........
of polychaetes
.......... Identification
Ostracoda and Rotifera; 2:00 p.m. Rolland
Pond Ex. 37
W.
8:30 a.m. .......... completion of study of freshwater pond
Th. 8:30 a.m. .......... T e r r e s t r i a l arthropods; 2:00 p.m. Sand-dune
animals, Ex. 39
I?.
8:30 a.m. .......... Further on t e r r e s t r i a l arthropods
S. ...................... Individual projects
M.
T.

8:30 a.m.
8:30 a.m.

Su.

Individual projects
Tideflats,Ex130and31
Optional f i e l d t r i p t o oyster bed, Ex. 33
Optional f i e l d tri t o Duxbury Reef and
Bolinas Lagoon,
26 and 28; 2:30 .m.
p r a c t i c a l -nation.
Property checE
10a.m.Final~ination

&.

27.

Ek. 25

F.

(-0.6,7:10a.m.)

APPENDIX B.
TT3E CLASSIFICAEON OF I.NVER-TES
FOLLOWED I N TBIS TEXT
(Modified from Hgman, Borradaile and Potts, and others)
Phylum PROTOZOA1
Classes MASTIGOPBORQ
SaRCODINA
SrnRGZOA

msom

Phylum PORIFERB

Phylum COEGWEEtAllA
C l a s s EIYDROZQA
Order EIYDROIDA

Order S i ~ h o n o ~ h o m
Three other orders (see &man)
Class ScYPHOZOA
Class ANTHOZOA
Subclasses ALCYONARIA
ZOANTBARIB (Sea Anemones )
Phylum CTENOPHOU
Phylum p
m
Class TORBEI,LlmA

Classes TREUTODB
CrnTODA

Phylum Nemat omorpha

'Explanation of meaning of four d i f f e r e n t ways of p r i n t i n g group names:

1. Those groups i n UNDERLINED CAPITALS a r e given s p e c i f i c treatment i n the ex-

e r c i s e s , o r form an e s s e n t i a l p a r t of t h e c l a s s i f i c a t i o n of such groups.

2. Those groups i n CAPITALS a r e not given s p e c i f i c treatment but should be

known t o a l l students of i n v e r t e b r a t e zoology. I n some cases, such a s

the Protozoa, these a r e very important groups which a r e not considered
because they form the subject matter of other courses. I n other cases
the groups a r e of moderate importance, but a r e r a r e l y encountered i n the
c o l l e c t i o n s d e a l t with here; i n s t i l l other cases t h e animals a r e import a n t but these group names a r e l i t t l e used.
3. Those group names i n underlined lower case r e f e r t o animals given i n the
l i s t s but e i t h e r t h e animals a r e of subordinate importance here o r these
group names a r e l i t t l e used.
4. Those groups i n lower case and not underlined do not appear i n t h e exerc i s e s o r l i s t s , but a r e included h e r e , because, i n most cases, they a r e
of equal taxonomic rank with better-known groups.

219

-+s

t
220

LABOMTORY AND FIELD STUDIES IN INVERTEBRATE ZOOLOGY

Phylum Acanthocephala
Phylum ANNXLIDA
Class Archiannelida
Class
given in keys and list)
Class 0 GOCHAEW
Classes HImINEA
ECHmROIDEQ
Phylum ONYCHOPHORA (or subphylum of Arthropods)
Phylum ARTKROPODA
Subphylum CRUSUCEQ
Class BRAN~OPODA
Orders Anostraca
Notostraca
Class
class

&iRiHP

tzBEaP

Class CIRRIPEDIA
Orders Thoracica
Acrothoracica
Apoaa
Rhizoce hala
a&
Class WLACOSTRACA
Subclasses Leptostraca
Hoplocarida
Syncarida
Subclass Pericarida
Orders

&&siea

Tanaidacea
Order ISOPODA
Suborders
Order ,AWEIPODA
Suborders G m D U
a-pgB
Ca~rellidea
Subclass Eucarida
Order Eu hausiacea
Order D E ~ O D A

