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LIBRARY 
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ULLETIN OF THE »oce 


GARDEN 
Southern California 
Academy of Sciences 


LOS ANGELES, CALIFORNIA 


Vot. 61 January-Marcu, 1962 Parr 1 


CONTENTS 


Life History Notes on Epiplatymetra grotearia Packard. John 
Adams Comstock 

The Significance of the Petroliferous Nodules of Our Desert Moun- 
tains. W. Dwight Pierce 

A New Pasiphaea (Crustacea, Decapoda, Natantia) from Southern. 
Californian waters. John C. Yaldwyn 

Nonmarine Molluscs from Recent Sediments near Vernon, Apache 
County, Arizona. Robert J. Drake 

Records of Snakes from Joshua Tree National Monument, Cali- 


fornia. Richard B. Loomis and Robert C. Stephens 

Monarchistic Dominance in Small Groups of Captive Male Mos- 
quitofish,Gambusia affinis patruelis, Melba C, Caldwell and 
David K. Caldwell 


G. W. Horn’s Land Gastropod Locality in Arizona. Robert J. Drake 

Dependence on Temperature of Ca/Mg Ratio of Skeletal Structures 
of Organisms and Direct Chemical Precipitates out of Sea 
Water. George V. Chilingar 


Report of a Scyphozoan Stephanoscyphus simplex Kirkpatrick from 
the Arctic ocean. Carolyn Brahm and John L. Mohr 


Issued May 11, 1962 


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Southern’ California 
Academy of Sciences 


OFFICERS 


Theodore Downs). 21M 6b saa. he en ae hc Op er President 
Richard! Belioomilisy 6 sea. 8. << chalin oe eee ee ee First Vice President 
Mol maPAG Wilber ee ae Re geo Second Vice President 
Gretchen: Sibley) 1.0.) on ae eG tins, sets Secretary 
loi IVE arin sy. cee eter ae ae teh Slee met teeta Assistant to Secretary 
WY, Dwight Pierce™ ic!) is /.ahs ie 60 a eens Ve ee: er Treasurer 
David Ke Caldwell. 20), 22/508. a2 se Napa. ae erates, ene ree Acting Editor 
DIRECTORS 

Charles Burch Lloyd M. Martin 

John A. Comstock W. Dwight Pierce 

Theodore Downs Gretchen Sibley 

Hildegarde Howard © Ruth D. Simpson 

Richard B. Loomis Fred S. Truxal 


John A. White 


ADVISORY BOARD 


John L. Baird Theodore Payne 
J. Stanley Brode Kenneth E. Stager 
A. Weir Bell Louis C. Wheeler 
Russell E. Belous Richard Stone 
Thomas Clements Richard H. Swift 
Dorothy Martin Peter P. Vaughn 
Charles A. McLaughlin Sherwin Wood 


STANDING COMMITTEES 


Finance Publications 

Russell E. Belous, Chairman ' Richard B. Loomis, Chairman 
Conservation : Hospitality 

Donald D. Shipley, Chairman Fred S. Truxal, Chairman 
Library Membership 

Dorothy E. Martin, Chairman Jay M. Savage, Chairman 

Junior Academy 
Laurel Woodley, Chairman 


SCIENCE SECTIONS 


Anthropology Botany 
Charles E. Rozaire, Chairman Richard G. Lincoln, Chairman 
Earth Sciences Experimental Biology 
Peter P. Vaughn, Chairman John L. Baird, Chairman 
Invertebrate Zoology Physical Sciences 
Elbert L. Sleeper, Chairman George V. Chilingar, Chairman 
Vertebrate Zoology 


Dennis G. Rainey, Chairman 


OB RIES OU WL UU) 


WHEREAS the Southern California Academy of Sciences has re- 

gretfully accepted the resignation of 
DR. JOHN ADAMS COMSTOCK 

as Editor of the BULLETIN 

AND WHEREAS Dr. Comstock joined this Academy on June 23, 
1919, was elected a member of the Publications Committee immedi- 
ately thereafter, and has served continuously since then as Editor of 
the BULLETIN 

AND WHEREAS he has he and loyally served this Academy in 
many other official capacities, to wit: Secretary, 1921-1926; President, 
1926-1927; Treasurer, 1937-1939; Secretary-Treasurer, 1939-1948, 
and member of the Board of Directors since June 17, 1920 

AND WHEREAS under his editorship the monographic series 
known as the Memoirs of the Academy was introduced in 1938, and 
six numbers have been published 

AND WHEREAS he has contributed to the BULLETIN many out- 
standing scientific articles on the subject of Lepidoptera, accompanied 
by meticulously rendered illustrations of life histories drawn from life, 
all of which have enhanced the reputation of the BULLETIN as a 
scientific journal 

AND WHEREAS he has led other scientists into active work for the 
Academy by his verbal eloquence, his congeniality, and his own ex- 
ample of devoted service 


Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


THEREFORE BE IT RESOLVED that the Southern California 
Academy of Sciences hereby expresses to Dr. Comstock the sincere 
thanks of all members and officers who have profited by his long, out- 
standing service as an eloquent literary spokesman in the promotion of 
scientific knowledge, and extends to him hearty wishes for his continu- 
ing good health and success in scientific achievement. 


to 


THEODORE Downs, President 
GRETCHEN SIBLEY, Secretary 


LIFE HISTORY NOTES ON EPIPLATYMETRA GROTEARIA 
PACKARD 
JoHn Apams Comstock 
Del Mar, California 


During the summer field work of the Los Angeles County Museum 
group of entomologists in the Tonto Creek area of Gila County, Arizona, 
we were able to obtain eggs of Epiplatymetra grotearia, on July 1, 1956. 
These were reared to maturity, which made possible the following rec- 
ord of its life history, no portion of which has heretofore been published. 


Eco 


Barrel-shaped, with rounded ends. Length, 1.1 mm, Width, 0.75 to 
0.89 mm. The eggs were laid on their sides. In captivity they were 
placed in a single row. 

The color is a delicate ivory-green when first laid, changing later to 
ivory-white, speckled with small brown dots and dashes, as shown on 
Figure 1. At first there appears to be a number of faint longitudinal 


Figure 1. Ege of Epiplatymetra grotearia. Enlarged X 50. 


lines on the surface of the egg, barely discernible under X 16 magni- 
fication. Later these lines become indistinguishable. 

There is apparently a cap-like micropyle at one end of the egg, but 
this is faintly indicated in only a few examples, and is absent in the 
others. 

The 18 eggs in our original lot all hatched on July 14, 1956. The 
young larvae were tried on leaves of oak, peach, cherry, pine, rasp- 
berry and willow. They accepted the willow, and several were carried 
to maturity on it. 


4 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


First Instar Larva 


The newly emerged larvae were long and cylindrical, of the charac- 
teristic “looper” type. They averaged 5. mm. in length. The head was 
wider than the first segment. The face and front were dull yellow, the 
mouth parts slightly darker, and the antennae black. The ocelli were 
prominent, and jet black. 

Body: Ground color, translucent light yellow. There is a very nar- 
row middorsal longitudinal yellow stripe, bordered by a very wide 
olive-green band. Below this, and on the ventral surface, the color is 
hght yellow. 

The legs are concolorous with the venter as are also the single pair 
of prolegs. No setae are distinguishable with a X 16 hand lens. 


LARVA OF 13 MM. LENGTH 


Head, 0.9 mm. wide, which is also the average width of the body seg- 
ments. The upper half of the head is light orange-yellow and the lower 
half white. 

The body is regularly cylindrical throughout. The ground color is 
light yellow-green. The segmental junctures are yellow. 

There is a middorsal longitudinal band of light green margimed with 
yellow. Lateral to this is a narrow dull green stripe, on which there are 
minute black dots placed approximately one to a segment. Lateral and 
below this are several narrow longitudinal stripes, alternately green 
and yellow. 

The infrastigmatal fold is yellow. The venter is striped as is the lat- 
eral surface. The spiracles are indistinguishable. The legs and prolegs 
are concolorous with the body. 

On September 10, 1956 most of the larvae were mature, anda typical 
example was described as follows: 


Mature Larva 


Length, 30. to 34. mm. Widest segment of body (7th) 2.5 mm. The 
shape is unusually elongate, and cylindrical. 

Head width, 2. mm., very flat. Ground color, light tan. The adfrontal 
sutures are barely discernible as delicate brown lines. 

The front is speckled with light brown, which is absent down the 
center, where the tan ground gives the appearance of a light stripe. 
Near the clypeal juncture there are two dark brown dots, one each side 
of the light middle area. 

The cheeks are mottled with light brown dots and crossed longi- 


Life history notes on Epiplatymetra grotearia is 


tudinally by two lines of blackish-brown dots, these lines ending on 
each cheek at the outer margin of the clypeus. 

The mandibular area, including the clypeus, is of the same shade as 
the ground color, except for a darker shading medially. 

The mandibles are tipped with black, the antennae are hyaline light 
tan, and the ocelli are black. 

In the middorsal area of the body there is a longitudinal crenulated 
and partly discontinuous pair of stripes, occasionally accented with 
black dots. Lateral to this is a pair of crenulated black stripes which ap- 
pear and disappear in an irregular manner. This line is an extension 
of the similar dark line on the cheeks, and it tends to become obsolescent 
near the cauda. 

Below this the body is mottled with numerous light brown dashes, 
dots, and broken longitudinal stripes. There is a semblance of a wavy 
white longitudinal band substigmatally. This becomes well defined 
near the cauda, and is there edged superiorly with a dark brown line. 
On the first three segments this light substigmatal band is bordered in- 
feriorly with a broad band made up of black dots. 

The spiracles are conspicuous and are margined with black. 

The venter is longitudinally marked in stripes of various shades of 
brown. The legs are concolorous with the body. The single pair of pro- 
legs bear a whitish longitudinal band edged with dark dots, and the anal 
prolegs are marked somewhat similarly. 

The crochets are dark brown. The few scattered setae are very short | 
and inconspicuous. The larva is illustrated on Figure 2. 


7 eA 


Length, 13 mm. Greatest width, 3.6 mm. The ground color is light 
tan on the cephalothorax and wing cases, and yellow-tan on the ab- 
dominal segments. The cremaster is deep blackish brown. The three 
middle segmental junctures of the abdomen are wide bands of deep 
brown, the remaining junctures being narrow. 

The outer edges of the antennae are widely bordered with deep 
brown and the fronds are marked by well defined narrow brown lines. 

The eyes are tan, with a marginal shading of light brown. The wing 
cases are striated over the venules. 

On the third and fourth abdominal segments there are protruding 
papillae over the spiracles resting on large dark brown blotches. The 
remaining spiracles are small and deep brown. 

A few broken blotches occur middorsally on the first five abdominal 
segments. 


6 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


The cremaster terminates in a long cone, topped by two straight 
black shafts that are only slightly recurved at the tips. On each side of 
these there are three short yellow hooklets, markedly recurved, 


hie Sa 


pera 


Figure 2. Larva and pupae of Epiplatymetra grotearia. Central, dorsal aspect of 
mature larva. Left, dorsal aspect of pupa. Right, ventral aspect of pupa. All figures 
enlarged X 414. Reproduced from drawings by the author. 

All of the structural features of the pupa are adequately shown on 


Figure 2. The first imago emerged September 27, 1956. 


THE SIGNIFICANCE OF THE PETROLIFEROUS NODULES 
OF OUR DESERT MOUNTAINS! 
W. DwicutT PIERCE? 
Los Angeles County Museum 


In the last six or seven years there have come to notice multitudes of 
odd-shaped nodules containing beautiful silicified fossils and petroleum. 
Prior to 1954, they had been written off as pseudofossils. 

The first collection of the nodules was made by W. E Foshag of the 
U.S. National Museum, at the abandoned Town of Borate in the Calico 
Mountains of San Bernardino County, in the early 1930’s. It was not 
then known that they contained fossils. 

In January 1954, Dr. Allen M. Bassett of the U.S. Geological Survey, 
while studying boron deposits, broke open a nodule which contained a 
fossil dragonfly. He examined other nodules and called them to the at- 
tention of Dr. Allison Palmer, of the Survey in Washington. Palmer 
and Bassett collected from a number of sites opposite Camp Rock on 
Mule Canyon Drive; and by acid treatments Palmer found many in- 
teresting fossil msects. 

I had the pleasure of seeing the Palmer collection in Washington in 
1955, and shortly after, had a visit from Mr. and Mrs. John H. Rouse, 
amateur collectors, who had independently found insects in several 
nodules by cracking them open. They had seen a fossil fish skeleton in 
a nodule found by H. G. Kirkpatrick, and were in search of another fish 
when a dragonfly was disclosed. 

The first Los Angeles County Museum party visited the Calico 
Mountains in May 1956, and we have had four expeditions with a total 
personnel of 51 persons, collecting 29,000 nodules (each one numbered 
and recorded by site, altitude, and collector). Other people have col- 
lected great numbers of nodules, and some very beautiful specimens 
have been extracted by them. 

A total of 100 sites are now known to the writer. While the Calico 
Mountains in San Bernardino County was the original discovery area, 
we have now added the following as nodule sources: Tick Canyon, near 
Lang, in the San Gabriel Range of Los Angeles County; several sites 


1Read at the Annual Meeting of the Academy on May 19, 1961. 


2Curator Emeritus. 


™“N 


8 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


on Mt. Pinos, in the Frazier Mountains, Ventura County; Skull Can- 
yon, a branch of Horse Canyon in the Tehachapi Mountains, in Kern 
County; Lane Mountain, northeast of Barstow and northwest of the 
Calico Mountains. and also Lenwood Hill, south of Barstow; and the 
Horse Shoe Hills. east of the Calico Mountains. all in San Bernardino 
County; and finally Ryan, in Death Valley, Inyo County. 

The significance of this distribution is that all of the sites are sources 
of borax, and are located on or close to andesitic volcanoes. 

It makes an interesting pattern when we consider the triangle of the 
Mojave Desert, bounded on the north by the Garlock Fault, margining 
the Tehachapi Mountains, and with Death Valley at the northeastern 
tip; and on the southwest by the San Andreas Fault, margining the 
transverse mountain ranges of the San Gabriel and San Bernardino 
Mountains; with Tejon Pass at the junction, and Mt. Pinos the fulcrum 
of the great Mojave Squeeze. And then on the axis of the Squeeze are 
Lane Mountain. Lenwood, the Calico Mountains. and Horse Shoe Hills. 

However, I expect more sites to be uncovered in the Panamints, and 
elsewhere in our desert volcanic areas. 

One of the first significant features of these findings is that possibly 
by means of the insect and other fossils, we can correlate the various 
boron deposits of California. 

All of the material so far found is of Miocene age; that is, from 20 to 
25 million years old. 

There are two series of fossil and petroleum bearing nodules in the 
Calico Mountains and the Frazier Mountains. The upper courses of 
nodules are bluish gray in color, while the older deposits contain only 
pale gray to whitish nodules. The nodules brought out from the borax 
mines are bluish. Inasmuch as all of the borax mines have been sealed 
off by the State, we have not been able to get much matrix material for 
the blue nodules, The white nodules are in lake bed deposits exposed by 
erosion in the Canyons. 

From both series of nodules we obtain several kinds of fossils. When 
a new lot of nodules is brought in, each nodule is numbered and ex- 
amined under lens for external specimens. Two percent of all the nod- 
ules examined have actual petrified insects, impressions, or moulds of 
insects, including many wings, visible on the outside. 

Certain types of nodule invite cracking. and display cavities formerly 
occupied by dragonflies, other large insects, and even a beautiful beetle. 
Some of these odinles were actually formed by concretion around the 
insect. Many others, on cracking, display specimens and moulds of in- 
sects which have not been preserved, but the space they occupied is so 


Significance of petroliferous nodules 9 


clearly outlined that the origmal occupant can be readily determined. 
Most of this type are in the upper or blue layers, and silification did 
not always take place. The strontium or celestite fossils are all in these 
upper deposits. 

The finest fossil material in rr blue and white nodules is obtained 
by digestion of the nodules by acids which do not injure insect tissues, 
or the silicified replicas of insects. The acids so far effective are 20 per- 
cent solutions of formic, acetic, and hydrochloric acid, and a weaker so- 
lution of nitric acid. In the case of externally exposed fossils we use the 
slower hydrogen peroxide. The processes take from a day to weeks. 
When we find an impression we pour in latex and obtain a cast of the 
original occupant, so perfect that we can identify many of them. 

To date, the findings in the nodules obtained by Dr. Palmer at Wash- 
ington, Mrs. Ruth Kirkby at Riverside, Mr. John Gibron, Sr., at Camp- 
bell, and myself include specimens in 26 orders of invertebrates as fol- 
lows, from the Calico Mountains, unless otherwise noted: 


Turbellaria, or flat worms of three families, and also many strings of 
eggs, some showing embryos. 
Mollusca. 

Anostraca, or fairy shrimps, in various stages of growth, and multi- 
tudes of their coprolites, and even the eggs in the ovarium. 
Ostracoda, from Calico Mountains, and also from Lenwood Hill. 

Copepoda, from Lane Mountain. 

Acarina, water mites, and their eggs, several species. 
Arachnida, water spiders, several species. 

Scorpionida, claw of a Scorpion. 

Chelonethida, pseudoscorpion, several claws. 
Myriapoda, a centipede. 

Collembola, springtails. 

Odonata, dragon flies and damsel flies. 

Ephemerida, may flies, and clusters of eggs. 

Plecoptera, Stone fly nymph. 

Corrodentia, bark louse young. 

Isoptera, five species of termite wings, and soldier head. 
Thysanoptera, both suborders of thrips, several species. 
Chermodea, jumping plant lice, adults, and wings. 
Aphidodea, aphids. 

Homoptera, several species of leafhoppers. 

Hemiptera, several families of bugs, including egg of one. 
Orthoptera, fragment of one grasshopper. 


10 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


Coleoptera, several families of beetles, cluding larvae and adults. Also 
found in the Frazier Mountains. 

Lepidoptera. crystallized aquatic moths, and impression of large moth. 

Hymenoptera, a crystallized Braconid parasite, an ant, and impression 
of a wasp. 

Diptera. several families of midges and flies, including eggs, larvae, 
pupae, adults. Also found in the Frazier Mountains. 

Pisces, two fish skeletons. 

Aves, crystallized feathers, and also feather impressions. 

Mammalia, crystallized hairs of mammals. 


In the plant kingdom: 


Diatoms of several types. 

Algae of many kinds. 

Mosses of several types. stems and leaflets. 
Pond weeds, entire plants. 

Seeds of higher plants. also impression of a leaf. 