APPENDIX B

CLCLSSIRCA'PIONS OF THE DECAPODP

Parker and Haswell

Borradaile and Potts

Pratt; Schmitt
&borders:
Sections:

Subphylum MYRUPODA

Sub hylum INSECTB

(givisions too numerous to give here-see
Subphylum BRBCHNIDA
Classes p~o~ionida
1p osura
Acarina
r e h ; ; % % @

texts)

terrestrial classes)

Subphyla (? ) Tardigrada
Pentastomida
Phylum MOLLUSCA
C~SB
Orders POLPPLACOPHORA
Aplacophora
Class WTROPODB.
Order STREFTO=
(Prosobranchiata. prosobranchs, or
strepotoneurous gastropods)
Suborder DIOTOCARDIA (A~~idobranchiata.
Aspidobranoha)
Tribe ~iaidoalossa
chiata (with paired gills;
S e ~ ~ ~ ~ r keyhole
a n limpets)
~ s ,
Section Azygobranchiata (with single gill;
azygobranchs; Ternla, Calliostoma)
Tribe Docoalassa (true limpets, Acmaea)
Suborder M0NOMGBRDI.A (Pectinibranchiata; pectinibranchs: all other snail-like forms)
Order OPISTBOHRBNCHIllTA (opisthobranchs)
Suborder TECTIBRANCHIAW (tectibmchs: sea hares )
Suborder NUDIBRANCHIATA (nudibranchs)
Order WLUANATA (pulmonates: land snails and slugs)
(Opisthobranchiata and Pulmonata are considered by some
to be suborders of the order Euthyneura)
Streptoneurnus vs Euthyneurous
Prosobranchs vs Opisthobranchs
Pectinibranchs vs Aspidobranche

CJ~~osedTerms
Zygobrancha vs Azygobranchs
Tectibranchs vs Nudibrancha
Pulmonates vs Gill-breathing gastropods

222

LBBORATORY AND FIELD TEXT IN INVERTEBRATE ZOOLOGY

Class SCAPHOPODA
Class PELECYPODA
Orders Profobranchiata
Filibranchiata
Eulamellibranchiata
Septibranchiata
Class CEPHALOPODA
Phylum I(BMPTOZ0A (Endoprocta)
Phylum BRYOZOA
Orders P lacto emata

l!%a&h-

Phylum BRACHIOPODA
Phylum Chaeto natha
phylum PHORO~DU
Phylum ECXIN0DEFWT.A

Phylum CHORDATA
Subphylum HEMICHORDATA
Subphylum DROCHORDATA
Classes Larvacea
Ascidiacea
Thaliacea
Subphylum CEPHALOCHOFDATA
Subphylum CRANUTA
Class PISCES
Class Amphibia
Class Re tilia
Class A&
Class M
A
M
M
u
A
,I

APPENDIX C.
ATTEUT AT A FIELD KEY TO SOME OF !PEE COWOIWR ALGAE OF MOSS BEACH
(For further identification see mounted specimens and monographs by Setchell and Gardner)
A. Green Algae
B. Brown Algae
C. Red Algae (oftentimes blackish or brownish)
A. Green Akae

. . . . . . . . .. .. .. .. .. .. .. .. .. .. .. .. .Ulva
.2

1. Made up of slender filaments

1. Broad, flat, thin, bright green. Sea Lettuce
2. A fine, green felting over tops of rocks covered only at high tide

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enteromorpha
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .? Cladophora

2. Small, but thick, felted masses; common on Moss Beach Flats, scattered

B. Brown Ahme
1. With naked,stout, cylindrical sti e or nstem," blades only at its apex;
2
found chiefly in outer zone of freakers
3
1. No conspicuous unbranched stipe
2. Stipe less than two feet long, erect, with a crown of many short blades.
Sea Palm
Postelsia palmaej'onis
Laminaria
2. Stipe averaging two feet or more, with a single long blade
2. Stipe very long, up to 60 feet, stipe hollow and ending in a swollen,
Nereocystis
hollow bulb; many long blades
3. Unilateral branching along one side of long cylindrical stipe; blades
broad, with a bulb-like float at base; edges of blades with scattered
tag-like outgrowths
Macrocystis
3. Bilateral branching along sides of flattened rachis; blades small;
Egregi a
interspersed with bladder-like floats
3. Dichotomously branohed fronds; branches less than a centimeter wide

. . . . . .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..
....................
...
...............
.....................
..............

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . ficus fircatus
1. Fronds cylindrical . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1. Fronds more or less flattened . . . . . . . . . . . . . . . . . . . . . 3
2. Forming low, stiff reddish-brown masses, seldom more than an inch in
height; densely branched, covered with very tiny, spinose branches
Gndocladia

...............................
.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gastroclonium
3. Fronds very broad, little-branched, iridescent in color
. . . . . . . . . . . . . . . . . . . . . . . . . . . Iridopsis (Iridaea)
3. Fronds narrower, much branched . . . . . . . . . . . . . . . . . . . . 4
2. Fronds blackish, elongated, hanging, several inches in length, ultimate
branches several millimeters long, all of about same length Microcladia
2. Fronds longer, more open, ultimate branches thicker, scattered

4. Fronds only slightly flattened, finely branched and delicate, bright

...................
...........

red; often epiphytic

.Microcladia
4. Fronds flattened, dichotomowly branched, varying from broad red blades
to narrow, much-branched brownish blades
Gigart ina

LIST OF REFERENCES
GENERBL ANATOMY AND CLASSIFICATION OF INVERTEBRATES
k r r a d a i l e and Potts. 1935. T
he(probably the best one-volume
text )
Bronn. Klassen und Ordnwen des Tierreichs (many volumes, a large part of
wh'lch remains t o be published
Cambritke Natural H
1895-1969 (several volumes)
throw& Cteno~hom.
Byman, L. H. 1904. Z E i e r t e b r a t e ! .
vol. 1.
(other volumes t o be published
K i i k e n t W. 1923 t o date. Handbuch der Zoo1 '
several volumes-many parts
yet t o be published: bibliographies v e r y E d
Lankester, E. Ray. 1900-09. A t r e a t i s e of zoolo
several volumes )
Parker and Haswell. 1940. Textbook of zooloa. g v i s e d by Lowenstein. Vol. 1.
Invertebrates.
Yierreich. Das. A series of'systematic monogra hs on various parts of the
animal king om, published during the past 0 years)

GENERAL OR BMERICAN FAUNA

Johnstone, Scott and Chadwick. 1924. The marine ~ l a n k t o n
Morgan, A. H. 1930.
Needham and Needha
Pratt, H. S. 1935.
Ward and Whipple. 1918. Freshwater bioloay
PACIFIC COAST FAUNA
Guberlet. 1936.
of the seashore
Johnson and Snook. 1927. Seashore of the
Ricketts and Calvin. 1939. &tween Pacific tidea

..

ECOLOGY-4-L
Allee, W. C. 1938. The social l i f e of animals
Chapman, R. N. 1931. Animal e c o l o ~
Crowder, W. 1928. A naturalist a t the seashore
Ekman, S. 1935. Tieri?eo~raphiedes k e r e s
Elton, C. 1927. Animal ec l o
Flattely and w
a
l
d biolow of the seashore
Hesse, Allee and Schmidt. 193
co o i c a l anima eo ra
r. Midland
MacGinit ie, 6. E. 1938.
l?tt ; ~ lcommunitte~,
ist (see bibliography)
Pearse, A. S. 1939. Anlmal ec o
Shelfod, V. E. 1 9 2 9 d a n d f i e l d ecologp

"Marid.