One beetle had died and been filled with fungus, and then crystal- 
lized, and the masses of fungus protrude from the segments of the crys- 
tal skeleton. 

Thus we have a good cross section of Miocene life in 3-dimensional 
preservation. The best of it is that in numerous specimens the internal 
structures, muscles, and nervous system have been preserved. 

While crystal replication of insects and plants is absolutely new to 
Paleontology since 1954. We may also add as new records the finding of 
fossil TR pSIESGA or flat worms, never before reported. Another first is 
the twenty or more kinds of fossil eggs. Dr. Hermann Weyland in 
Bavaria has, since our first announcement, reported insect eggs from 
hgnite. 

The beautiful fossils have another story to tell; that of sudden death 
and quick preservation. In proof of this are a may fly, which has an 
active life of one day. in front of her mass of eggs: several Heleid 
midges caught half way out of their pupa cases; a pair of bugs, and a 
pair of flat worms in copula; strings of eggs showing embryos in vari- 
ous stages of development; and fairy shrimps w ne eggs in the brood 
pouch. In fact, the multitude of insect larvae and pupae in perfect 3- 
dimensional preservation tell of rapid kill and preservation. There are 
other cases of decomposed bodies preserved in that condition. The tiny 
pond weed with corm, rootlets, and leaves in perfect form is a good ex- 


Significance of petroliferous nodules 14 


ample. Whatever the state of the biological material at the moment of 
catastrophe, that is the state that has been preserved for us. 

The nodules are of several types. There are true concretions formed 
around dragonflies and other large insects; however, the insects in these 
are not silicified, sometimes strontium preserved. There are concretions 
formed by sudden solidification of boiling minerals, in which the for- 
mation is radial from a nucleus. These do not contain fossils. The com- 
monest type are flat layered nodules, which were originally part of a 
layered bottom formation, and they almost uniformly display biological 
material. 

I interpret from the nodules that life in the lake was normal often 
for many years, and then sudden catastrophe killed everything in the 
lake and preserved it. The fossils are not confined to nodules, for we 
find good specimens in matrix lake bed strata. In such matrix there is 
also petroleum. Such catastrophes occurred many times in the thou- 
sands of years represented by this lake bed series. 

In addition to the deposit in the lake bed of volcanic gases, liquid and 
ash, there was violent earthquake tumbling of the rocks, and we see 
many evidences of upturned lake beds, of folding and twisting, and in 
the anticlmes and synclines we find nodules in position in strata which 
had been flat when they rolled in. 

My interpretation of the majority of the fossil bearmg nodules is 
that the lake beds were uplifted and tilted by earthquakes, and then 
sunbaked, cracked, and broken, and the nodules thus formed fell back 
mto the waters, rolled and tumbled and got their rounded form; and 
new deposits grew up around them, setting them in the matrix in 
which we now find them. Often a nodule was broken at the point of fold 
showing that the folding took place when the layers were soft. There 
are many evidences that the nodules are older than the matrix m which 
they now are, and it can be seen that the surrounding matrix was grad- 
ually built around them. 

We also find matrix layers unbroken, containing the same kinds of 
insect remains. The matrix was laid down in two annual layers, which 
are of different colors; and in most areas these annual double bands 
measure 25 to the inch. At this rate the vertical wall of 88 feet in the 
southwest quarter of section 24, containing three great bands of nodule 
bearing strata could have taken up to 26,400 years to deposit. Thus we 
know that the lake existed either constantly or periodically for many 
thousands of years, and that there were many periods of volcanic dis- 
turbance. 

The oldest bed, judging from its lowest position, contains more spe- 


12 Bull. Sou. Calif. Acad. Sct. / Vol. 61, Pt. 1, 1962 


cies of insects, and a greater diversity of species than the upper beds; 
and tentatively we might suggest that some species which originally 
existed in the lake or lakes, were extinguished, while only certain forms 
persisted through the centuries. 

There are two peculiarities to the fossil deposits: one is that the bio- 
logical material is crystallized either as silicate, or as strontium celestite. 
The other is that petroleum is present in practically every fossil bearing 
nodule. 

The strontium preservation is commonest in the upper strata of blue 
nodules, or in the white concretions containing dragonflies. The silica 
preservation is in two states, crystal and colloidal. The colloidal silica 
fossils have perfectly clear mes, and are complete replications of the 
skeletons of the plants or msects, even to the hairs on the antennae and 
the septa in the compound eyes. 

A replication like this requires a catalytic process of exchange of 
silica for chitin. 

The petroleum phenomenon is unusual and I believe very important 
to us. The odor of petroleum in the mines and caves is so strong that one 
can hardly stand it. If you rub two nodules together you smell petro- 
leum. If you dissolve the nodule in acid, the petroleum foams up in 
great quantities. 

When you break a nodule you may find a great quantity of hardened 
petroleum in the heart of the nodule, and sometimes it is still Liquid. 
Specimens containing insects may have a petroleum film surrounding 
the insect. The crystallized beetle with the fungus was full of petro- 
leum, which came out when the specimen was put in xylol. Sometimes 
silicification has not taken place and the insect tissues are red; however, 
when dropped in xylol the entire specimen is dissolved. We solved this 
and saved the insect tissues by avoiding xylol and clearing in terpineol 
before mounting in balsam. 

In the past it has been said that petroleum was formed from plant 
tissues and from diatoms. In this lake there were pond weeds, algae. 
diatoms, great layers of coprolites of fairy shrimp, and the animal life 
of the lake. I hold that all forms, plant and animal, which were con- 
verted into silica, contributed the substance which became petroleum. 

In other words, there was another catalytic process: the formation of 
petroleum from the biological material freed by silicification, and even 
that which was not silicified. I have now concluded that the two cata- 
lytic actions were independent of each other, and there are cases where 
each process was incomplete. There are insects with half of the body 
silicified and half not. 


Significance of petroliferous nodules 13 


It is well known that catalysis is brought about by the presence of a 
substance which does not itself enter into the final product. There was 
present at every site so far investigated, a well known catalyzer, boron. 
Boron is a primary volcanic product, and the primary deposits are those 
we must seek for our fossils. Borax is water soluble and has washed 
down from the volcanic primary sites to settle in brine basins as sec- 
ondary boron, forming the dry lakes of the desert. 

Boron is well known for its affinity with silicon, and boron silicate 
glass is one of the finest glasses known for laboratory work. Boron is 
also recently in the limelight as a catalyzer for the improvement of the 
quality of gasoline. Boron is volcanic, and has been mined at every site 
where the petroliferous fossil bearing nodules are found. Colloidal sili- 
con replications of living creatures have never been found elsewhere 
than in the neighborhood of borax deposits on andesitic volcanoes. Be- 
cause of these facts I reason that boron was the catalyzer responsible, 
when in highly heated condition, for the killing, preserving, and repli- 
cating of the insects and plants; and for the formation of the petroleum 
from their remains. 

Since there is every evidence that the two catalytic processes were 
rapid, I reason that what happened once can happen again. Therefore, 
I challenge the chemical profession to produce in their retorts, petro- 
leum from the waste biological material of our cities on a 24-hour basis. 
If I am right, the fuel supply of the future is assured us forever. 

The problem may not be simple until we know all the ingredients 
and proportions. There is another factor that must not be overlooked. 
Many important chemical processes require a living, active agent at 
some step in the process. Some of the known agents in manufacture are 
bacteria, yeasts, fungi, and algae. 

There were in these lakes algae and diatoms. The nodules yield beau- 
tiful silicified specimens of many types of algae, and often multitudes 
of diatoms. 

Algae in lakes are known to take carbon dioxide from calcium bi- 
carbonate and deposit the calctum carbonate as sediments. Likewise, 
algae have an affinity for silica in water, transforming it into colloidal 
silica. As long as I believed the two catalytic processes dependent one 
upon the other, I felt that perhaps the algae would be necessary to the 
commercial production of petroleum, and then the process might be 
slower. The latest findings indicate that petrolification was not depend- 
ent upon silification, and in that case colloidal silicon in the presence of 
boiling boron, might release the tissues for immediate conversion into 
petroleum. Of course our catalytic process will require the presence of 


14 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


certain gases; and we are certain that in volcanic gases there are hydro- 
gen, carbon dioxide, and sulphur gases. 

Strontium entered the picture only im a small part of the nodules; 
however, it must not be discounted. 

Even if we prove that the petroleum in the volcanic mountain nod- 
ules was produced by boron catalysis, this does not necessarily mean 
that petroleum has not also been formed by other processes. We may 
have, however, pointed the way for solving the other types of forma- 
tion. 

It has long been thought that petroleum was formed in salt water; 
however, now we have evidence of its formation in fresh water into 
which volcanic salts came suddenly, and fresh water life continued in 
subsequent years. 

Most reports on boron indicate that it is primarily volcanic; second- 
arily is leached into sinks and flat beds; and finally seeps into the ocean. 
May we not suspect that perhaps boron can still have catalytic powers 
in its secondary and tertiary phases? 


A NEW PASIPHAEA (CRUSTACEA, DECAPODA, 
NATANTIA) FROM SOUTHERN 
CALIFORNIAN WATERS? 


JouHwn C. YaLpwyn?” 
Allan Hancock Foundation, University of Southern California 


Since 1949, members of the Allan Hancock Foundation and of the De- 
partment of Biology, University of Southern California, have under- 
taken an increasingly intensive study of the fauna and ecology of the 
mid-waters of the deep basins, especially the San Pedro Basin, off the 
coast of Southern California. During the course of this work several new 
or unusual species of macrurous decapod Crustacea (shrimps and 
shrimp-like forms) have been obtained. The bulk of the collections 
made before September 1960 had already been examined and identified 
by Dr. John S. Garth of the Hancock Foundation, and specimens of the 
Pasiphaea described here had been sent to Dr. Fenner A. Chace, Jr., of 
the U.S. National Museum and had been recognized by him as a new 
species, though not described. Thus it has been my privilege, while 
working on these deep-water Crustacea during 1960-61, to describe this 
Pasiphaea in the first of a series of studies on this interesting material. 

The San Pedro Basin, lying between San Pedro and Santa Catalina 
Island, has an area of about 655 square kilometres and a maximum 
depth of about 912 metres, while the sill depth (the depth of the lowest 
point on its rim) is about 737 ‘metres. The new Pasiphaea has been 
taken as shallow as about 300 metres over this basin, as well as in nets 
that have touched the basin floor (actual depth of capture unknown), 
but it does occur in abundance, at least during daylight, at depths of 
about 550 to 700 metres. It has almost always been associated with 
Sergestes similis Hansen and appears to occur in a shallower zone than 
that in which the larger, more intensely pigmented, Pasiphaea emar- 
ginata Rathbun is usually taken. 

Though occurring in numerous hauls in the San Pedro Basin, as well 
as in the deeper Catalina and San Nicolas Basins further offshore, the 
new Pasiphaea is recorded here from only four selected stations as its 


1Allan Hancock Foundation Contribution No. 245. 


On leave from the Dominion Museum, Wellington, New Zealand; visiting Ful- 
bright Advanced Scholar supported by a grant from the National Science Founda- 
tion. 


15 


Fig. 1, lateral view carapace (arrow indicates extent of dorsal carina); fig. 2, 
lateral view abdomen; fig. 3, lateral view anterior part of carapace; fig. 4, dorsal 
view tip of telson; fig. 5, antennule; fig. 6, scaphocerite; fig. 7, mandible; fig. 8, 
ist maxilla; fig. 9, 2nd maxilla; fig. 10, 1st maxilliped; fig. 11, 2nd maxilliped; 
fig. 12, 3rd maxilliped; fig. 13, 1st pereiopod; fig. 14, 2nd pereiopod; fig. 15, 3rd 


Va 


i 


\ 


i 


pereiopod; fig. 16, 4th pereiopod; fig. 17, 5th pereiopod; fig. 18, male ist pleopod; 
fig. 19, male 2nd pleopod. Figs. 1 to 17 from holotype, 18 and 19 from paratype, 
carapace length 17mm. Figs. 5 to 19 of right appendage. Figs. 1 and 2 to same 


scale; figs. 4, 7 to 11, 18 and 19 to same scale; figs. 5 and 6 to same scale; figs. 12 
to 17 to same scale. 


Z. 


@) 


18 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


occurrence and bathymetric distribution will be discussed in detail in 
a future paper on the bathypelagic natant decapod Crustacea taken by 
the Velero IV off Southern California. 


TABLE 1 
Selected Allan Hancock Foundation stations in the San Pedro Basin from which 
Pasiphaea chacei n.sp. was obtained. All the specimens from these four stations are 
to be regarded as type material. 


Net Down Net Up 
Sta. No. Date Position Time Fms_ Position Time Fms Remarks 
7186-60 Oct.28 33-36-11 1035 475 33-30-52 2 metre net 
1960 118-32-13 118-22-48 hit bottom 
during 114 
hours at 
depth. 
7221-60 Dec.9 33-37-36 1008 463 33-27-13 1510 461 Isaacs-Kidd 
1960 118-32-28 118-17-24 Mid-Water 
Trawl at 1931 
ft. by gauge; 
4 hours at 
depth. 
7273-61 Jan.23 33-39-07 1015 458 33-27-47 1503 IKMWT at 
1961 118-31-02 118-16-06 1986 ft. by 
gauge; 34% 
hours at 
depth. 
7299-61 Feb.24 33-38-03 1005 468 33-25-48 1510 458 IKMWT at 
1961 118-32-39 118-16-48 2341 ft. by 
gauge; 4 
hours at 
depth. 


SECTION CARIDEA 
Family PAsIpHAEIDAE 


Genus Pasiphaea Savigny, 1816 
Pasiphaea chacei new species 


Ty pes: Holotype, female, carapace length 19mm. from Station 7186-60 
(Allan Hancock Foundation Catalogue No. 601). Paratypes, 1 female 
11mm.° from Station 7186-60; 2 males 12 to 17mm., 30 females and 
immature 6 to 15mm., 2 ovigerous females 19mm., from Station 7221- 
60; 10 males 10.5 to 15mm., 24. females and immature 7.5 to 20mm. 


“The carapace length, in mm., is used as the standard length throughout. 


New Pasiphaea from southern California 19 


(including 1 ovigerous 20mm.) from Station 7273-61, and 190 speci- 
mens, of which a sample of 50 consisted of 19 males 11.5 to 19.5mm. 
and 31 females 9.5 to 22mm. (3 ovigerous 19 to 21mm.), from Station 
7299-61. 

Diagnosis: Telson truncate distally. Carapace dorsally carmate, with 
a characteristic, slender, acute rostrum projecting anterodorsally at 
least as far as cornea. Abdomen smooth, not carinate. 1st pereiopod 
with 0 to 12 meral spines, 2nd with 6 to 23 meral spines. Fingers of 2nd 
pereiopod subequal with palm. 

Description: Rostrum (gastric spine of some authors) prominent, 
relatively long and slender, acute, directed obliquely dorsally, without 
a curve, to extend anteriorly at least as far as cornea. Carapace with a 
distinct carina on anterior 2% of dorsal surface, posterior third smooth 
and rounded. Lateral surface of carapace with suprabranchial carina. 
Anterior margin produced dorsally mto a blunt, convex lobe, not ex- 
tending as far anteriorly as the broadly rounded lower orbital angle; 
this angle continuing ventrally mto a weakly concave emargination, 
which in turn passes through a broadly rounded angle to trend oblique- 
ly posteroventrally into a concave emargination at level of branchio- 
stegal spine, then into the deep branchiostegal sinus. Branchiostegal 
spe prominent, arisimg behind, and projecting beyond, anterior 
margin. 

Abdomen smooth, all segments dorsally rounded and unarmed; 6th 
strongly compressed dorsally but not carinate. Pleura of 1st and 2nd 
broadly rounded; pleura of 3rd to 5th produced somewhat anteroven- 
trally, 5th with concave ventral margin. 6th half as long again as 5th, 
with weak dorsolateral carina. Telson a little shorter than 6th segment, 
with a broad, longitudinal groove dorsally and a truncate distal mar- 
gin armed with four pairs of spinules graded in size from a short me- 
dian pair to a relatively long lateral pair. 

Eyes well developed, cornea rounded, well-pigmented and broader 
than ocular peduncle. 

Antennular peduncle with first segment reaching with less than 1 
its length beyond eyes. Stylocerite narrow in dorsal view and relatively 
widely separated from first antennular segment proper; in lateral view 
dorsal margin forming an acute point distally which passes ventrally 
into the rounded anterior margin of the broad ventral portion of the 
stylocerite. Two antennular flagella present, dorsolateral flagella with 
about 13 enlarged basal segments in female, about 17 in male; enlarged 
basal section in male considerably broader and more abruptly passing 
into the flagellum proper than in the female; this character is especially 


20 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


useful for sexual differentiation. Antennal peduncle reaching to about 
midpoint of antennular peduncle, basal segment with strong, obliquely 
directed spine. Scaphocerite reaching distally beyond antennular pe- 
duncle to about last enlarged flagellar segment; outer margin weakly 
convex, produced into a strong distal tooth projecting full length be- 
yond lamella, which is 31 times as long as wide. 

Mandible consisting of toothed incisor process only, palp absent. 1st 
maxilla with small, rounded proximal endite, strongly toothed distal 
endite and long, simple endopod armed with a single stout seta. 2nd 
maxilla with no endites visible, simple, well developed endopod and 
large scaphognathite. 1st maxilliped reduced to large elongate lamella, 
articulated distally. 2nd maxilliped simple, with distal segment articu- 
lated normally with penultimate, no epipod or exopod. 3rd maxilliped 
reaching a little beyond scaphocerite, ultimate segment long and slen- 
der, somewhat less than twice length of penultimate and shorter than 
antepenultimate, exopod well developed. 