ECOLOGY-SPECIFIC

LOWJTIES OR SITDATIONS

.,

Allee, W. C. 1923. Studies i n marine ecolow. I and 11. Biol. kl

%
Allee. 1. 0. 1934. w ~ o n c e r n the
i ~ ormmization
of marine coastal'conarmnit i e s , "
-

communities," Amer, Midland Naturalist,

Hewatt, N. G. 1937. "Ecological studies on selected marine i n t e r t i d a l communit i e s of b n t e r e y Bay, California," Amer. Midland Naturalis ,l!2
MacGinitie, G. E. 1935. nEcological aspects of a California marjne estuary
(Elkhorn Slough),"
r. Midland Naturalis
1937. "!be use of mucus
by marine plankton E d e r s , "
&: k 8 - 1938. "Notes on the natur a l history of some marine animals,"
Newcombe, C L. 1935. "Certain e n v i r o n m e n t % a c ~ c h* ,

m,

r. &I@
23 .,
Pierce, Dwlght, and Pool.
sand dunes. I-V,

s.

J8

1938, 1939. The fauna and f l o r a of the El S e p d o

p
.

'1

P'
i

LIST OF REFERENCES

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Shelford and Tower. Animal communities of San Juan Channel and adjacent
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PORIFERA
de Laubenfels, M. W. 1932. "The marine and fresh water sponges of California,"
Pmc. U.S. Nat. ms., 81. 1936. "A discussion of the sponge fauna of the
Dry Tortugas, with material for a revision of the families and orders of
the Porifera Cam. Inst. Publ., No. 467.
COErnTERATA
Hsdrozoa
Fisher, W. K. 1938. "Hydrocorals of the North Pacific Ocean," Proc. U.S. Nat.

."

A!&?*, 84

Fraser, C. McL. 1937. Hydroids of the Pacific Coast of Canada and the United
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.,reyaM
1910."Medusae of the world. I. The Hydromedusae." Cam. Inst. Bull.
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Scmbzoa
Inst.
Mayer A. G. 1910. nMedusae of the world. 11. The Scyphomedusae."
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Anthozoa
Harrison, A. H., Zool. 112 Report. A classification of the sea anemones of the
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Kemp, N. E., 'Lool. 112 Report'.The anemones of the Moss Beach area
pLA-TBES
Turbellaria
Boone E. S. 1929. "Five new polyclads from the California coast," Ann. bkq.
fiat. Hist., ser. 10, vol. 3
Heath and McGregor, 1912. "New polyclads from Monterey Bay, California,"-.
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Coe, W. R. 1904. Nemertinea, Harriman Alaska l?medition, 2 (pts. 1,2).

1905. "Nemerteans of the west and northwest coasts of America," bll. hs.
Com~.Zool., Harvard, 47. 1940. Revision of the Nemertean fauna of the
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(For identification of freshwater rotifers, see Ward and Whipple or
Pratt, Revised Edition)
u.s* Nat*
Harring, H. K. 1913. "Synopsis of the Rotatoria,"
Hudson. C. T. and Doas, P. H. The Rotifera. 2 volsyl886. Supplement, 1 9

Hartman O., Ph.D. Dissertation. P 1 chae ous annelids of the li toral z ne of

~aiiforn'a. 1936. "Nomencla trTa ct es invo ving a i okia o & m e t e
d
u
. !ash. Acad: So!.,
%6
and dter. !kp;rs on
Families Spionidae! Nereidae, Eunicidae, Stauronereidae, Opheliidae,
Phyllodocidae,"
a 'f
b
ool., 41. 1936. New s ecies of polychaetous annelic8tEA&ig kZidae E m ~alifornia,%roc. U.S ~t
83. 1938. "A revision of the Nephtyjdae from the no heast Pacific,"
Pmc. Ts.
Nat. &IS., 85. 1939. "New species of polychaetgs annelids
from Southern California," Alan Hancock Pacific Expeditions, 2. 1939,
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an
,"
cock Pacific Emedi&ggl 5 1941.nSome contrihiionsd
th%oey
and life history of
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Newby. W. W. 1940. "The embryology of the echiuroid worm Urechis caupo ," k r .