All pereiopods with well developed exopods, but no epipods. 1st pe- 
reiopod reaching with fingers and half palm beyond scaphocerite. Fin- 
gers slender, cutting edges toothed, tips curved and capable of crossing 
one another, ¥4 length of elongate, parallel-sided palm. Palm with 
medioventral row of about 5 to 7 slender, movable spines, as shown 
alongside fig. 13. Carpus 13 palm, armed with distoventral spine. 
Merus 5/4. palm, armed ventrally with from o to 12 (usually 2 to 9) 
unevenly sized and irregularly placed spines. Ischium about 14 length 
merus, unarmed. Basis with strong, distoventral spe. 2nd pereiopod 
reaching with fingers and half palm beyond scaphocerite. Fingers 
elongate, slender, cutting edge toothed, tips curved and capable of cross- 
ing one another, subequal in length to elongate palm, which is some- 
what narrower distally than proximally. Carpus a little less than 14 
palm, armed with strong distoventral spine. Merus 5/3 length palm, 
armed with from 6 to 23 (usually 14 to 21) unevenly sized spines 
spaced relatively evenly along entire ventral margin. Ischium about 
1/, length of merus, unarmed. Basis with distoventral spine. 3rd pereio- 
pod slender, attenuated, reaching to distal end of antennal peduncle, 
dactyl subequal to carpus and about 1/10 propodus which is 24 merus. 
Ischium a little less than 14 merus, all segments unarmed. 4th pereio- 
pod short, reaching to basoischial articulation of 2nd pereiopod, dactyl 
1, propodus, carpus about 14 propodus, merus equal to carpus and pro- 
podus combined, ischium to 1 carpus, propodus and dactyl combined. 
5th pereiopod distinctly longer than 4th, reaching a little beyond is- 
chiomeral articulation of 1st pereiopod. Dactyl Halle eley broad, and 


New Pasiphaea from southern California BL 


rounded distally, a little more than 14 propodus and a little more than 
1 carpus, carpus subequal to ischium and 2/5 merus. 

First pleopod male and female with broadly ovate endopod and short 
appendix interna bearing some hooks distally. 2nd to 5th pleopods male 
and female with slender appendix interna, 2nd pleopod male with in 
addition a slender and shorter appendix masculina. Uropods elongate, 
exopod longer than endopod, exopod with lateral margin produced into 
distal spine reaching beyond distal margin of lamella. 

Eggs large and relatively few in number, measuring after preserva- 
tion, 1.79 to 1.88 < 1.26 to 1.38 mm. 


TABLE 2 
Branchial Formula. 
Maxillipeds Pereiopods 
1st Qnd 3rd 1st 2nd 3rd 4th 5th 
Pleurobranchiae : i : 1 1 1 1 { 
Arthrobranchiae : ; ‘ 1 1 1 
Podobranchiae 
Epipodites 
Exopodites : ‘ 1 1 1 1 1 1 


Color in Life: The body and appendages are mainly transparent, 
while the eyes are dark golden: brown. There is an irregular scattering 
of large, stellate, red chromatophores on the body and appendages, dif- 
fering greatly in concentration with the size of the specimen. Small 
specimens are almost colorless, while in larger specimens the carapace 
at least appears pink, the fingers are very red with the curved tips grey- 
black in color. Eggs in an early stage of development are colorless. The 
viscera appears through the carapace as a dark mass anteriorly and 
yellow gonads posterodorsally (in mature specimens). 

In comparison with Pasiphaea emarginata, taken in the same area, P. 
chacei is considerably less pigmented, especially on the carapace, the 
pereiopods and the anterior segments of the abdomen. 

Maturity and Sexual Differentiation: The smallest ovigerous female 
examined in the material available had a carapace length of 19mm., 
however, some females at a carapace length of 15mm. had a small, but 
quite distinct, ovary, containing ova, clearly visible posterodorsally 
through the transparent cuticle of the carapace. Males with a carapace 


22 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


length of 15mm. or less had an extremely small appendix masculina, 
which at a carapace length of 12mm. and below was usually not dis- 
cernible. The difference in width of the enlarged basal segments of the 
antennular flagella, as described above, is, however, a clear sexual dis- 
tinction at all sizes down to at least a carapace length of 1omm., and in 
larger specimens can be easily used for “in hand” sexing without 
magnification. 

The size at maturity then is a carapace length of about 15 to 16mm., 
with all males above this size bearing a well developed appendix mas- 
culina and all females above this size with a clearly visible, developing 
ovary. 

Variation in Meral Spinulation: The number of spmes on the meri 
of the 1st and 2nd pereiopods has been widely used in this genus as a 
systematic character. Considerable variation within a species has been 
recorded previously (e.g. in Pastphaea sivado by Stephensen, 1923). 
but the extent of this variation both within a species and between the 
right and left members of a pair of limbs im the same individual is often 
not fully appreciated. Fifty specimens selected at random from Station 
7299-61 gave the following figures. (Data for 2nd pereiopods in paren- 
thesis. ) 

The spines on the merus of the 1st pereiopod (2nd pereiopod) varied 
from o to 12 (6 to 23), with 90% (80%) of the sample having from 
2 to g (14 to 21) on the right member. Sexually mature specimens 
tended to have more than immature specimens. Thus the variation in 
the 23 mature specimens was from 2 to 12 (15 to 23) with an average 
of a little more than 6 (18), while in the 27 immatures the variation 
was from o to g (6 to 19) with an average of a little more than 3 (a 
little less than 14.). The difference in the number of spines between the 
right and left member of the 1st pereiopods (2nd pereiopods) varied 
from o to 3 with an exceptional 5 (0 to 4); in 38% (28%) there was 
no difference between the right and left member, in 40% (38% ) there 
was a difference of one, and in 18% (18%) a difference of 2, one speci- 
men had a difference of 3 and one of 5 (five specimens had a difference 
of 3 and three a difference of 4.). 

In a general way the greater the number of spines on the 1st pereio- 
pod the greater the number on the 2nd. The ratio of the number of 
meral spines on the right 1st to the number on the right 2nd in the 
sample varied from 1:1.6 to 1:9 (96%) with one at 1:12 and one at 
1:16. The difference between the number of spines on the right 1st and 
the number on the right 2nd varied from 4, to 17 with 76% between 10 
and 14, and the average at a little over 11. 


New Pasiphaea from southern California 23 


Finally it should be stated that there is a great variation in the size of 
the meral spines present, and that all recognizable spines have been 
counted no matter how small. Sexual significance in variation was 
tested for, but no correlation could be made. 

Systematic Position: The carinate carapace, the smooth abdomen and 
the distally truncate telson clearly distinguish this species from all but 
five of the approximately 34 described and recognized species and 
forms of the genus Pasiphaea. These five species are as follows: P sivado 
(Risso, 1816) from the North Atlantic, Mediterranean and Indo-West 
Pacific; P unispinosa Wood-Mason, 1893, from the Indian Ocean; P 
magna of Rathbun, 1904. (only questionably identified by Rathbun as 
P. magna Faxon, 1893) from Californian waters; P flagellata Rathbun, 
1906, from Hawaiian waters, and Pasiphaea sp. oc. de Man, 1920, from 
Indonesian waters. 

Pasiphaea chacei differs from P sivado (first adequately described by 
Zariquiey, 1957) in that the latter has the carapace compressed but not 
actually carinate and a small but distinct spine posterodorsally on the 
6th abdominal segment (personal observation from Mediterranean ma- 
terial) ; it differs from P unispina in which the condition of the abdo- 
men is undescribed, in the number of spines on the meri of the 1st and 
and pereiopods, only o and 1 respectively in the latter species; it differs 
from P magna of Rathbun (which I am convinced is conspecific with 
P. magna Faxon) in that Rathbun’s specimens have the 2nd to 5th ad- 
dominal segments carinate (personal observation from Rathbun’s orig- 
ial material ) ; it differs from P flagellata, in which the condition of the 
dorsal surface of the carapace is undescribed in the literature, in that 
the carapace is non-carinate in the latter (personal observation from 
Rathbun’s original material), and finally P chacei differs from Pasi- 
phaea sp. oc of de Man in that the carapace in the latter is compressed, 
but not actually carimate dorsally, and the number of spines on the 
meri of the 1st and 2nd pereiopods is, as in P unispina, o and 1 respec- 
tively. In addition to the above differences the characteristic, slender, 
prominent rostrum, extending beyond the anterodorsal margin of the 
carapace, Clearly distinguishes P. chacei at a glance from these and other 
similar species. 

Comparison with Pasiphaea emarginata Rathbun: The only other 
species of Pasiphaea taken commonly in the San Pedro Basin is P emar- 
ginata Rathbun (P. pacifica Rathbun, with a deeply forked telson, oc- 
curs very rarely). Though P emarginata has a carinate abdomen and 
a weakly, but distinctly, distally-emarginate telson in adult stages, in 
juveniles these features are not at all apparent and may, especially in 


24 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


damaged material, be impossible to make out. It has been found, how- 
ever, that in juvenile P emarginata (at carapace lengths of less than 11 
mm. ), though the telson may be apparently distally truncate, there are 
always at least 5 pairs of spines on the distal margin of the telson and 
often a small medial spine as well, while in P. chacei (juveniles and 
adults) there are almost always only 4 pairs of distal spines. In the rare 
cases Where more than 4 pairs are present, identification must be 
made from rostral shape, strength of branchiostegal spine and depth of 
branchiostegal smus. 

In general P emarginata, in contrast to P. chacei, is usually bigger, 
specimens with a carapace length up to about 45mm. are quite common 
from San Pedro Basin (compared to a maximum of about 22mm. in 
P. chacei), and more brightly colored; in P emarginata the carapace 
appears deeper anteriorly as the posterior margin of the branchiostegal 
sinus is longer; the front (the anterodorsal margin of the carapace) is 
more prominent and reaches as far as the suborbital angle; the rostrum 
does not extend up to the front and is separated from the front (in lateral 
view) by a much less acute angle or more often a smooth curve (in 
juveniles the rostrum may extend almost vertically); the branchio- 
stegal spine is considerably shorter and does not extend much beyond 
the anterolateral margin of the carapace; there is a distinct, submar- 
ginal, low carina around the dorsal portion of the branchiostegal sinus; 
the dorsal carina of the carapace extends nearly to the posterior margin 
and the 2nd to 5th abdominal somites are distinctly cariate. 

Remarks: It gives me great pleasure to name this species for Dr. 
Fenner A. Chace, Jr., whose work on Crustacea, especially Western 
Atlantic Caridea, is so well known, and to memorialize his name, along 
with those of other eminent carcinologists—Alcock, Doflein, Faxon and 
Rathbun—in the wide-ranging and typically bathypelagic genus. 
Pasiphaea. 

I wish to thank Dr. John S. Garth for suggesting and aiding this 
study, and for permission to examine these specimens, and also the Al- 
lan Hancock Foundation for the use of laboratory space and equipment. 


LITERATURE CITED 
STEPHENSEN, Kk. 
1923. Decapoda-Macrura excl. Sergestidae. Rep. Danish Ocean. Exped. Mediter- 
ranean, 2 (1D3):1-85. 27 figs. 
ZARIQUIEY ALVAREZ, R. 


1957. Decadpodos espanioles XIII—Las Pasiphaeas del Mediterraneo occidental. 
Trabajo Museo Zoologia Barcelona, n.s. Zool., 11 (5):1-31, 9 pls. 


NONMARINE MOLLUSCS FROM RECENT SEDIMENTS 
NEAR VERNON, APACHE COUNTY, ARIZONA 


Rosert J. DRAKE 
The University of British Columbia 


INTRODUCTION 


Mr. James Schoenwetter, of the Geochronology Laboratories of the 
University of Arizona, had in 1960 worked out a B. C. 1500 to A. D. 
1300 pollen sequence for the Vernon area as part of studies on post- 
glacial climatic and resulting cultural changes for northern Arizona. 
In the summer of 1961, Mr. R. H. Hevly continued work in the region, 
concentrating on the pre-B. C. 1500 time. Mr. M. E. Cooley of the U. S. 
Geological Survey collaborated in stratigraphic field studies of the Re- 
cent geology. 

In connection with gathering material for C-14 dating, samples for 
pollen analysis were also taken; invertebrate remains also discovered 
were turned over for checks on the preliminary paleo-environmental 
mmplications shown by pollen and lithologic work. In order to put on 
record findings from stratigraphy and malacology for the immediate 
area, the following description is presented. 


Unirs IV & Ill 


Six miles north of the town of Vernon lies old Laguna Salada with an 
associated series of beach deposits with archaeological materials of early 
man and later puebloid affinities. It isin T. 11 N., R. 25 E. A large ar- 
royo drains southeasterly into the northwest end of Laguna Salada; the 
elevation is ca. 6,300 feet. Messrs. Helvy and Cooley took samples from 
the arroyo wall in fluviatile and/or lacustrme deposits. Four strati- 
graphic units were in sequence in the arroyo; their approximate ages 
were determined. In the top two units (IV & IT) invertebrate remains 
were collected with matrix. For the molluscs, mostly freshwater, the 
return was low and only genus-level determinations are presented at 
this time. 


Unit IV—Rust stained sand and gravel; with Cretaceous (?) brachio- 
pod shells, apparently of one kind. 
Sample No. LS-3, 13-F: 
(Grey sand and gravel. ) 


to 
Or 


26 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


GYRAULUS 


PURIEL ID 
: LYMNAEID 


SUCCINEID 


Figure 1. Commonly occurring nonmarine molluscan remains in Recent deposits 
near Vernon, Arizona. Scale lines = 1 mm. 


Gyraulus, rare in comparison to Unit III samples. Fragments of 
brachiopod shells or a Cretaceous (2) oyster-like bivalve. 


Unit III.—Clays, sands, and gravels. 
Sample No. LS-3, 12F-“B” [superior position | : 
(Dark brown clay.) 
1 lymnaeid freshwater snail shell; illustrated in Figure 1. Abun- 
dant Cretaceous (?) foraminifera; large, some up to 1.5 X 7.0 mm 
and illustrated in Figure 2. 1 framentary Gyraulus, a freshwater 


Nonmarine molluscs from Arizona Recent sediments 27 


snail shell. Fragments of the body whorl of a large snail; possibly 
freshwater, and a lymnaeid. 

Sample No. LS-3, 12-F: 
(Dark brown clay. ) 
Numerous succineid terrestrial, [ ?amphibious], snail shells; ex- 
ample illustrated in Figure 1. Foraminifera; same kind as in previ- 
ous sample. Fragments of Cretaceous (?) brachiopod shells. Many 
Gyraulus. Fragments of body whorls of freshwater snails or suc- 
cineids or both. 

Sample No. LS-3, 11-F: 
(Grey sand and gravel. ) 
Many Gyraulus; example illustrated in Figure 1. 1 fragmentary 
terrestrial pupillid snail shell; illustrated in Figure 1. 

[ Yellow brown clay. No invertebrate remains discovered. | 


ye aD Caz 
Figure 2. Examples of Cretaceous (?) foraminifera occurring in Unit III of 
reported Recent sediments near Vernon, Arizona. Scale lines = 1 mm. 
DiscussIon 


In Pleistocene and Recent alluvial deposits in the Southwest, it 1s some- 
times suspected that redeposition has occurred. In many areas where 
the Recent geology has been little studied, it cannot be readily known if 
often present terrestrial and freshwater molluscan fossils are i situ or 
not. Therefore, the presence of Cretaceous (?) brachiopods and fora- 
minifera with Pleistocene or Recent snail shells in the old Laguna 
Salada area is phenomenal. 


28 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


SUMMARY 
The presence, by very evident redeposition of much older organic fos- 
sils, Cretaceous (?) brachiopods and foraminifera, in beds of Units IV 
& III points to the existence in the immediate region of a marker for 
possible future value to correlation. Additional interdisciplinary studies 
are needed to determine the extent in time and space of the particular 
alluviation pattern depositing sediments in Units IV and III. 


ACKNOWLEDGMENTS 


I am grateful to Mr. Hevly and Mr. Cooley, also to Dr. P. S. Martin of 
the Chicago Natural History Museum for the opportunity to study the 
material and situation. Drs. W. H. Mathews, V. J. Okulitch, and A. E. 
Cockbain of my University kindly examined the unusual forams. This 
report constitutes partial results of research supported by Grant NSF- 
184.72 from the Anthropology Program of the National Science Foun- 
dation, 1961-1962. 


RECORDS OF SNAKES FROM JOSHUA TREE NATIONAL 
MONUMENT, CALIFORNIA? 


RicHarp B. Loomis AND RoBERT C. STEPHENS 
Long Beach State College 


The following preliminary report represents the compilation of rec- 
ords of snakes obtained in or near Joshua Tree National Monument 
from August, 1959 through 1960. These specimens were taken in con- 
nection with detailed studies of the flora and fauna being conducted by 
the authors and others at Long Beach State College, with the generous 
aid of the Monument staff. 

The records listed below include those which fill certain distribu- 
tional gaps, represent range extensions and in most cases represent the 
first published account of the species within the Monument. 

Fifteen species of snakes were taken within the Monument. A total 
of 150 snakes are listed below, including 125 records from the Monu- 
ment. Most of the snakes were found on the roads, either dead (DOR) 
or alive (AOR). Some identified examples were not saved either be- 
cause of their poor condition or when alive, because of adequate samples 
available from the area. 

Examination of the seasonal data reveals that most of the snakes were 
taken between late March and the first of July (130 of 150 records). 
This coincides in part with the amount of collecting since the reduction 
of snakes obtained per night discouraged additional trips. However, col- 
lecting trips were conducted in and around the Monument in every 
month of the year, and the Monument staff picked up specimens when 
possible. A summary of the records is as follows: 

Jan.—o, Feb.—o, March—24, April—38, May—46, June—23, July— 
5, August—5, Sept.—3, Oct.—6, Nov.—o, Dec.—o. 


ACKNOWLEDGMENTS 


We wish to thank the following persons who have aided in the assembly 
of specimens and observations: Dr. Dennis G. Rainey, Dr. Elbert L. 
Sleeper, Mr. South Van Hoose, Mr. Julius C. Geest, Mr. Kenneth D. 


1Contribution No. 3 from the Biological Sciences Department, Long Beach State 
College, California. 


30 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


Peyton and others, all from Long Beach State College; Superintendent 
William R. Supernaugh, Mr. James R. Youse, Park Naturalist and 
others on the staff of Joshua Tree National Monument. To Mr. Super- 
naugh and to the National Park Service, we extend our appreciation 
for permission to study the flora and fauna and to collect specimens in 
the Monument. Investigations in the field were supported (in part) by 
a research grant (E-34.07) from the National Institutes of Health, Pub- 
lic Health Service. 


ACCOUNTS OF THE SPECIES 


The records of the snakes (15 species, 150 records, 108 specimens) are 
listed from southwest to northwest, m the Monument and adjacent to 
the Monument (to the north). Unless otherwise indicated, all localities 
are within the Monument and the dates are for the year 1960. 

Several localities mentioned below are not on the map in the folder 
for Joshua Tree National Monument, or on other maps usually avail- 
able. These are as follows, with their position from well known points: 
Cottonwood Spring Y (= 1 mi. NW Cottonwood Spring), Pinto Y (= 
g mi. S Monument Headquarters in Twentynine Palms) and Lower 
Covington Flat Camp, in Riverside County, 0.5 mi. S of San Bernardino 
County line (= 8 mi. S of the town of Joshua Tree ). 

The 108 specimens indicated in parentheses are in the Herpetological 
Collection of Long Beach State College. 


Lichanura roseofusca gracia Klauber 
Desert Rosy Boa 


Specimens examined.—Total 5, as follows: Riverside County: 3.5 mi. 
S Cottonwood Spring Y, AOR, 13 May (1); 3.2 mi. N Cottonwood 
Spring Y, AOR, 16 June (1); Road W of Hidden Valley at junction of 
Stokes Road, 4.000’, 2 May (1); San Bernardino County: Indian Cove 
Camp No. 4, 3300’, 21 May (1). ADJACENT TO MONUMENT, San 
Bernardino County: 0.3 mi. N Monument Entrance, Lower Covington 
Flat, 4.100’, 7 Oct. (1). 

Additional Record.—Total 1, as follows: ADJACENT TO MONU- 
MENT, San Bernardino County: La Contenta Road, 2 mi. S of Twenty- 
nine Palms Highway (3 mi. N of Monument Entrance), AOR, 11 
April (Sight record, E. L. Sleeper). 

All of the Rosy Boas examined were similar in coloration, being typ- 
ical of the desert subspecies to which they are assigned. The northwest- 


Snakes from Joshua Tree National Monument 31 


ern examples have more irregular (serrated) edges to the reddish body 
stripes than do those from near Cottonwood Spring. 


Arizona elegans eburnata Klauber 


Desert Glossy Snake 


Specimens examined.—Total 12, as follows (AOR, unless otherwise 
noted): Riverside County: 1.1 to 1.5 mi. S Cottonwood Spring Y, DOR, 
15 May (1), 16 June (1); 2.7 mi. NW Cholla Cactus Garden, 15 June 
(1); 0.2 mi. W Pinto Y, 24. June (1); 6.8 mi. W Pinto Y, 4400’, 9 Aug. 
(1); Lower Covington Flat Camp, 4.700’, Can Trap, 11 June (1), 17 
July (1). San Bernardino County: 7 mi. S Monument Headquarters, 
Twentynine Palms, 3200’ (2.8 mi. S Monument Entrance), 2 April 
(1); 1.8 mi. N Lower Covington Flat Camp, 6 Sept. (1); 4.6 mi. N 
Lower Covington Flat Camp, 23 July (1). ADJACENT TO MONU- 
MENT, San Bernardino County: 1 mi. W of Joshua Tree, 2900’, 9 Aug. 
(1); 0.5 mi. S, 4, mi. W town of Joshua Tree, La Contenta Road, 14, 
May (1). 

The Glossy Snake is widely distributed throughout the Monument, 
having been taken as high as 4.700 feet. It was the most abundant noc- 
turnal snake at higher elevations. 


Chionactis occipitalis occipitalis (Hallowell) 
Mojave Shovel-nosed Snake 
Specimens examined.—Total 2, as follows: Riverside County: 2 mi. NW 
Old Dale Junction, 2200’, AOR, 15 May (1). ADJACENT TO MON- 
UMENT, San Bernardino County: Twentynine Palms, 2000’, 15 May 
(Oe 


Hypsiglena torquata deserticola Tanner 
Desert Night Snake 


Specimens examined.—Total 5, as follows: Riverside County: 0.5 mi. 
N Old Dale Junction, 2500’, AOR, 14, May (1); 1 mi. S Old Dale Junc- 
tion, DOR, 21 May (1); 1 mi. N White Tank Campground, 3800’, 
AOR, 29 May (1); Lower Covington Flat Camp, both in can trap, 
4,700’, 16 June (1), 23 June (1). 

The specimen from White Tank seems to possess certain character- 
istics of the coastal subspecies, H. t. klauberi. Additional specimens are 
needed to determine if this represents intergradation between the two 
subspecies, or retention of characters of the coastal form as it is being 
submerged by the desert subspecies. 


32 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


Lampropeltis getulus californiae (Blainville) 
California Kingsnake 
Specimens examined.—Total 2, as follows: Riverside County: 3.1 mi. 
N Cottonwood Spring Y, AOR, 15 June (1). ADJACENT TO MON- 
UMENT, San Bernardino County: 1 mi. E Joshua Tree, 2800’ DOR, 15 
May (1). 
Masticophis flagellum piceus (Cope) 
Red Racer 

Specimens examined.—Total 11, as follows: Riverside County: 0.2 mi. 
NE Hidden Valley Entrance, DOR, 24 April (1); 1.5 mi. W Pinto Y, 
DOR, 9 April (1). San Bernardino County: 5 to 7 mi. S Monument 
Headquarters, ‘Twentynine Palms, DOR, 8 May (1), 6 September (1) ; 
2.9 mi. N Lower Covington Flat Camp, 29 May (1). ADJACENT TO 
MONUMENT, San Bernardino County: Twentynine Palms, 31 Au- 
gust 1959 (1); 2 to 3.6 mi. N Monument Boundary, La Contenta Road 
(1 to 3 mi. S, 4.mi. W town of Joshua Tree), March (3), AOR, 8 May 
(1), AOR, 23 June (1). 

This is another widespread species, probably being present through- 
out the Monument. It is more abundant at elevations over 2800 feet. 


Masticophis lateralis lateralis (Hallowell ) 
California Striped Racer 
Specimens examined.—Total 4, as follows: Riverside County: 1 mi. N 
Salton View, DOR, 8 April (1); 1 mi. N Hidden Valley, DOR, 9 April 
(1); Lower Covington Flat Camp, 4.700’, 26 March (1). San Bernar- 
dino County: Lower Covington Flat, 2.5 mi S Monument Entrance, 
4300’, AOR, 2 April (1). 

Additional records.—Total 2, as follows: Riverside County: Eureka 
Point, 2 mi. W Lower Covington Flat Camp, 5300’, 7 August 1959 (1, 
dead, not saved) ; Lower Covington Flat Camp, 4.700’, 30 April (Sight 
record, E. L. Sleeper). 

The striped racer is found in the Pinon-Juniper association at eleva- 
tions above 4.000 feet. The example from San Bernardino County re- 
gurgitated an adult Eumeces gilberti. The easternmost record (Hidden 
Valley) represents a range extension of approximately 26 miles to the 
east of the nearest records from the San Bernardino Mountains. 


Phyllorhynchus decurtatus perkinsi Klauber 
Western Leaf-nosed Snake 


Specimens examined.—Total 20, as follows (AOR, unless otherwise 


Snakes from Joshua Tree National Monument Be 


noted): Riverside County: 1.6 to 5 mi. S Cottonwood Spring Y, 13 May 
(2), 14 May (2), 21 May (1), 28 May (1); 1.4 mi. SW Old Dale 
Junction, DOR, 21 May (1); 0.4. to 4.3 mi. NW Old Dale Junction, 13 
April (1), 14. May (4), 15 May (1); Cholla Cactus Garden, 2 April 
(1); 2 to 4.2 mi. NW Cholla Cactus Garden, 15 April (1), 16 June 
(1); 2 mi. SE White Tank, 29 May (1); White Tank, 3800’, 15 June 
(1). San Bernardino County: 4.mi. S Monument Headquarters, ‘Twen- 
tynine Palms, DOR, 13 April (1), 3 June (1). 

Additional records.—Total 10, as follows (all AOR and released): 
Riverside County: 0.5 to 2 mi. S Cottonwood Spring Y, 21 May (2), 
3 June (1); Old Dale Junction, and 0.3 to 4.3 mi. NW, 21 May (3), 
29 May (1); Cholla Cactus Garden and 2.5 mi. NW, 3 June (2); 2.8 
mi. SE White Tank, 26 June (1). 

This is perhaps the most abundant snake in the Monument although 
it seems to be restricted to the lower elevations, rarely above 3000 feet 
although one record is from 3800 feet. Most of the records (19 of 30) 
are from the month of May. 


Pituophis melanoleucus deserticola Stejneger 
Great Basin Gopher Snake 

Specimens examined.—Total 9, as follows (DOR, unless otherwise 
noted): Riverside County: 0.5 mi. N Pinto Y, 25 March (1). San Ber- 
nardino County: 1 mi. N Pinto Y, AOR, 23 June (1); 4. mi. S Monu- 
ment Headquarters, Twentynine Palms (0.1 mi. S Monument En- 
trance), 3000’, AOR, 22 May (1), 12 April (1). ADJACENT TO 
MONUMENT, San Bernardino County: 3 to 3.5 mi. S Monument 
Headquarters, Twentynine Palms, AOR, 5 March (1), 24, April (1), 
25 April (1); 1 mi. W Indian Cove turnoff (7 mi. W Twentynine 
Palms), 21 May (1); 1 mi. E town of Joshua Tree, 15 May (1). 

This subspecies seems to be restricted to the northern edge of the 
Monument. No examples of the Gopher Snake were taken in the south- 
ern half of the Monument where the subspecies affinis is to be expected. 


Rhinocheilus lecontei lecontei Baird and Girard 
Western Long-nosed Snake 
Specimens examined.—Total 5, as follows: Riverside County: 1.8 mi. N 
Cottonwood Spring Y, AOR, 22 October (1); 0.3 mi. S Old Dale Junc- 
tion, DOR, 12 April (1) and 12 July (1). San Bernardino County: 6 
mi. S Monument Headquarters, Twentynine Palms, DOR, 11 April 
(1). ADJACENT TO MONUMENT, San Bernardino County: Road 
to Indian Cove, 1 mi. N Monument Entrance, DOR, 23 April (1). 


2A, Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


All of the specimens possess the desert pattern usually referred to as 
the subspecies Rhinocheilus lecontei clarus Klauber. All of the localities 
are below 3000 feet in elevation. 


Salvadora hexalepis (Cope) 

Western Patch-nosed Snake 
Specimens examined.—Total 14, as follows (DOR, unless otherwise 
noted): Riverside County: 4,.mi. S Cottonwood Spring Y, 2500’, 14 May 
(1); 1.6 mi. S Old Dale Junction, 27 March (1); 2 mm. S Pinto Y 
(White Tank) 3800’, 12 April (1); 0.5 mi. S to 0.5 mi. N Pinto Y, 26 
March (1), 2 April (2); 1.5 W Pinto Y, 21 Oct. 1959 (1); Entrance 
to Hidden Valley, 23 April (1). Lower Covington Flat Camp, 4.700’, 
caught near camp, 12 April (1). San Bernardino County: 1.5 mi. N 
Lower Covington Flat Camp, AOR, 10 April (1). ADJACENT TO 
MONUMENT, San Bernardino County: 1.5 to 2.5 mi. S Monument 
Headquarters, ‘Twentynine Palms, 28 Oct. 1959 (1), 13 April (1); 1.5 
mi. N Monument Entrance, Lower Covington Flat, AOR, 23 April 
(1); 0.5 mi. S, 4,mi. W town of Joshua Tree, 9 Oct. (1). 

Additional records.—Total 2, as follows: Riverside County: Cholla 
Cactus Garden, 3 June (1, DOR, not saved) ; 4, mi, SE Pinto Y, 21 May 
(1, DOR, not saved). 

No subspecific determination has been made of the specimens of 
patch-nosed snakes from the Monument. The two specimens from the 
south edge of the Monument (near Cottonwood Spring Y and Old Dale 
Junction) seem to represent typical examples of S. h. hexalepis (Cope), 
whereas those specimens at the northern edge of the Monument ex- 
hibit characters of both S. h, hexalepis and S. h. mojavensis Bogert. 
Additional specimens are needed to determine the subspecies present, 
the ranges of each and the areas of intergradation. 


Trimor phodon vandenburghi Klauber 
California Lyre Snake 
Specimen examined.—Total 1, as follows: Riverside County: 1.9 mi. 
NW Cholla Cactus Garden, 2800’, DOR, 5 September 1959 (1). 
This represents a slight range extension to the east from the west side 
of the Little San Bernardino Mountains. 


Crotalus cerastes Hallowell 
Sidewinder 
Specimens examined.—Total 26, as follows (AOR, unless otherwise 
noted): Riverside County: Cottonwood Spring Y, 12 July (1); 2.6 mi. 


Snakes from Joshua Tree National Monument 35 


S to 7 mi. NW Old Dale Junction, 1900’ to 2700’, 12 March (1), 25 
March (7), 26 March (1), 15 April (1), 14, May (1), 15 June (1), 22 
Oct. (1); 3 mi. NNE Old Dale Junction, 12 July (1); 1 mi. SE to 3 mi. 
NW Cholla Cactus Garden, 25 March (1), 27 March (1), 9 April (2), 
13 April (1), 15 April (1), 15 June (1); 0.5 mi. S Belle Campground, 
3800’, 9 May, DOR (1), 14. May (1). San Bernardino County: 5 mi. S 
Monument Headquarters, ‘Iwentynine Palms, 26 March (1); ADJA- 
CENT TO MONUMENT, 1.2 mi. S Monument Headquarters, 2400’, 
g August (1). 

Additional records.—Total 4, as follows: Riverside County: Old Dale 
Junction, 26 March (1, AOR and released) ; 4..3 to 5 mi. NW Old Dale 
Junction, 26 March (1, AOR and released), 29 May (1, AOR and re- 
leased); 4,.m1. NW Cholla Cactus Garden, 26 March (1, DOR, not 
saved ). 

Determination to subspecies was not made for these specimens. 
Klauber (1944: 102) indicated that the area of intergradation between 
C. c. cerastes and C. c. laterorepens Klauber is to be expected along the 
Riverside-San Bernardino County line. Specimens from the northern 
edge of the Monument seem to be C. c. cerastes. Most of the specimens 
were taken below 3000 feet, although one adult was taken alive at dusk 
on the road at 3800 feet near Belle Campground. 


Crotalus mitchelli pyrrhus (Cope) 
Southwestern Speckled Rattlesnake 


Specimens examined.—Total 12, as follows: Riverside County: 2.5 mi. 
NW Old Dale Junction, DOR, 23 April (1); 5.4. mi. NW Old Dale 
Junction, 27 March (1); 1.5 mi. SE Cholla Cactus Garden, AOR, 13 
April (1); Cholla Cactus Garden, 2300’, AOR, 15 May (1); 0.2 mi. 
S Eureka Point, 2 mi. W Lower Covington Flat, 5200’, 12 June (1). 
San Bernardino County: 8 mi. SE town of Joshua Tree (3 mi. SE Mon- 
ument Entrance), AOR, 15 June (1); Lower Covington Flat, 0.3 to 3 
mi. SE Monument Entrance, AOR, 29 April (1), AOR, 15 May (1), 
DOR, 29 May (1), AOR, 11 June (1). ADJACENT TO MONU- 
MENT, San Bernardino County: 3.5 mi. S Monument Headquarters, 
Twentynine Palms (0.5 mi. N Monument Entrance), 15 May (1); 1 
mi. N Monument Entrance, Lower Covington Flat, AOR, 15 May (1) 

The Speckled Rattlesnake has been found throughout the Monument 
in rock habitats. Several specimens were taken in areas some distance 
from conspicuous rock outcrops, but they usually were in higher eleva- 
tons. 


36 Bull. Sou. Calif. Acad. Sct. / Vol. 61, Pt. 1, 1962 


Crotalus viridis helleri Meek 
Southern Pacific Rattlesnake 


Specimens examined.—Total 3, as follows: Riverside County: Lower 
Covington Flat Camp, 4.700’, 11 April (1), 30 April (1); Jumbo Rocks, 
4000’, 11 mi. S Twentynine Palms, DOR, 5 August 1959 (1). 

Additional record.—Total 1, as follows: Riverside County: Lower 
Covington Flat Camp, 4.700’, 23 May (Sight record, E. L. Sleeper). 

These localities represent a range extension of more than 30 miles to 
the east for the southern subspecies which has been reported from the 
San Bernardino Mountains. 


LITERATURE CITED 


KRLAUBER, -I: M. 
1944. The Sidewinder, Crotalus cerastes, with description of a new subspecies. 
Trans. San Diego Soc. Nat. Hist., 10(8):91-126. pls. 6-7, fig. 1 map. 


MONARCHISTIC DOMINANCE IN SMALL GROUPS 
OF CAPTIVE MALE MOSQUITOFISH, 
GAMBUSIA AFFINIS PATRUELIS 
Mesa C. CaLpwE.t! and Davin h. CaLpwe.u” 


Los Angeles County Museum 


Dominance-subordinance interrelations among vertebrates have re- 
ceived considerable attention in recent years. Allee (1952) summarized 
the literature dealing with the more commonly reported of these, the 
peck-right or peck-dominance hierarchy, which has been described 
for all classes of recent vertebrates. A less frequently reported social 
order (also discussed, briefly, in the same paper by Allee) is that in 
which the despot not only dominates all of its associates, but also in- 
hibits any interaction between the remaining members of the group. 
Collias (1944: 90) used the term “‘monarchistic” for this type of social 
organization. 

Such a monarchistic dominance has been reported for captive male 
albino laboratory mice (Uhrich, 1938), for captive male domestic 
cats (Winslow, 1938), by Cooper (1942) for captive African lions 
(males over males and females), and possibly for captive male Ameri- 
can chameleons (Evans, 1936). Allee (1952) mentioned that it is 
known to occur in fishes, but did not cite a specific species. 

We now report the finding of monarchistic dominance behavior in 
captive male mosquitofish, Gambusia affinis patruelis Baird and 
Girard. Monarchistic was the: usual dominance behavior demonstrated 
by the groups studied, in March and April, although one group of four 
fish for a time demonstrated a peck-right (or thrust-right ) domimance 
of sorts in which three of the four interacted (the fourth neither chased 
nor was chased). This display of different types of dominance in 
mosquitofish is thus similar to the variations in types of social orders 
shown for the laboratory mouse (Uhrich, 1938). Although the fish were 
not actually measured, our notes indicate that in each of the dominance 
relations established during this study, the largest fish was the despot. 
This was true in the case of the American chameleons studied by Evans 
(1936), but was not true for the domestic cats reported on by Winslow 


(1938). 
1Also Department of Zoology, University of California, Los Angeles. 


2Also Research Associate, Florida State Museum, and Collaborator in Ichthyol- 
ogy, Institute of Jamaica. 


3, 


38 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


MATERIALS 


Male Gambusia affinis patruelis were collected from a stream running 
through the botany garden on the University of California campus at 
Westwood Village, Los Angeles, California, and were identified using 
keys provided by Hubbs (1926). 


PROCEDURE 


Under varying conditions, up to four male Gambusia were integrated 
in a five-gallon aquarium and the water level reduced to one gallon, to 
reduce the area of confinement. Although in this series of observations 
there were many forms of aggressive behavior, possibly including some 
nipping or biting, thrust was taken as the most clear-cut demonstration. 
The thrust is usually directed at the victim’s flank. Aside from thrust, 
preliminary dorsal fin erection forms a major part of the aggressive 
pattern. 