2.

fd
. .

u.,

::

LABORATORY AND FIELD TEXT IN INVERTEmTE ZOOLOGY

226

SIP[TNCIJMIDEIL
Brown, H. E., Zool. Report. The si~unculidsof the Moss Beach. California. area
Fisher, W. K. 1928. "New sipunculoidea from California," Ann. k. Nat. Hi-st.,
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Crustacea. Eutomostraca
(For freshwater Entomostraca, see in addition to those iven below,
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Cladocera
Johnson,.D.,Zool. 112 Report. Freshwater Cladocera of the San Francisco Bap
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~uda~;ih907. "Cladocera of the San Diego region (mrine)." Univ. California
Publ. Zool., 3
Ostracoda
Juday, C. 1906-7. Ostracoda of the San Diego region (marine). Univ. Calif.
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Skogsberg, T. 1928: "Studies on marine ostracods," Occ. Papers. California
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Coae~oda(kine )
Esterly, C. 0. 1924. The free swimming Copepoda of San Francisco Bay. U a .
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Fraser, J. H. 1936. 'nhe occurrence, ecology, and life history of Tigriopus
,fulurs (Fisher)," Jour. W. Biol. Assoc., 20. 1936. "me distribution of
rock pool copepods according to tidal XeveT;" Jour. Animal Ecol., 5
Light and Hartman. 1937. A review of the genera Clausidiun and Hemicyclops.
Univ. Calif. Publ. Zool., 41
-Monk, C. R. 1941. "Marine harpacticoid copepods from California," Trans. Amer.
Micr. Soc., 60
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T
American parasitic Copepoda," Pmc. U.S. Nat W.,
35. 1932. The Copepoda of the Woods Hole region, Massachusetts.
Nat. Mus., 158 (both marine and freshwater). 1935. "Parasitic copepods from the P a z i c Coast," Amer. &&land Naturalist, &
Various special reports on the red tide pool copepod, T i g r i opus c a l i fornicus
(= T. &lous or T. t r i a n p l u s )
Co~eaoda(Freshwater)
Coker, R. E. 1934. "Contribution to knowledge of North American harpacticoid
freshwater copepod Crustacea," Jour. Elisha Mitchell Sci. Soc., 2
111%. Paul. Ph.D. thesls. Csclouoid ToDepoda of California
Kiefer, F. 1929. "Cmstacea Copepoda. 2. Cyclopoida Gnathostoma," in Das Tier-

manual^

.
u.'

m.

subgenera and species of diaptomid copepods, Univ. Calif.

1939. New American subgenera of Diapt onus. Trans. Amer

and key of the North American copepods of the

Nat. b.,
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Branchiura
Meehean, 0. L. 1940. "A review of the parasitic Crustacea of the genus A r p l u s , "
Pmc. U.S. Nat. b.,
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Cirri~edia
Cornwall, I. E. 1935. "On the nervous system of four British Columbian barnacles,"
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U.S. Nat hs. , 88. lE0. Cirripedia of m e t Souna with a key to ~ e ~ a ~ e c i eJJiir.
s . Wasr Publ. Oceanoq-

1
I

:.'

&*

II

LIST OF REFERENCES

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Pilsbq, H. A. 1916. "The sessile
Shelford, ewer, Worley, Rice. 193
Sound Biol. Sta., 1
-

ISODO~~
Holmes, S. J. 1909. "Four new species of isopods from California," Proc. U.S.
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~ o f o i m r3
, al. 1927. Limnoria and its allies, the crustacean borers,
in Hill and Kofoid, Marine borers and their relation to marine construction on the Pacific coast
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omparative ecological studies on the terrestrial isopod
Cru:iakal%?iz
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Richardson, H. 1905. Monograph on the isopods of North America. EhllXl.~.
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54
Van &me, W. G. m36. The American land and freshwater isopod Crustacea. &
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)her. h.Nat. Hist., 11
Am~hi~oda
Alderman, A. L. 1936. Some new and little known amphipods of California.
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H O ~ B
l1
phi@cis collected by the 'Albatross' off the west coast
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E k a ? R c. U.S. Na
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Rodholm, A. K., M.A. thesis0 copy in Z~ll?coil~tion). Contributions to the
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Shoemaker, C. 1925. The Amphipoda collected by the "Albatross" in 1911. Bul
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Stebbing, T. 8. 1906. The Bmphipoda. 1. Gammarid%, 1n-s
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228