Fighting, on the other hand, is carried out by the fishes making an 
“*S” shape of their bodies and giving a sharp flip to the caudal fin. This 
propels a current of water against the opponent. On one occasion, two 
Gambusia fought continuously for 15 minutes using these “tail blows” 
(such as those described for other species of fishes by Tinbergen, 1958: 
25), with intermittent thrusts at each other’s flanks, before the domi- 
nant individual was established. 

This dominant individual, or despot, roams freely throughout the 
tank. Once beaten, the subdued individual usually flees from the agres- 
sor. Submission is indicated by holding quite still, or by swimming very 
slowly. The body is held parallel to the bottom of the tank. Fast swim- 
ming almost invariably brings on an attack by the despot. On occasion, 
a submissive individual will elude observation by the despot by smking 
down into the aquarium where it seems to attract less attention. 

Marking.—100 grams of Alizarin Red S dye were dissolved in 100 
cc. of distilled water, and between 0.1 and 0.2 cc. were injected intra- 
muscularly either in the caudal region, in the dorsal fin region, or both. 
This method allows for three separate markings and for one unmarked 
control. 

In a series of 15 markings or remarkings, between April 12 and 
April 21, death was caused twice. Death was immediate, and mechani- 
cal marking injury was considered responsible. At no time was further 
ill effect noticed from this freshly-made solution. It should be noted 
that old Alizarin Red S solution is toxic to fish. On one occasion a solu- 
tion 30 days old was used to mark eight males. Six died immediately 


Monarchistic dominance in male mosquitofish 39 


(apparently not from mechanical injury), one showed distress, and 
one showed no apparent ill effect. 

Specimens retained their stain for varying lengths of time (two to 
seven days) and remarking did not interfere with the experiments. 


RESULTS 
Experiment 1 (13 March) 

In a preliminary observation, definite aggression was shown by one 
of the larger males and in the space of one hour no interaction was 
noted between the other three. The aggressive individual thrust at the 
other three, who would flee from it. The agressive actions by the despot 
were almost continuous. 


Experiment 2 (21 March) 


In this experiment the despot (which suppressed interaction between 
the remaining fish) was removed as soon as its dominance was estab- 
lished (in less than 30 minutes). There was an occasional interaction 
between submissive fish (three such interactions were noted in one 
hour), but this was rare once dominance was established. As the despot 
established himself, he was removed from the aquarium and placed in 
a container marked A. Interaction by the remaining three fish was be- 
gun almost immediately, and when another became dominant (atter 
four to five minutes) he was transferred to a container marked B. Be- 
fore his removal, he suppressed interaction between the remaining two 
fish. When one of these remaining two fish became dominant (after 
one to two minutes ), he was placed in a container marked C. The fourth 
individual was placed in a container marked D. 


Experiment 3 (11 April) 

Four fish were starved for four days and then were integrated. A 
piece of shrimp was introduced and two of the fish carried out a 15- 
minute battle directly above the shrimp. During the fight, both of the 
other fish came under and fed. Once both fish stopped fighting to chase 
off a feeding fish, but immediately resumed their fight. After a domi- 
nant alpha individual was established, he curtailed interaction among 
the other individuals by (1) chasing any fast-moving fish m the 
aquarium—this could be either a chasing or a chased individual, or 
both, but his presence would stop the interaction, and (2) by juxta- 
posing his body (on two occasions) between an aggressor and his vic- 
tim. In this latter instance, all three individuals held quite still. 


40 Bull. Sou. Calif. Acad. Sct. / Vol. 61, Pt. 1, 1962 
Experiment 4 (12 April) 

In this series of experiments, times of establishment of successive 
dominance were noted and the individuals were placed in separate 
tanks marked A, B, C, and D in their successive order of rank when 
the preceeding despot was removed. The time required for A to be- 
come despot was not noted, but after his removal, B became despot in 
10 minutes, and after his removal C became despot in four minutes. A 
was marked in the caudal region, B was left unmarked, C was marked 
in the region below the dorsal fin, and D was marked in both the caudal 
and dorsal regions. 


Experiment 5 (12 April) 

One hour after Experiment 4, the fish were integrated again and the 
same successive ranking manifested itself as each despot was removed. 
A was despot within ten minutes. After A was removed, B became 
dominant over C and D within four minutes, and after the removal of 


TABLE 1 
Number of thrusts in 30-minute periods in Dominance-Subordinance relationships 
of the mosquitofish, Gambusia affinis patruelis, after monarchial hierarchy estab- 
lished. Upper, four fish; Middle, after A removed, three fish remaining; Lower, 
after A and then B removed, two fish remaining. 


(Submissive individual on top of table, aggressive individual on side ) 


A B C D 
A - 8 2 2 
B 1 - 0 1 
C 0 0 - 3 
D 0 0 0 - 
B C D 
- 11 8 
0 - 0 
0 0 
C D 
C - 8 


Monarchistic dominance in male mosquitofish AA 


B, C, became dominant over D in four minutes. This was the same suc- 
cession of monarchy as shown before marking. 


Experiments 6 and 7 (13 and 14 April) 

Experiment 4. was repeated twice within the next two days, with the 
same succession of monarchy resulting, using the same four individ- 
uals. In Experiment 6, the successive times in which the despot as- 
sumed his position were three, five and ten minutes. ‘Times were not 
recorded in Experiment 7. In both of these experiments, the fish were 
well fed before they were integrated, after bemg kept in separate con- 
taimers for nearly 24. hours. 

During Experiments 6 and 7 the number of thrusts made by each 
individual was tabulated. Table 1 shows these data and clearly indi- 
cates that interactions among individuals below the despot were in- 
frequent. It should be stated that they occurred early in the experi- 
ment, though this is not indicated in the table. In addition, thrust fre- 
quency is not very stable. Various factors seemed to influence the num- 
ber of thrusts: (1) The existing despot varies as to degree of aggressive- 
ness, and (2) The existing submissive individuals vary as to degree of 
submussiveness. If the submissive individual flees easily, he is harassed 
more than one that does not—as fleemg almost always stimulates chas- 
ing and thrusting. Also, if the submissive individual remains quiet, 
rather than moving around, he is usually ignored. 


Experiment 8 (20 April) 

In order to observe the effect of prior residence, B of Experiment 4. 
was placed in the aquarium and A was removed to a separate container 
for five days. Then A was returned to the aquarium. B thrust at A who 
did not flee, but could be edged around in a circle by B for the first 
ten minutes. For the second ten minutes, the fish usually avoided each 
other. Then A thrust at B twice within the next ten minutes, after 
which A became completely dominant. The pattern of A’s thrusts at 
B after 50 minutes was as follows: Over a ten-minute period; five 
thrusts, rest, search, rest, six thrusts, rest, seven thrusts, rest, seven 
thrusts, rest, eight thrusts, rest, two thrusts, rest, five thrusts, rest. Dur- 
ing this time, the submissive individual, B, would try to avoid contact 
if approached by the despot, A, or would hold very still. B fled quickly 
if thrust at. 


Experiment 9 (21 April) 
In the last experiment of this series, the effect of hunger on the 
dominance order of the fishes from Experiment 4. was studied. Indi- 


42 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


viduals C and D were separated and starved for seven days. Individuals 
A and B, also separated, were well fed. The four fish were then inte- 
grated and food was introduced. For the first six minutes, D (the 
omega individual in the succession of monarchy) thrust, chased, and 
inflicted tail blows at A, B, and C. The following four mimutes, C was 
dominant, engaging in the same aggressive actions against the other 
three fish. A then engaged B. During this period, C and D were inter- 
mittently feeding when not fighting. For a period of three hours and 
20 minutes there was no continuing despot, and fighting occurred be- 
tween all individuals. After this period, A reestablished himself as the 
alpha despot. Therefore, hunger, like prior residence, played a dis- 
rupting role in the dominance order, but did not have a permanent 
effect. 


SUMMARY 
There is definite dominance and subordination in small groups of 
captive male Gambusia affinis patruelis in which: 


1. The dominance-subordination relationship is not affected by the 
marking methods employed during the study. 


to 


. Usually, instead of a descending peck-order or thrust-order within a 
group, a single individual is dominant and represses aggression by 
other members of the group. This may be termed monarchistic 
dominance. 

3. Within a short while after removal of a despot, a new despot mani- 

fests himself. 

4. The order of succession of monarchy shows constancy over a period 

of time. 

5. Dominance order is disrupted for a brief period of time by prior resi- 

dence of a usually submissive mdividual, and for a much greater 

length of time by starvation of a usually submissive individual. In 

each case, the usually submissive individual becomes despot for a 


brief period. 


jo) 


LA peck-right or thrust-right of sorts is sometimes demonstrated. 


ACKNOWLEDGMENTS 


We wish to thank Dr. Nicholas E, Collias of the Department of Zo- 
ology of the University of California, Los Angeles, for suggesting the 
experiments and for his critical examination of the manuscript. Dr. 
Richard E. Whalen of the Department of Psychology of the same uni- 


Monarchistic dominance in male mosquitofish 43 


versity also made many helpful comments on the completed manu- 
script. 


LITERATURE CITED 
ALLEE, W. C. 


1952. Dominance and hierarchy in societies of vertebrates. Colloques internation- 
aux du centre national de la recherché scientifique. Structure et physiologie 
des sociétés animales, 34: 157-182 pls. 1V-VII 


COLLIAS, N. E. 
1944. Aggressive behavior among vertebrate animals. Psysiol. Zool., 17: 83-123. 


COOPER, J. B. 
1942. An exploratory study on African lions. Comp. Psychology Mono., 17(7): 
1-48. 


EVANS, L. T. 
1936. A study of a social hierarchy in the lizard, Anolis carolinensis. J. Genetic 
Psychol., 48: 88-111. 


HUBBS, C. L. 

1926. Studies of the fishes of the order Cyprinodontes, VI. Material for a revision 
of the American genera and species. Univ. Michigan Mus. Zool., Misc. Publ. 
No. 16., 87 p. 


TINBERGEN, N. 
1958. The study of instinct. Clarendon Press, Oxford, England., 228 p. 


UHRICH, J. 
1938. The social hierarchy in albino mice. J. Comp. Psychol., 25: 373-413. 


WINSLOW, C. N. 
1938. Observations of dominance-subordination in cats. J. Genetic Psychol., 52: 
425-428. 


4A. Bull. Sou. Calif. Acad. Sct. / Vol. 61, Pt. 1, 1962 


G. W. HORN’S LAND GASTROPOD LOCALITY IN ARIZONA. Camp Grant 
in southern Arizona is the designated type locality for several species of verte- 
brates and invertebrates. It is where George H. Horn (Ewan 1955: 52), the 
coelopterist, collected terrestrial molluscs in the 1860’s; he later gave them to 
William M. Gabb in Philadelphia. Three kinds were described as new by Gabb 
(1866). 

E. L. Cockrum (1960), in commenting upon the collecting spot for the type 
material of Onychomys torridus torridus (Coues), a grasshopper mouse, has given 
a history for changes of name and localities for long deactivated Camp Grant. His 
summary was necessitated by the mammal locality being considered in some re- 
cent publications to be in Graham County instead of at the junction of Aravaipa 
Creek and San Pedro River in Pinal County (2160 feet elevation, T. 7 S., R. 16 E., 
8 mi. N. & 5 mi. W. of Mammoth). In Gabb’s paper and H. A. Pilsbry’s mono- 
graph (1940, 1948), the confluence of the streams was indicated; the molluscan 
type material was probably from stream rejectamenta as Pilsbry has noted. 

Although Gabb described three species of minute pulmonates from the locality, 
two remain valid. The Pupa (Modicella) arizonensis was shown (Pilsbry 1948: 
921) to be Pupoides albilabris (C. B. Adams). Pupoides hordaceus (Gabb) [=Pupa 
hordeacea Gabb, 1866] is now represented by only one worn shell from the type 
lot (Pilsbry 1948: 924-5). Helix hornii Gabb 1866 has become Thysanophora 
horni (Pilsbry 1940: 986).—Robert J. Drake, University of British Columbia. 


LITERATURE CITED 


COCKRUM, E. LENDELL 
1960. The type locality of the southern grasshopper mouse. Jour. Mammal. 41 
(4): 515-516. 


EWAN, JOSEPH 

1955, San Francisco as a mecca for nineteenth century naturalists. pp. 1-63 in A 
CENTURY OF PROGRESS IN THE NATURAL SCIENCES. California 
Academy of Sciences, San Francisco. 


GABB, WILLIAM M. 
1866. Descriptions of three new species of land shells from Arizona. Amer. Jour. 
Conch. 2 (4): 330-331, pl. 21. Teste 1 October 1866. 


PMGSBRY. IH. 7AG 

1940. Land Mollusca of North America (north of Mexico). Vol. 1, Part 2. Acad. 
Nat. Sci. Philadelphia, Mono. 3, Vol. 1, Part 2. 

1948. Ibid. Vol. 2, Part 2. 


DEPENDENCE ON TEMPERATURE OF Ca/Mg RATIO OF 
SKELETAL STRUCTURES OF ORGANISMS AND DIRECT 
CHEMICAL PRECIPITATES OUT OF SEA WATER 


GEORGE V. CHILINGAR 
University of Southern California 


RELATIONSHIP BETWEEN Ca/Mg Ratio oF SKELETAL STRUCTURES 
OF ORGANISMS AND TEMPERATURE 


As shown by Chilingar (1953: 206) and Chave (1954.), there is an 
mverse (“hyperbolic”) relationship between the Ca/Mg ratio in the 
skeletons of organisms and the temperature of the water in which they 
live. Figures 2 through 11 show the relationship between the mean 
yearly temperature of sea water and the Ca/Mg ratio’ of various groups 
of organisms, arranged in the order of increasing phylogenetic com- 
plexity. Sixty two per cent of the data plotted in these graphs are based 
on the analyses by Clark and Wheeler (1922) and 23 per cent on the 
results obtained by the writer. Although many analyses of organisms 
by Clark and Wheeler are not accompanied by temperature data, the 
exact description of location and depth enables one to determine the 
temperature from oceanographic literature. Additional data (15 per 
cent) were obtained from Chave (1954), who plotted MgCO, content 
of skeletons versus temperature of sea water, The “Ca/Mg ratio versus 
temperature” curves generally have a “hyperbolic” shape. The average 
Ca/Mg ratios of various groups of organisms, however, are different, 
and there is very little relationship between the average Ca/Mg ratio 
and the phylogenetic level (Figure 1). 

Some scattering of the points in Figure 3 can be explained by the 
fact that samples of Foraminifera analyzed contained both benthonic 
and pelagic forms, whereas the temperature recorded is that of the bot- 
tom water. 

Very high Ca/Mg ratios of the skeletons of madreporarian corals 
(Figure 5) are due to the fact that aragonitic organisms contain very 
small amounts of magnesium. Chave (1954.) demonstrated that ara- 
gonitic organisms seldom contain over 1 per cent magnesium carbonate. 
The Ca/Mg ratio, therefore, largely depends on the mineralogic form 
of the carbonate. For example, in the case of gastropods and pelecypods 


1Weight ratios. The Ca and Mg contents were determined by the writer by using 
wet chemical technique with double precipitation. 


45 


ATIVDILINIOOVAHG 


YSIHOIH 


———= SWC a 


46 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


AVERAGE Ca/Mg RATIO 
5 § & 6 Circus, Be 
(e) 8 {e) (e) {e) (e) 3 


= WwW b (0) 
Oo oO P53 Oo ce) o O ce) Oo 
IBS co ae 
| 
Py [rokaminiFers (| 
| ALGAE 
oe 
A 


Peek teehee 
Pac eae 
MA 


Ci ebetobed 
PP fecbofee 


alas 
Bir pes lirica 


Figure 1. Relationship between Ca/Mg ratio and the phylogenetic level of or- 
ganisms. 


the presence of a few per cent of calcite causes a marked decrease in the 
Ca/Mg ratio. 

The X-ray analysis conducted by the writer showed that madre- 
porarian corals do not contain calcite, whereas the organisms plotted 
in Figures 2, 3, 4, 6, 7, 8, 9, 10, and 11 are devoid of aragonite. 

Even small variations in temperature are reflected in the Ca/Mg 
ratios of organisms. For example, Rhipidogorgia flabellum Linné at 
24.5° and 25° C. had Ca/Mg ratios of 9.04:1 and 8.3:1, respectively 
(Figure 4,). The spines of Tripneustes ventricosus (Lamarck) have a 
Ca/Mg ratio of 14.3:1 at 24.5° C. and 12.1:14 at 26° C. 


Ca/Mg ratio of skeletal structures AT 


EFFrect OF TEMPERATURE ON Ca/Mg Ratios or CHEMICAL 
PRECIPITATES FROM SEA WATER 


Mixtures of CaCO; and MgCO,; have been precipitated from 500 
cc. samples of sea water on adding 300 cc. of saturated solutions of 
Ca(HCO;).>. Figure 12 shows inverse relationship between the tem- 
perature and Ca/Mg ratios of precipitates obtained at the end of 48 
hours. The “hyperbolic” shape of ““Ca/Mg ratio versus temperature” 
curve is possibly due to the rapid rate of precipitation of CaCO, at high- 
er temperatures, which enables CaCO, to trap more MgCOs;. 

In another set of experiments, 300 cc. of saturated solutions of 
Ca(HCO;).2 were added to 500 cc. samples of sea water with enough 
Na.CO; to bring the solution to the verge of clouding. The higher mag- 
nesium content of the precipitates in these experiments (Figure 13) 
was possibly due to the precipitation of magnesium as Mg (OH )>, be- 
cause the solubility product of Mg (OH). is exceeded at a pH of around 


9.49. 
DiscussIoN 


The similarity in shape of “Ca/Mg ratio versus temperature” curves 
of primitive invertebrates and direct chemical precipitates suggests that 
the Ca/Mg ratios of these organisms are either controlled to some ex- 
tent by the conditions in the surrounding environment (namely, varia- 
tion of the solubility products of CaCO,, MgCO;, Mg(OH)s, etc. at dif- 
ferent temperatures ), or that the internal processes somewhat resemble 
the external processes. The former explanation becomes even more 
plausible when one remembers that primitive invertebrates do not have 
autonomous blood systems (open to outside environment) and their tis- 
sues are transfused by sea water. 

Different organisms have different Ca/Mg ratios; hence, the chem- 
ical composition of protective and skeletal structures is not entirely con- 
trolled by the physical-chemical properties of the surrounding environ- 
ment. Inasmuch as different organisms possibly attain different pH 
within their tissues, the Ca/M¢g ratios of their skeletons is probably also 
related to this pH. The writer (1956a: 32) had previously shown that 
the Ca/Mg ratio of direct precipitates decreases with higher pH. 
Examination of Figures 12 and 13 also suggests that the “Ca/Meg 
versus temperature” curves of organisms, which could attain high pH 
(—S> 9.49) within their tissues and precipitate magnesium as 
Mg(OH)>., might approach a straight line. 

It is also interesting to note that the writer (Chilingar, 1956b: 211) 
proved that the Ca/Mg ratios of Strongylocentrotus pur puratus (Stimp- 


48 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


RATIO 


oO 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 
TEMPERATURE ies 


Figure 2. Relationship between the Ca/Mg ratio of Lithothamnium skeletons and 
temperature. Analyses by Chave (1954, p. 273): 1, 2—Lithothamnium sp., Alas- 
ka; 3—Lithothamnium sp., Maine; 4—Lithothamnium sp., California; 5, 6— 
Lithothamnium sp., Japan; 7-8—Lithothamnium sp., Bermuda. Analysis by 
Chilingar: g—Lithothamnium sp., Guam. 


Ca/Mg ratio of skeletal structures 49 


Omorr4e 6 8 10 l2 4 16 te 20 22.124 °26 28° 30°32 
TEMPERATURE cle 


Figure 3. Relation between Ca/Mg ratio in skeletons of Foraminifera and tem- 
perature. Analysis by Clarke and Wheeler (1922, p. 2): 1—Globorotalia menardii 
d’Orbigny. Analyses by Chilingar: 2—Foraminifera sample No. 7, off San Diego, 
California, depth of 2560 feet (Bandy, 1953); 3—Foraminifera sample No. 4, off 
San Diego, California, depth of 380 feet (Bandy, 1953); 4—Foraminifera sample 
No. 1261, off San Diego, California, depth of 375 feet (Bandy, 1953); 5—Vora- 
minifera sample No. 960, off San Diego, California, depth of 260 feet (Bandy, 
1953); 6—Foraminifera sample No. 345, off San Diego, California, depth of 200 
feet (Bandy, 1953); 7—Foraminifera sample off Louisiana, Gulf of Mexico, depth 
of 102 feet; 8—Sorites sp., South of Tortugas, Florida; g—Sample of Foraminifera 
from Bikini; 1o—Miniacina alba Linné, Bahamas; 11—Quinqueloculina auberi- 
ana @’Orbigny, south of Tortugas, Florida; 12—Archaias adunca Fichtel and Moll, 
Key West, Florida; 13—Sorites mar ginalis Lamarck, south of Tortugas, Florida, 
depth of 29.3 meters. 


0 2 4 6 8 10 le 14 16 I8 20° 22 "24m2eures 
TEMPERATURE TAS 


Figure 4. Relation between Ca/Mg ratio in skeletons of Alcyonarian corals and 
temperature. Analyses by Clarke and Wheeler (1922, p. 9): 1—Aleyonium car- 
neum L,, Agassiz, 2—Lepidisis caryophyllia Verrill, 3—Pennatula aculeata Dana. 
4—Rhipidogorgia flabellum Linné, 5—Gorgonia acerosa, Pallas, Florida. Analysis 
by Chilingar: 6—Rhipidogorgia flabellum Linné, Bahamas. 


Ca/Meg ratio of skeletal structures 51 


TEMPERATURE °C. 


Figure 5. Relation between Ca/Mg ratio in skeletons of Madreporarian corals and 
temperature. Analyses by Clarke and Wheeler (1922, p. 6): 1—Flabellum ala- 
bastrum Moseley, 2—Deltocyathus italicus Michelotti, 3—Desmophyllum ingens 
Moseley, 4—Dasmosmilia lymani Pourtales. Analyses by Chilingar: 5—Madre- 
pora sp., 6—Madracis sp., 7—Dendrophyllia sp. 


Bull. Sou. Calif. Acad. Sci. / Vol: 61, Pt. 1, 1962 


Or 
LS) 


2 4 6 8 10 12 14 16 18 20 22 24 26 28 3 
TEMPERATURE =G: 


Figure 6. Relation between Ca/Mg ratio in skeletens of Sea Urchins and tempera- 
ture. Analyses by Clarke and Wheeler (1922, p. 22): 1—Echinus affinis Morten- 
sen, 2—Strongylocentrotus fragilis Jackson, 3—Lytechinus anamesus H. L. Clark, 
4—Tretocidaris affinis Philippi, 3—Echinometra lucunter Linné, 6—Mellita sex- 
iesperforatus Leske. Analyses by Chilingar: 7—Eucidaris sp., Bahamas, 8—Echi- 
nometra lucunter Linné, Bahamas. 


Ca/Meg ratio of skeletal structures 53 


Ca “Mg RATIO 


7 
OM 2 A> 6 8 10 l2 14 16 't8 20 22 e4 26 28 30 
TEMPERATURE FG 


Figure 7. Relation between Ca/Mg ratio in skeletons of Star Fishes and tempera- 
ture. Analyses by Clarke and Wheeler (1922, p. 26): 1—Ctenodiscus crispatus 
Retzius, 2—Benthopecten spinosus Verrill, 3—Plutonaster agassizii Verrill, 4— 
Leptasterias compta Stimpson, 5—Odontaster hispidus Verrill, 6—Ctenodiscus 
procurator Staden, 7—Orthasterias tanneri Verrill, 8—Asterina minuta Gray, 9— 
Linckia guildingti Gray. Analyses by Schmelck (1901): 10, 11—Arcaster tenuts- 
pinus (Diiben and Koren). 


54 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


Oo 2 4 6 8 10 l2 14 16 18 20 22 24 26 28 30 
TEMPERATURE Ce 


Figure 8. Relation between Ca/Mg ratio in skeletons of Ophiurans and tempera- 
ture. Analyses by Clarke and Wheeler (1922. p. 29): 1—Ophiomusium lymani 
W. Thomson, 2—Ophioglypha sarsii Liitken, 3—Ophiocamax fasciculata Lyman, 
4—Ophioglypha lymani (Ljungman), 5—Ophiocoma pumila Litken, 6—Ophio- - 
myxa flaccida Say. 


Ca/Mg ratio of skeletal structures 55 


Ca / Mg RATIO 


@ Ce) (o) 


a 
a 


Figure 9. Relation between Ca/Mg ratio in Crinoid skeletons and temperature. 
Analyses by Clarke and Wheeler (1922, p. 17): 1—Heliometra glacialis (Leach) 
var. maxima (A. H. Clark), 2—Promachocrinus kerguelensis Carpenter, 3—An- 
thometra adriani Bell, 4—Ptilocrinus pinnatus Clark, 5—Florometra asperrima 
Clark, 6—Pentametrocrinus japonicus Carpenter, 7—Hathrometra dentata Say, 
8—Hypalocrinus naresianus Carpenter, 9—Metacrinus rotundus Clark, 10— 
Parametra granulata Clark, :1—Crinometra concinna Clark, 12—Tropiometra 
carinata Lamarck. 


56 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


60 


50 


O©-2 4 6 8 10 le l4 16 I8 20 22 24 26 28 


TEMPERATURE °C. 


Figure 10. Relationship between temperature and Ca/Mg ratio in skeletons of 
Ostracods (analyses by Chave, 1954. p. 273). 


Ca/Mg ratio of skeletal structures Ba 


RATIO 


Ca /Mg 


Ore 4 6 8 10 2 14 16 18 20 22 24 26 28 30 


TEMPERATURE Ge 


Figure 11. Relationship between Ca/Mg ratio in skeletons of Barnacles and tem- 
perature (analyses by Chave, 1954, p. 273). 


58 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 


TEMPERATURE mG 


Figure 12. Relationship between temperature and Ca/M¢ ratio of direct chemical 
precipitates from sea water on adding saturated solutions of Ca( HCOs)>. 


O 10 20 30 40 50 ‘60 70 80 GOFFIOOM IO 
TEMPERATURE bal OF 


Figure 13. Relationship between temperature and Ca/Mg ratios of direct chemical 
precipitates from sea water on adding saturated solutions of Ca(HCOs). with 


NaeCOs. 


Ca/Mg ratio of skeletal structures 59 


son) and Mytilus californianus Conrad are proportional to the Ca/Mg 
ratio of sea water in the aquarium. This finding suggested that possibly 
other invertebrates also assimilate more magnesium in the environ- 
ment having a higher concentration of magnesium. 


CoNCLUSIONS 


The findings of the present study can be summarized as follows: 

1. There is an inverse (“hyperbolic”) relationship between the 
Ca/Mg ratio in the skeletons of organisms and the temperature of the 
water in which they live. 

2. Different organisms have different Ca/Mg ratios and there is very 
little relationship between the Ca/Mg ratios and the phylogenetic level 
of organisms. 

3. Inverse relationship exists between the Ca/Mg ratios of direct 
chemical precipitates out of sea water and the temperature. 

4. The Ca/Mg ratios of direct chemical precipitates out of sea water 
are also controlled by the pH of the medium of deposition. 

5. The similarity in shape of “Ca/Mg ratio versus temperature”’ 
curves of invertebrates and direct chemical precipitates suggests that 
the Ca/Mg ratios of these organisms are controlled to some extent by 
the effect of temperature on solubility products of CaCO;, MgCOs, 
Mg(OH)., etc. The differences in magnitude of Ca/Mg ratio in dif- 
ferent organisms may be related to the growth mechanism and compo- 
sition and pH of the body fluids. 

The future line of research suggested by this work is to make a de- 
tailed study of the variation in the Ca/Mg ratios of skeletal and protec- 
tive structures of organisms on varying the pH and chemical composi- 
tion of the sea water. 


ACKNOWLEDGMENTS 


This author is greatly indebted to Drs. T: Clements, W. H. Easton, K. O. 
Emery, O. L. Bandy, C. M. Beeson, and R. H, Merriam of the Uni- 
versity of Southern California, whose help and critical advice were in- 
valuable in carrying the present study to completion. 

The investigator would also like to express his appreciation to G. P. 
Kanakoff of the Los Angeles County Museum and A. H. Clark of the 
Smithsonian Institution for supplying numerous skeletal and protective 
structures of organisms. 

Some wet chemical analyses have been done by A. Dollar of the 
Griffmn-Hasson Laboratories, Los Angeles, California. The help ex- 
tended by H. A. Lowenstam and W. Orr is also greatly appreciated. 


60 Bull. Sou. Calif. Acad. Sci. / Vol. 61, Pt. 1, 1962 
BIBLIOGRAPHY 


CHAVE, Kk. E. 

1954. Aspects of biogeochemistry of magnesium, 1. Calcareous marine organisms. 
Jour. Geology, 62(3): 266-283. 

CHILINGAR, G. V. 

1953. Use of Ca/Mg ratio in limestones as a geologic tool. Compass, 30(4,): 202- 
209. 

1956a. Note on direct precipitation of dolomite out of sea water. Compass 34.(1): 
29-34. 

1956b. Use of Ca/Mg ratio as a geologic thermometer and bathometer. Abstract 
of paper presented at XX International Geological Congress, Mexico, p. 211. 

1956c. Use of Ca/Mg ratio of limestones and dolomites as a geologic tool. Ph.D. 
Dissertation, University of Southern California, 140 pp. 


BANDY, O. L. 
1953. Ecology and paleoecology of some California Foraminifera, Part I. Jour. 
Paleontology, 27(2): 161-183. 


CLARKE, FE W, and WHEELER, W. C. 
1922. The inorganic constituents of marine invertebrates. U. S. Geol. Survey Prof. 
Paper 124, 62 pp. 


SCHMELCK, L. 
1901. Chemi om syandets faste bastanddele. Norske Nordhlaus Expedition, 9(28): 
1-71. 


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64 Bull. Sou. Calif. Acad. Sct. / Vol. 61, Pt. 1, 1962 


REPORT OF A SCYPHOZOAN STEPHANOSCYPAUS SIMPLEX KIRK- 
PATRICK FROM THE ARCTIC OCEAN. The scyphozoan Stephanoscyphus 
simplex Kirkpatrick has recently been collected from drifting stations in the 
Beaufort and Chukchi Seas. The specimens agree with the description of Kramp 
(1959) in both internal and external characteristics. 

The following data are available for the two collecting stations: 


Station Number 


204 411 

location 

longitude (W) 144° 55’ 165° 48’ 

latitude (N) ANS TNO SAS 
date December 24, 1959 January 30, 1961 
water depth (meters ) 1540 471 
collecting device orange peel bucket orange peel bucket 
collector Roger W. Lewis John Tibbs 


The colony collected from station 411 contained thirty-two individuals clustered 
on a rock, whereas a single loose specimen was collected from station 204, Water 
temperatures at the bottom were not recorded from these locations but from read- 
ings taken at about the same depth and location approximate temperatures can be 
extrapolated: —0.4°C for station 204 (Muguruma 1961) and +0.5°C for station 
411 (Church 1961). 

Stephanoscyphus simplex has previously been reported from several localities 
in the Atlantic Ocean, the Mediterranean Ocean, the Indian Ocean, the waters 
of the Malayan Archipelago, the Gulf of Panama, the Kermadec Trench, the 
Tasman Sea, and the Antarctic Ocean with depths ranging from 430 to 7000 
meters. Temperatures had previously ranged from 0° to 11°C. This report ex- 
tends the range of distribution and temperature tolerance. 

These studies were aided by a contract between the Office of Naval Research, 
Department of the Navy, and the University of Southern California, NR 107-567. 
We also gratefully acknowledge the use of the laboratory facilities of the Allan 
Hancock Foundation and the technical assistance of Yuk Maan Leung for section- 
ing the specimens.—Carolyn Brahm and John L. Mohr, Biology Department, Uni- 
versity of Southern California, Los Angeles 7. 


LITERATURE CITED 
CHURCH, P. E. 


1961. Personal communicaion. 


KRAMEP: Po. 
1959. Stephanoscyphus (Scyphozoa). Galathea Rep. 1:173-187. 


MUGURUMA, J. 
1961. Oceanographic Observations of Fletcher’s Ice Island T-3 Winter 1959-1960. 
Arct. Inst. N. Am. Sci. Rep. No. 10:1-18. 


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BULLETIN OF THES%o 


Southern California “™ 
Academy of Sciences 


LOS ANGELES, CALIFORNIA 


Vot. 61 Aprit-JUuNE, 1962 Part 2 


CONTENTS 


Analysis of the Habitat, Web Design, Cocoon and Egg Sacs of the 
Tube Weaving Spider Diguetia canities (McCook). (Aranea, 
Diguetidae). M. A. Cazier and M. A. Mortenson 


Studies in Nearctic Desert Sand Dune Orthoptera. Part VI. A New 
Genus and Three New Species of Large Sand-treader Camel 
Crickets from the Colorado Desert with Keys and Notes. Ernest 
R. Tinkham 


A New Megahippus from the Barstow Formation San Bernardino 
County, California, Richard H. Tedford and Raymond M. Alf 113 


Report of an Echiuroid Worm Hamingia arctica Danielssen and 
Koren from the Beaufort Sea. Carolyn Brahm and John L. 


Issued June 29, 1962 


Southern California 
Academy of Sciences 


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Mheodore Downs 0 ye ees sage O08 ek Se or President 
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ANALYSIS OF THE HABITAT, WEB DESIGN, COCOON AND 
EGG SACS OF THE TUBE WEAVING SPIDER DIGUETIA 
CANITIES (McCOOk). (ARANEA, DIGUETIDAE) 

M. A. Cazrer' 

University of California, Berkeley 
and 


M. A. MorTENSON 


Southwestern Research Station, Portal, Arizona 


The spider family Diguetidae, also known as tube and net weavers, is 
endemic to North America where it occurs in southwestern United 
States and northern Mexico. It is represented by a single genus, Digue- 
tia, which contains five species (Gertsch, 1949) of which Diguetia 
canities (McCook) is the most common and widespread. It has been 
recorded from west Texas, Arizona, southern Utah, southeastern Cali- 
fornia (Gertsch, 1935:6) and is now known to be abundant in at least 
two areas on the eastern side of the Chiricahua Mountains, Cochise 
County, Arizona. The family is one of four archaic types known to per- 
sist in the southwest (Gertsch, 1949:265). 

The present studies were started on December 20, 1960, when a 
number of cocoons were brought into the laboratory from a thorn shrub 
area located 1.5 miles N.E. of Portal at an elevation of 4600 feet. No 
adult D. canities were found at this time so the cocoons, egg sacs and 
parasites were cultured, worked over in a preliminary fashion and set 
aside awaiting the capture of the spider for verification of the determi- 
nation. This was accomplished on October 31, 1961, when many co- 
coons were collected in the same area and most fresh ones were found to 
have the female in the entrance to the cocoon. ‘Two cocoons were each 
found to contain an immature specimen of Phidippus formosus Peck- 
ham and Peckham and one had an immature specimen of Chiracan- 
thium inclusum (Hentz). In 1960, one cocoon contained an immature 
female of the latter species along with the eggs, deutova and spider- 
lings of D. canities. These two species apparently occupy deserted or 
old cocoons of D. canities for purposes of overwintering but might also 
feed on the spiderlings of the latter as they emerge from the egg sacs 
and enter the passageway. 


HaBITATS 


When it was discovered that the spiders were selecting primarily one 
species of plant in which to hang their cocoons in the Portal area, three 
other areas each with different dominating vegetation cover were either 


1Department of Entomology and Parasitology. 


65 


66 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


surveyed or studied. From two to three miles N.E. of Portal the prin- 
cipal cover plant is Flourensia cernua (Tar bush) imtermixed with 
Ephedra trifurca (Joimt-fir), Parthenium incanum (Mariola), Yucca 
elata (Yucca) and a few species of perennial and annual herbs. Several 
hundred plants were examined on November 28 and December 1 but 
no cocoons were found. In an area from five to six miles N.E. of Portal 
the dominant plant is Larrea tridentata (Creosote bush) sparsely inter- 
mixed with Flourensia cernua (Tar bush) and Parthenium incanum 
(Mariola). An examination of hundreds of plants again gave negative 
results. Chew (1961:8) studying the “Ecology of the Spiders of a Des- 
ert Community” and working on a plot six miles N.E. of Portal, col- 
lected one specimen of D. canities on “Larrea divericata” in 817 spiders 
swept from the above mentioned three plant species. The fourth area, 
dominated by Opuntia, Fouquieria and Agave, gave positive results 
which will be discussed along with the account of the Portal plot. 