Terrestrial Arachnids
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Ellsworth, J. d. Zool. 112 Report. A key to the local millipedes
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Needham, J. G., and P. R. Needham. 1930. A wide to the study of fresh-water
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Hungerford, H. 8. 1919. The biology and ecology of aquatic and semiiquatic
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Lloyd, J. T. 1921. The biology of North American caddis fly larvae. kll. Llo~d

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A. Claassen. 1925. A monoara~hof the Plecootera of America

B. Heywood. 1929. A handbook of the draaonflies of North

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Needh-G.
and J. R. Traver. 1935. The biolons of masflies with a s~stemtip
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Chamberlin, J. C. and G. F. Ferris. 1929. On Li arocephalus and allied genera
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Michener, Chas: D. 1939. Zool. 112 report. Some intertidal insects of the Moss
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Van Dyke, E. C. 1918. New inter-tidal rock-dwelling Coleoptem from California.
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MOLIJJSCA
General
abrid~edcheck list of west North American Mollusca
Keen, A. M. 1937.
Kee and Bailey. 1 3
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Schenrand Een. 1936. "hrine Molluscan provinces of west North America,"
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Tryon and Pilsbry. 1 8 m . G u a l of concholow. structural and ssstemtic

Berry S. S. 1917, 1919. "Notes on western American Chitons," Proc. Calif.

a d S i . ser. 4: vols. 7, 9
Shelden, F., 001. 112 Report.

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Bonnot, P. 1930. Ahalones in California.
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Grant, A. R., Ph.D. Dissertation (copy in
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Turner, I?.
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Vokes. H. E.. Zool. 112 Re~ort The smaller Gastrouoda from the littoral zone
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Villee, C. A. and T. C. Grood 1940. "The behavior of limpets with regard to
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Hewatt, W. G. 1935. l'Ecologicalsuccession in the Myt ilus ?a1i f o r n i a w s habitat,'? fc9hB$, 16
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Ceuhalopoda
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CHAETOGNArn
Michael, L. L. 1911. Classification and vertical distribution of the Chaetopatha of the San Diego region. Univ. Calif. Publ. Z A . , 8
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Bullock T. H., Zool. 112 Re ort. Phoronidea from Marin County
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Verrill, A. E. 1914. bnogra h of the shallow water starfishes of the northern

Pacific Coast from the Lctic Ocean to California. E&rriman Alaska Ser~err,
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Clark, H. L. 1911. North Pacific ophiurans. Bull. U.S. Nat. h.,
75
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Tunicata
Ritter, W. E. 1913. "The simple ascidians from the northeast Pacific in the
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Fishes
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Jordan, D. S. 1925. Fis es

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Birds
Hoffman, Ralph. Guide to the birds of the Pacific States

Dillon Beach, 185, 205

Diplopoda, 117

Amphipoda, 93; key to Gammaridea, 95;

list of, 102; of Point Richmond,
Amphipods, see Amphipoda, 93
Armelidti, 28; of Point Richmond, 18
Anthozoa, see sea anemones, 19
Arachnids, 118
Araneida, 118