Although the female D. canities, when out of her natural environ- 
ment (Fig. 1), is seen to be strikingly marked with white hairs and 
bands on the abdomen and cephalothorax, and with black rings on her 
long legs, she is seldom seen unless prodded out of her hiding place in 
the entrance to the cocoon. When disturbed from her normal venter-up 
position under the cocoon she blends with the background of the foliage 
or the cocoon to a remarkable degree. Most of the nest and web build- 
ing activity takes place at night and although much of the feeding ac- 
tivity probably occurs during the day it would consist of a brief dash 
onto the web to capture the prey and take it back to the cocoon entrance 
to be eaten. Although sometimes difficult, it is easier to look for the 
cocoon and capture her at the entrance where she is guarding the egg 
sacs rather than looking for the spider. 


PortTaAL HaBIraT 


In the study area near Portal the cocoons were found suspended in vari- 
ous species of small leafy shrubs and herbs, beautifully camouflaged to 
match their surroundings (Figs. 2, 3, and 4). Even though they are 
fairly common they are easily missed because of their secretive position 
in the plants and the nature of the materials used in the construction of 
the cocoon. The effectiveness of this camouflage is lost only after the 
leaves have fallen or the plants have dried up for the winter, at which 
time, some of the cocoons have also fallen to the ground and are almost 
indistinguishable from the debris beneath the plants. 

The principal shrubs or small trees in this area are Prosopis juliflora 
(mesquite), Mimosa biuncifera (wait-a-minute), Acacia constricta 


Tube weaving spider biology 67 


(white-thorn), Acacia gregi (catclaw), and Chilopsis linearis (desert 
willow). Intermixed with these are perennial herbs such as Gutierrezia 
muicrocephala (snake-weed) , Solanum eleagnifolium (horse-nettle) and 
many annual herbs including Salsola kali (Russian thistle), Eriogo- 
num abertianum (buck-wheat), Tidestromia lanuginosa, Amaranthus 
palmeri (pig-weed), and A. fimbriatus (Kearney and Peebles, 1951 ). 
Many species of grasses are also represented and most of these plants 
figure prominently in the cocoon making activities of D. canities at 
lease in this locality. According to Gertsch (1949:234) “Cocoon re- 
treats from different areas differ markedly in color, texture and general 
composition.’ The preferred plant was found to be Acacia constricta 
and most of the cocoons were in young plants from one to three feet in 
height and were suspended from four inches to two feet from the 
ground. In some instances the sheet webbing beneath the cocoon was 
im touch with the ground and anchored to it. Other plants used in the 
order of frequency were P. juliflora, G. microcephala, S. kali, M. 
biuncifera and T: lanuginosa. The cocoons are suspended in small open- 
ings in the plants by means of guy lines to adjacent leaves and twigs 
attached to the top half or two-thirds of the cocoon (Fig. 5). In the im- 
mediate vicinity of the cocoon many tangle lines are irregularly at- 
tached to the guy lines and to the various objects making up the cocoon. 
From the lower half or one-third to the edge of the bottom there is an 
area free of supporting lines extending around the cocoon. From the 
edge of the opening at the bottom other guy lines extend radially out- 
ward and shghtly downward to the adjacent vegetation, each connected 
to the other by a loose sheet web. Near the edge of the cocoon there are 
one or more openings in this sheet web which allows the spider access 
to the top surface to gather in prey that fall upon it or in the tangle 
lines around the upper portion of the cocoon. They also have access to 
the bottom of the sheet but whether or not they pull prey through it 
as do the sheet web weavers isn’t known. In a number of instances a 
second sheet is suspended beneath the first and is attached to the 
lower edge of the cocoon by one or more lines that probably act as trap 
lines similar to those constructed by the orb weavers to detect the pres- 
ence of prey in the web. This second sheet may also serve to protect the 
spider from attack from beneath as it apparently does in the sheet web 
weavers (Gertsch, 1949:170). 


Cocoons 


Incomplete or small cocoons tend to be trumpet shaped, narrowing 
gradually from bottom to top, whereas complete or large ones are elon- 


68 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


gate or oval with a narrow constriction at the top and are nearly parallel 
sided to the bottom opening. The narrower, loosely closed end is always 
at the top and the bottom opening is slightly flared and enlarged so as 
to accommodate the female. On the side of the opening opposite to the 
side on which the egg sacs are placed, a few leaves or other debris are 
woven in to form a small protruding lip. In case of threat the female 
may pull this over the opening to protect herself and block entry to 
the passageway extending upward over the egg sacs for the length of 
the cocoon. It may also be used as support while she is completing the 
last egg sac which is very near the opening. These cocoons have been 
reported to be three or more inches in length (Gertsch, 1949:234) but 
in the Portal area they ranged from 20-45 mm., average 31 mm., in 
length and 11-18 mm., average 14 mm. in width. In 1960, one 85 mm. 
long cocoon was found but appeared to have been torn and another 65 
mm. long contained 19 normally arranged egg sacs. None this large 
has been seen since. 

The cocoon is a rather loosely but strongly woven conglomeration of 
materials found in the immediate vicinity (Fig. 6) and also contains the 
bodies of various prey consumed by the female which have been incor- 
porated into the cocoon especially around the open end. The following 
table gives the identifiable contents of 25 cocoons and the total number 
of cocoons in which each type of material was found. 


Number 
Materials Cocoons 
Plants 
Eriogonum abertianum, seeds ..........).). seeker 25 
Acacia constricta, leaves and twigs . ....... 1.2223 21 
Tidestromia lanuginosa, leaves and flowers ................. 20 
Gramineacea, seedsiand stems ..).......)-: Jen 20 
Amaranthus palmeri, seeds .............0 2.5) eee 18 
Amaranthus fimbriatus, seeds ........2..... 2) eee 12 
Gutierrezia muicrocephala, flowers’... ..... >. 2] eae 8 
Solanum eleagnifolium, leaves .......... =. ..)-- see 7 
Prosopis juliflora, leaves and stems ©. .-. 70... 2p eee 6 
Mimosa biuncifera, leaves and stems ......... ssa eee 1 
Salsola kali, leaves... ... 0... 3 s.0..545 0) 00) eee 1 
Twigs, miscellaneous ..... 8.00. ss. sob See 95 


Pebbles:and dirt): . 22 oo3.344 ociwes eee ee ee ee 95 


Tube weaving spider biology 69 


Number 
Materials (continued ) Cocoons 
Excrement 
MIO MSC MEPRER ar etl Se etna e CARR NAR Ue 5 Ae Ny A Re 13 
TRYAIGIOREE: 5 oo: .ot 6 oi RAL NORE Ite aiURS ocala eR ae) ee 15 
(Caattiereypil lieie oi abt ba east ec See anaes i ane eran en ane 3 
Arachnida 
SaliciGlaceeee net hse Prec Mie ac Rede yk 1 
Insecta 
(COMSOVDUEIRA, oes tileeene aaa ent Hiei cy aan ee it eara ce ir aera 12 
Carabidacy 2 Chrysomelidae ...... 8 
ATHTTUCICRS oo na ooo e 2 Staphylinidae ...... 2 
Mialachidae ........ 1 Melyridae ......... 1 
Scarabaeidae ....... Y Clenidace aa 1 
Pleo MICK Amuem Nr ln kann wea eee a mentan Ate 2D 
Hormieidae —) 5.25. 18 Dxpidaeynttenis ceive 1 
Chaleidae ss. 8)... 1 Small parasitic wasps . 19 
HOI OM LCkme Mae «hs iets ia ee epee SRN 95 
@icadellidae......... 25 Chermidae ye 12 
Membracidae ...... 1 uloordae peers: 1 
J SUSTaaUTOUNSTEE) Sven en pues Mee ea ar ee PC ar ng ae ae pe eee 15 
IMinnerdaer et. oe le 4 Anthocoridae ....... 11 
liyaerdaenn 2 5s 7 Mineidaeae. 2555 oe). 1 
IDI cetaMPP Nee kes ee tae Ra aI ad AN 15 
IMiumscidaes...:...8 15 Chironomidae? ..... 1 
Vhysanoptera .......... URN AG ote Saran HUAN Cla ea NS Ht Lid ee 1 
Me pido ptenaperi a ahs aire eta e Mee, Lae aay he alate aan etter 14 
Gelechiidae, Smallimoths <->... - if 
larval cocoons .... 11 Gaterpillarsee ee ae 1 


The exterior of the cocoons consists primarily of small twigs, leaves, 
seeds, pebbles and excrement, tied together with a maze of silk. Cocoon 
building, although not observed, probably begins in June or early July 
and judging from the number of seeds represented, continues into 
late fall. A number of cocoons collected on October 31, 1961, contained 
fresh green leaves of Acacia constricta, fresh flowers of Tidestromia 
lanuginosa, seeds of Amaranthus palmeri and A. fimbriatus m the 
loose outer covering which would indicate building activity in Septem- 
ber and October. By November the females are content to rest in the 
cocoon opening (Fig. 7) and there was no indication of recent con- 


70 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


struction or egg laying activities in the field. Two females, that were 
brought into the laboratory and kept with their cocoons in petri dishes, 
each spun an additional egg sac and laid a few eggs in them around 
the middle of November. Since the females had no building materials 
available, the sacs protruded from the cocoon entrances and were out 
of line with the other sacs because the cocoons were lying on their 
sides. In 1960, by December 20 all the females had died or at least 
departed from the cocoons. 


PREY 


If the remains of prey incorporated in the cocoons is indicative of the 
dietary preferences of D. canities, irrespective of relative abundance, 
they feed primarily on Cicadellids (leafhoppers ), small wasps of many 
species, Formicidae (ants), Chermidae (Psylids or jumping plant 
lice), Diptera (flies), and Anthocoridae (flower bugs or minute pirate 
bugs). However, all the above plus many more insects are common in- 
habitants of the plants in which the spiders occur and it would seem 
that the size of the prey rather than quality governs the selection. With 
only one exception (Scarabaeidae—Diplotazxis ) all the prey was small. 
Although the larval cocoons of the Gelechiidae are listed above with the 
prey, it is not at all certain that they don’t represent building mate- 
rials rather than food. These larval cocoons of Gelechia aulaea Clarke 
(det. by Lloyd Martin) are common in Acacia constricta from mid- 
summer through the winter months and are occupied by the cater- 
pillars until October and November when they drop to the ground to 
pupate. Eleven cocoons of D. canities were found to contain from one 
to six of these Gelechia coccons and since the latter are firmly attached 
to the plants and seldom drop to the ground during the spider’s period 
of activity, it is assumed that some, if not all, were collected on the 
plants and therefore might have contamed caterpillars. The remains of 
a caterpillar were found in one D. canities cocoon. 


Hasrirat, 7.8 m1Les N.E. oF Porta 


Another population of D. canities was located 7.8 miles N.E. of Portal 
on an east facing limestone hillside where the vegetation cover was 
entirely different from that on the adjacent valley floor and at 1.5 miles 
northeast of Portal. The principal plants in the order of their abun- 
dance were as follows: Fouquieria splendens (Ocotillo) , Opuntia phaea- 
cantha, Opuntia engelmanni, Ferocactus wislizent (Barrel cactus), 
Acacia gregii (Catclaw), Parthenium incanum (Mariola), Callian- 
dra sp. (False mesquite), Larrea tridentata (Creosote bush), Agave 


Tube weaving spider biology 71 


schottu (Century plant), Flowrensia cernua (Yar bush), Gutierrezia 
muicrocephala and a number of small herbs. The dominant vegetation 
along the base of the hill and extending on out into the valley was L. 
tridentata (Creosote bush) and F. cernua (‘Tar bush). Both plants were 
also represented on the hillside although outnumbered by the first 
seven species listed above. 

On November 29, 1961, 25 cocoons were collected in an area rough- 
ly 100 yards square, all plants being examined. Fourteen were sus- 
pended between the leaves of Opuntia phaecantha which is a low grow- 
ing nopal that usually has grass, Calliandra sp. or other vegetation 
growing around it, seven were well camouflaged in Parthenium in- 
canum, three were suspended between low leaves of the taller growing 
Opuntia engelmanii and one was exposed in a dead Gutierrezia micro- 
cephala plant. Even though Creosote bush and ‘Tar bush were inter- 
mixed with the above plants no cocoons were found on them which was 
also true of two other areas examined in which they were the dominant 
species. Neither of these plants are represented on the Portal plot where 
D. canities is known to be present. 


Cocoons 


The cocoons that were suspended in Opuntia (Fig. 8) were made up 
primarily of materials from underlying or adjacent plants and only 
three out of 17 contained recognizable pieces, spines (5), from the 
cactus. Most of the leaves that were used were from Calliandra sp. 
Those suspended in Parthenium were made up primarily of materials 
from this plant with a liberal use of leaves especially on the outer cov- 
ering. The coccon on Gutierrezia was an old one but appeared to con- 
sist mostly of materials from this plant with the addition of pebbles 
and grass stems. All were found within a foot of the ground. The in- 
sect remains incorporated in the cocoons were in almost the same rela- 
tive numbers and kinds as in the Portal plot. In the 25 cocoons there 
were three live and two dead D. canities, one live Chiricanthium in- 
clusum and one dead Phidippus sp. which represents a considerable 
reduction in adult population of D. canities in a month’s time between 
the two collecting dates—October 31 and November 29. 

In D. canities the cocoons appear to serve in three capacities for the 
benefit and survival of the species. (1) As a protective covering for the 
egg sacs; (2) As a centralized protective and feeding retreat for the 
female; (3) As a central anchor for the sheet web that spreads out be- 
neath it for trapping prey. For what is probably its primary function, 
that of protecting the egg sacs against the climatic elements, predators 


72 Bulletin So. Calif, Academy Sciences / Vol. 61, Pt. 2, 1962 


and parasites, it has a number of advantageous features but unfortu- 
nately these are not entirely infallible. Its camouflaged construction, 
the space at the entrance and habit of the female to repose there prob- 
ably have a deterring effect on predators and parasites but some of 
these have found ways of getting around these protective features. The 
construction of the cocoon and arrangement of the egg sacs appear to 
afford the best protection against the climatic elements and probably 
developed as such because the species overwinters in three vulnerable 
stages—eggs, deutova and spiderlings. 


Eco Sacs 


In many spiders the female lays a mass of eggs, covers them with a 
silken sheet and then molds the mass into the egg sac characteristic of 
her species (Gertsch, 1949: 33). Others use viscid secretions or fine silk 
lines to hold the eggs together. D. canities apparently uses neither of 
these techniques as the eggs are loose in the sac, the sac is only about 
half to three-quarters filled and its surfaces do not bear the imprints of 
the enclosed eggs. The fact that the eggs are laid loose or at most with 
a temporary adhesive probably accounts for why the sacs are only par- 
tially filled. Egg sac construction and egg laying have not been 
observed. 

The spinning of the egg sacs and laying of the eggs begins at the top 
of the cocoon since the contents of the upper sacs are more advanced 
in development than those at the lower end, at least by the end of Octo- 
ber. The sacs are spun on one side of the hollow interior of the cocoon, 
they are shingled one on top of the other, with the inner edges elevated — 
above the outer edge at about a 45° angle to the long axis of the cocoon 
(Fig. 9). The first sac is located a short distance in from the narrowed 
upper end of the cocoon and the last is immediately inside the cocoon 
entrance with its lower outer edge almost flush with the edge of the 
opening (Fig. 10). When the female is inside the opening (Fig. 7) she 
is covering this last egg sac. The passageway through the cocoon nar- 
rows down immediately in front of the last egg sac and while it would 
allow for movement of the spiderlings it appears to be too small for the 
female to move through. Each sac is individually surrounded by a net- 
work of pale yellow silk attaching it to the margin of the cocoon and 
to the adjacent sacs. A heavier layer of silk, extending from side to 
side on the cocoon, covers all the sacs (Fig. 11). The opposite wall of 
the cocoon is also covered with silk but not as densely. 

Each egg sac is lenticular or discus shaped, from 5 to 6 mm. in di- 
ameter, and is made of pale yellowish silk woven so tightly as to give 


Tube weaving spider biology 73 


them a parchment-like appearance. The edges of the two halves are 
rather loosely silked together, probably to enable the spiderlings to 
escape readily (Fig. 12). In 54 cocoons gathered near Portal in 1961, 
the number of egg sacs in each varied from 2-10 with an average of 4.6. 
In 25 cocoons taken in 1961, at 7.8 miles N.E. of Portal, they varied 
from 1-8 egg sacs in each with an average of 3.9 per cocoon, The 
shingled and angled arrangement of these sacs would be a definite 
advantage during wet weather if moisture seeped in at the top of the 
cocoon. The first sac would direct it downward and outward, thus pro- 
tecting the remaining sacs beneath it. 


Eces 

The eggs are opaque, white, cylindrical and vary from .75-.80 mm. in 
diameter (Fig. 13). They are laid free in the sacs, never completely fill 
it and there is no impression of the eggs on the side wall of the sac. The 
deutova (Fig. 14) vary from .80-.95 mm. in lengths, the cephalothorax 
and legs are white, the eyes are black and the abdomen is a pale brown. 
The first molt spiderlings vary from 1.0-1.5 mm. in length and the 
color is the same as in the deutova. In 50 egg sacs taken at random the 
number of eggs or deutova in each varied from 41-164 with an aver- 
age of 113. Since there is an average of 4.6 and 3.9 sacs per cocoon this 
would give an average reproductive potential of 520 offspring per fe- 
male at Portal and 441 at 7.8 miles N.E. of Portal. However, as pointed 
out by Gertsch (1949:35) spiders that lay many more eggs than D. 
canities have a survival average of only one pair per female. This figure 
seems inordinately low for the maintenance of the poulation and it 
would be expected that more would survive under favorable environ- 
mental conditions and there would therefore be a fluctuation in the 
population from year to year. 

When multiple egg sacs are spun by a single female, the number of 
eggs tends to decrease in the later ones (Gertsch, 1949:35). In D. 
canities there is some irregularity in individual cocoons, for example, 
one had the following number of eggs in the order laid: 117, 150, 137, 
141, 109; and another 101, 147, 133, 130, 111, 94. However, if the 
average number is taken for the last five sacs in 25 cococns in the order 
laid it is 135, 132, 121, 114, 86 or a gradual decrease as is known to 
occur in some other species. When the cocoons were collected near 
Portal on October 31, 1961, all but one of the last sacs laid contained 
eggs, the other had deutova; in the next to last, 21 contained eggs, four 
had deutova; and in the third from the last, 11 contained eggs, eight 
had deutova, four contained both eggs and deutova, one had a preda- 


74, Bulletin So. Calif, Academy Sciences / Vol. 61, Pt. 2, 1962 


ceous larva and the last a parasite larva. In the cocoon that had ten egg 
sacs, the first four laid contained 1st molt spiderlings, the next three 
deutova, the eighth had 100 deutova and eight eggs and the last two 
were still in the egg stage. Out of the 25 cocoons only five had spider- 
lings in a total of ten egg sacs. 