Calyssozoa, 156

Echiuroidea, 65
Ecologic succession, 212
Ectoprocta, 156
Eelgrass, broad-leaved (Zostera), 211;
narrow-leaved (Phyllospadix), 197,
209
Emerita, 207
Enteropneusta, 165
Entoprocta, 156
Eucarida, 104
Field trips, Bird Island, 209; Bolinas
Lagoon, 210; Dillon Beach, 185;
Drake's Cave, 200; Duxbury Reef,
209; Frenchman's Reef, 197; freshwater pond, 191; Fruitvale Bridge,
190; Halosaccion Flats, 182, 196;
Iridaea Channel, 198; Inverness,
212; Lake Merritt, 188; list for
Dillon Beach, 206; Marine View
Rocks and Seal Cove, 199; Moss
Beach Flats, 195; Nye's Rock, 199;
of the spring semester, 177; oyster
beds 213; Perch Rock Point, 208;
point Richmond, 177; Sand Rocks,
196; Second Sled Road, 209; Strawberry Can on, 192; tide flats (Tomales ~ay? 210, 211, 212
Field work, directions for, 174; field
notes, 175; field trips, see above
Flatworms, 22
Frenchman1s Reef, 197, 210
Fruitvale Bridge, 190
Garden centipedes, see Symphyla, 117
Gastropods, 130
Gastropods, 130; key to freshwater
snails, 146; list of gastropods,
147; key to land slugs, 147; keg to
marine shelled, 123; key to nu&-

Cladocera, 70; key to genera, 71

Classification, 1, 219
Coelenterata, 11

231

232

LABORATORY AND FIELD TEXT IN lNV3RTEBRATG ZOOLOGY

Nemertea, 23; Pelecypoda, 150;

Polychaeta, 34; Porifera, 9; Pgcnogonida, 119; Rotifera, 26; sea anemones, 19; Sipunculoidea, 65; tunicates, 166; use of, 3
Lake Merritt, 188
Leeches, see Hirudinea, 65
Macoma, 210, 211, 212
Malacostmca, 84
Mammals, 173
Marine View Rocks, 199
Millipedes, 117
Mites, 118
Mollusca, 125
Moss Beach, 195
Myriapoda, 117
Mysidacea, 84
Nemertea, 23; key to, 23; list of, 25
Nemerteans, see Nemertea, 23
Nemertinea, see Nemertea, 23
Nestlers, 210
Nudibranchs, 144; key to, 144
Nye's Rock, 199
Octopus, 125
Oligochaeta, 64; key to earthworms, 64
Oniscoidea, see Isopoda, 84
Ostracoda, 73
Paroctopus, 125
Pelecypoda, 150
Pelecypods, 150; key to, 150; list of,
155
Peracarida, 84
Perch Rock Point, 208
Phalangida, 119
Phoronidea, 161
Phoronis, see Phoronopsis, 161
Phoronopsis, 161, 210-212
Phyllospadix, or narrow-leaved eelgrass, 211
Pisces, 169
Plankton, 213

Schedules, 216
Schizothaerus, 210, 211

Scientific names, use of, 4

Scorpionida, 118
Scorpions, 118
Scyphozoa, 11
Sea anemones, key to, 19; list of, 21
Sea cucumbers 161
Sea stars, 161
Seal Cove, 199
Second Sled Road, 209
Siphonaphora, 11
Sipunculoidea, 65; key to, 65
Slugs, key to, 147
Snails, 133; key to marine snails,
133; key to genera of freshwater
snails, 146; key to shelled land
puhnates, 146
Special problems, 193, 204, 214
Spiders, 118
Sponges, 6
Starfish, see sea stars, 161
Strawberry Canyon, 192
Summer Course, 176, 193
Symphyla, 117
Tanaidacea see Chelifera. 84
Taxonomy, 5 , Appendix B, 219
Terrestrial arthropods, 116
Ticks, 118
Tigriopus, 203
Tomales Bay, 185
Tomales Point, 185, Maps IV, V
Tooth-shells, see Scaphopoda, 125
Tunicates, 165; key to, 166
U ogebia, 207
Achorda, 165
Valvifera, see Isopoda, 84
Vernacular names, see common names, 4
Vertebrates, 169
Zostera, broad-leaved eelgrass, 211