In the 25 collected at 7.8 miles N.E. of Portal on November 29, 18 
of the last sacs laid contained eggs, two deutova, three were empty be- 
cause of predator activity and two had parasite larvae and pupae. In 
the 22 next to last sacs laid, 13 contained eggs, three deutova, one eggs 
and deutova, three a predaceous larva and two parasites. In 19 third 
from the last sacs, six contained eggs, eight deutova, three a predaceous 
larva, one parasite, one eggs and deutova. Predatism and parasitism 
appears to be higher in this location than at Portal and might be due to 
the more exposed position of the cocoons on the Opuntia plants (Fig. 
8). No parasites or predators were found in the seven cocoons on P 
incanum which were well camouflaged (Fig. 15). 


PREDATISM AND PARASITISM 


Predatism and parasitism are both operating against the early stages 
of D. canities. One predaceous larva will feed on the contents of as 
many as three egg sacs, thus destroying around 340 potential spiders 
and a single parasite larva working in one sac will destroy an average 
of 113 eggs or deutova to complete its development. So far as known 
no parasites of D. canities have previously been recorded and although 
two wasp parasites have been reared, the true association has been estab- 
lished for only one of these, Arachnophaga picea (Riley). This wasp 
(Figs. 16 and 17) is ant-like in appearance and behavior, belongs in 
the family Eupelmidae and has previously been recorded as a parasite 
of Chrysopa californica, Argiope argentata, A. sp., Epeira sp., and 
Phidippus opifex (Muesebeck, Krombein and Townes, 1951:512). 
The manner in which the egg is deposited in the egg sac of D. canities 
has not been ascertained but apparently the female lays only one per 
sac as only one larva is found in each (Fig. 18). As early as October 31 
these larvae are ready to pupate but in 1960 on December 20 both lar- 
vae and pupae were found in the sacs and the adults emerged in the 
laboratory between January 8 and February 3, 1961. Pupation takes 
place in the sac (Fig. 19) after all the D. canities eggs and deutova 
have either been eaten or destroyed. On November 29 one larva of 
A. picea was found with an ectoparasitic larva on its anterior end which 
killed it, but thus far we have been unable to rear the adult for deter- 
mination. 


Tube weaving spider biology AB; 


Figure 2. Suspended cocoon of Diguetia canities (McCook) in Salsola kali L. 
Photo by Mortenson. 


Figure 3. Suspended cocoon of Diguetia canities (McCook) in Acacia constricta 
Benth. Photo by Mortenson. 


Figure 4. Top view of Diguetia canities (McCook) cocoon suspended in Acacia 
constricta Benth. Photo by Mortenson. 


Tube weaving spider biology WH. 


Figure 5. Cocoon of Diguetia canities (McCook) showing guy lines, tangle 
lines and sheet web. Photo by Mortenson. 


Figure 6. Cocoon of Diguetia canities (McCook) showing composition. Photo 
by Mortenson. 


78 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


Figure 7. Entrance to cocoon of Diguetia canities (McCook) showing female in 
opening. Photo by Mortenson. 


Figure 8. Cocoon of Diguetia canities (McCook) suspended in Opuntia phaea- 
cantha, Photo by Mortenson. 


Tube weaving spider biology 79 


‘ : 
Figure 9. Cross section of cocoon of Diguetia canities (McCook) showing cover- 
ing over ege sacs and angle at which they lay. Photo by Mortenson. 


Figure 10. Entrance to cocoon of Diguetia canities (McCook) showing position 
of last ege sac and flap made of two rabbit droppings. Photo by Mortenson. 


80 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


Figure 11. Cross section of cocoon of Diguetia canities (McCook) showing cov- 
ering over egg sacs on bottom of passageway. Photo by Statham. 


Figure 12. Cross section of cocoon of Diguetia canities (McCook) with silk 


cover removed and egg sacs opened. Photo by Statham. 


Tube weaving spider biology 81 


Figure 13. Eggs of Diguetia canities (McCook) in egg sacs. Photo by Morten- 
son. 


82 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


Figure 14. Deutova and shed corions of the eggs in the egg sac of Diguetia 
canities (McCook). Photo by Mortenson. 


RG 


‘| =) 4. > ee ‘ : 5 xe 
t. a’ a. ie Ved 
Figure 15. Cocoon of Diguetia canities TS suspended in Parthenium in- 
canum. Photo by Mortenson. 


bee 


y 


Tube weaving spider biology 82 


Figure 16. Adult (dorsal view) of Arachnophaga picea (Riley). Photo by Mor- 


tenson. 


Figure 17. Adult (lateral view) of Arachnophaga picea (Riley). Photo by 
Mortenson. 


84 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


Figure 18. Mature larva of Arachnophaga picea (Riley) in egg sac-of Diguetia 
canities (McCook). Photo by Statham. 


Figure 19. Pupa of Arachnophaga picea (Riley) in egg sac of Diguetia canities 
(McCook). Photo by Mortenson. 


Tube weaving spider biology 8 


Figure 20. Larva of Cymatodera sp. in egg sac of Diguetia canities (McCook). 
Photo by Mortenson. 


Figure 21. Pupal cell of Cymatodera sp. in egg sac of Diguetia canities (Mc- 
cook). Photo by Mortenson. 


86 Bulletin So. Calif, Academy Sciences / Vol. 61, Pt. 2, 1962 


In 1960 no predaceous larvae or signs of their work were encoun- 
tered in any of the egg sacs examined at the Portal plot. In 1961 they 
were found to be not uncommon in this location and at 7.8 miles N.E. 
of Portal were even more abundant and had destroyed the contents of 
a number of egg sacs. Although none have been reared as of this writ- 
ing they belong to the beetle family Cleridae and probably to the genus 
Cymatodera (Fig. 20). Entry into the egg sac is made through the 
bottom or sides where small holes are cut by the larva. The adult beetle 
probably lays her eggs on or nearby the cocoon from where the larvae 
work their way into the egg sacs. As many as three larvae were found 
in some of the larger cocoons but the usual number was one. Each larva 
consumes the contents of from one to three egg sacs and usually pu- 
pates within one of the sacs although one pupal cell was found just out- 
side and another could not be found even though three sacs had been 
destroyed. They move from sac to sac by cutting holes in the parch- 
ment-like side walls near the bottom and are thus concealed from view 
from above. 

The pupal cell (Fig. 21) is constructed of coarse white strands, silk- 
like in appearance, on the outside which are firmly anchored to the 
spider silk and egg sac walls and an inner layer of finer strands which 
line the cell. These layers harden and give the cell a fairly rigid outer 
covering. This substance is probably the oral exudate which is used by 
many Clerids in the construction of their pupal cells (Balduf, 1935: 
110). The larvae of Cymatodera have previously been reported as 
predators of many kinds of wood boring beetles, larvae of various hy- 
menoptera and moth caterpillars (Balduf, 1935:107-111) but this 
appears to be their first recorded invasion of the Aranea. 


DIscussION AND SUMMARY 


It is known that in many desert areas these spiders prefer various spe- 
cies of cactus for their nest building sites. However, with a species as 
widespread as D. canities such plants are not always available and it 
is no surprise that their distribution is neither limited by nor confined 
to regions where such succulent vegetation dominates. In an area 
roughly eight miles in extent along the eastern foothills of the Chiri- 
cahuas between Portal and Harris Mountain there are four adjacent 
but distinct types of vegetative cover, two of which are occupied by 
D. canities. 

At 1.5 miles N.E. of Portal the dominant shrubs in this thorn shrub 
area are Acacia constricta and Prosopis juliflora which accur in about 
equal numbers but the spiders are found mostly on the former. In the 


Tube weaving spider biology 87 


next and adjacent area extending from two to three miles N.E. of Por- 
tal the dominant plant is Flowrensia cernua in which no D. canities 
were located. It is doubtful that this represents a sampling or survey 
error since hundreds of plants were examined closely on two different 
occasions. Their absence might better be explained by the fact that the 
plant is resmous, has a hop-like odor and there are not as many small 
insect prey in this type of vegetative cover as in the thorn shrub area. 
The next area surveyed was from three to six miles N.E. of Portal in 
which the dominant plant was Larrea tridentata sparsely intermixed 
with Flourensia cernua, Here again no spiders or their cocoons were 
found even though the same area was covered where Chew found one 
specimen on Larrea. This one may have been moving around from 
adjacent Parthenium incanum plants in which it is now known they 
construct their cocoons. The characteristic open growth at the base of 
the Larrea plants, the paucity of subsidiary growth beneath them and 
their strong characteristic odor might be the determining factors inhib- 
itmg or preventing the establishment of colonies of D. canities in this 
type of vegetative cover. Larrea also has a limited insect fauna as com- 
pared with many other plants. 

Since both of these plants were absent from the Portal plot there was 
no way of checking on them in this known D. canities colony. How- 
ever, at 7.8 miles N.E. of Portal the spiders were found in an area domi- 
nated by two species of Opuntia, the low-growing O. phaecantha being 
preferred, and even though both Larrea and Flowrensia were inter- 
mixed and in close contact neither contained cocoons. Parthenium in- 
canum which when growing in Larrea or Flourensia contained no 
cocoons, had them when in association with Opuntia. 

When the female D. canities constructs her cocoon in herbaceous or 
woody plants, she utilizes materials from these as well as from adjacent 
plants and material found on the ground. When they are suspended 
in succulents such as Opuntia, practically all of the building materials 
are obtained from the plants growing beneath or around these plants. 

The cocoons are usually suspended in openings within two feet of 
the ground and each has one and sometimes two sheet webs beneath 
for capturing prey which consists primarily of ants, small parasitic 
wasps and leafhoppers. The cocoons contain an average of from 3.9 to 
4.6 egg sacs and each of these has an average of 113 eggs. There is an 
overall decrease in the number of eggs laid between the first and last 
egg sacs and the upper sac is the first one constructed. 

One parasite, Arachnophaga picea (Riley) a wasp belonging to the 
family Eupelmidae, has been reared from the egg sacs of Diguetia 


388 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


canities and it has an ectoparasite on its larva which has not as yet been 
reared. The primary predator of the immature stages is a beetle be- 
longing to the family Cleridae and probably to the genus Cymatodera. 


ACKNOWLEDGMENTS 


Our appreciation is expressed to Dr. Willis J. Gertsch of the American 
Museum of Natural History for identifying the spiders, to Dr. B. D. 
Burks of the U. S. National Museum for the name of the parasite and 
to Miss Marjorie Statham of the American Museum of Natural His- 
tory for supplying some of the photographs. 


LITERATURE CITED 

BALDUF, W. V. 

1935. The Bionomics of Entomophagous Coleoptera. John S. Swift Co., Inc., pp. 
1-220. 

CHEW. R. M. 

1961. Ecology of the Spiders of a Desert Community. J. New York. Ent. Soc., V. 69: 
5-41. 

GERTSCH, W. J. 


1935. Spiders from the Southwestern United States, with Descriptions of New 
Species. Amer. Mus. Novitiates, 792:1-31. 


1949. American Spiders. New York: D. Van Nostrand Co., Inc., 285 p. 
KEARNEY, T. H. anp PEEBLES, R. H. 
1951. Arizona Flora. Berkeley: Univ. of California Press, 1032 p. 


MUESEBECH, C. EF W; KROMBEIN, Kk. V., ann TOWNES, H. Kk. 


1951. Hymenoptera of America North of Mexico, Synoptic Catalog. U. S. Dept. 
Agriculture, Agriculture Mono. No. 2, 1420 p. 


STUDIES IN NEARCTIC DESERT SAND 
DUNE ORTHOPTERA 


Part VI 


A NEW GENUS AND THREE NEW SPECIES OF LARGE 
SAND-TREADER CAMEL CRICKETS FROM THE COLORADO 
DESERT WITH KEYS AND NOTES 


Ernest R. TINKHAM 
Indio, California 


This is another treatise on the Sand Dune Fauna completed under a 
erant from the National Science Foundation. 

For quite some years now, especially during the course of four sum- 
mers of extensive exploration and intensive study on the sand dune 
biotae of the North American Deserts and with the accumulation of 
considerable materials representing well over a dozen new species, it 
has been increasingly apparent that the sand-treader camel cricket 
genus Ammobaenetes Hubbell 1936, is not a congeneric one. 

The genotype of Ammobaenetes is Dahinia phrixocnemoides Cau- 
dell 1907, a small Rhaphidophorid possessing, in the apical half of its 
straight caudal tibiae, a closely packed group of 5 to 6 pairs of long 
aciculate spurs forming a “sand basket” by which it jumps and digs 
readily in loose drifting sand and from which it derives both its scien- 
tific and common names. The female has a rather long, straight, slen- 
der ovipositor whose length is approximately one-half that of the body 
length and with which she oviposits rather deeply in the loose sand. 

Restricted to, and existing on, the sand dunes of the Colorado Desert 
and dune areas peripheral to it but pertaiming to the Gila Desert, both 
eremological components of the Great Sonoran Desert, is a group of 
much larger sand-treaders whose body size is two to three times that of 
members of true Ammobaenetes as based on the genotype. The large 
males of this group possess strongly arched or bowed caudal tibiae and 
the ovipositor of the female is very short and heavy, its length about 
one-sixth body length, or approximately the length of the pronotum. 
All species of this sand-treader complex have these important features. 
Their habits and life zones differ from those of the genus Ammobae- 
netes and it is obvious that here, contained in the genus Ammobaenetes, 
is a generic entity quite distinct from that genus. It is the purpose of 
this paper to describe this generic entity and the various species repre- 
senting 1t. 

To facilitate the student, a provisional key is here presented. 


89 


gO 


— 


bo 


Ov 


OV 


Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


ProviIsIONAL GENERIC KEY TO THE SAND-T READER 
CAMEL CRICKETS AND ALLIES 


. Mesotibiae with 3 to 5 pairs of dorsal spines( sometimes irregularly 


placed) exclusive'cf the’calcars -....... 2... 7. 2 ae 3 


Mesotibiae with 2 pairs of dorsal spurs (sometimes only 3) exclusive 
of the-calcars <0 2. vei id nen ae et be oe 2 


. Sand basket present and formed by crowding apically of 4 pairs of 


long aciculate spurs. Ovipositor equal to the pronotal length 
5g An Bens OM Ot ee ene Ne We dee on pee Rhachocnemis Caudell 


Sand basket absent. Ovipositor exceeding the length of the pronotum 
cee Ceuthophilus, Pristoceuthophilus, Udeopsylla, Styracosceles 


. Sand basket formed of 4 to 6 pairs of long aciculate spurs, somewhat 


flattened on their mner faces and crowded apically on the caudal 
tIDIAC™ ecco hi bk oe eh ne ee ee Sal eee 5 


Sand basket absent .....-. <. 02 0.2 J.0<) 4). eee 4 


. Caudal tarsomeres—3, their distoventral angles well rounded. Ovi- 


positor 1.5 to 2.0 times their pronotal length. ..Dathinia Haldeman 


Caudal tarsomeres—4, their distoventral angles strongly acute. Ovi- 
positor equal to pronotal length ............ Dathiniodes Hebard 


. Size small; external inferior keel of caudal femora untoothed or with 


a few minute teeth. Caudal tibiae straight. Ovipositor long, slender, 
about one-half body length or twice pronotal length ........ hex Sarg 


Size medium to very large; external inferior keel of caudal femora 
bearing strong teeth. Caudal tibiae straight or strongly arched. Ovi- 
positor short and heavy, its length about the pronotal length .... . 6 


. Size medium to medium large. Caudal tibiae strongly arched in the 


male, female straight. External inferior keel of caudal femora with 
row of uniform strong teeth; internal keel also dentate. Tarsomere 
ratio 3-4-4, the distoventral angles spinose or acute 

ee rere er Ae OL0IAOZIES Il. GBA. 


Size medium to very large. Caudal tibiae straight in both sexes. Ex- 
ternal inferior keel of caudal femora with 2 to 4 very large spike-like 
teeth centrally situated on the keel, preceded and followed by smaller 
teeth. ‘Tarsomere ratio 3-4-4, the distocentral angles spmose or acute 
Pe Ronn tre Oe os Re ew Be Daihinibaenetes Tinkham 


New sand-treader camel crickets g1 


7. Tarsomere ratio 3-4-4, their distoventral angles well rounded 


I ee PsP ke Walle wide IB oN gl als Dathiniella Hubbell 


Tarsomere ratio 3-4-3; their distoventral angles spined or acute 
5 010 6 6 6.5) 5, 8 NOE eet ee ean Sree Sooner Ammobaenetes Hubbell 


Macrobaenetes NEW GENUS 


The new genus Macrobaenetes is amply distinguished from Ammo- 
baenetes Hubbell by many characters as: much larger size, the strongly 
arched or bowed caudal tibiae of the male, the dentition on dorsal ridge 
and inferior keels of the caudal femora which in Ammobaenetes is 
nondentate; by the tarsomere ratio of 3-4-4 instead of 3-4-3 as in Am- 
mobaenetes; by the long first caudal tarsomere which im Ammo- 
baenetes is very short; by the short ovipositor bemg about half the 
length observed in Ammobaenetes in relation to body length and by 
other minor features as well. 

The males of Macrobaenetes are distinguished from all other large 
sand-treaders by the strongly arched caudal tibiae; the female separated 
from Dathinibaenetes Tinkham by the fine teeth on the external in- 
ferior keel of the caudal femora and by the same character from 
Dathinia and Daihiniodes which do not possess sand baskets, that is, at 
least 5 pairs of long aciculate spurs, somewhat flattened internally and 
crowded apically and without any small teeth intervening between 
these long spurs. 


Description: Form typical of the heavier-bodied Rhaphidophorids. Max- 
illary palpi with all segments slender and elongate, the apical palpo- 
mere broadened and recurved in the distal two-thirds, its ventral sur- 
face sunken or excavate to leave a sensitized periphery; labial palpi 
with segments elongate, the apical segment enlarged in distal half. 
Pronotum with foremargin slightly concavely emarginate; lobes of 
all thoracic notites with slight narrowly reflexed margins. Leg spina- 
tion as follows: forefemora unspined dorsally, ventrally unspined ex- 
cept for single tooth on internal keel. Foretibiae unspined dorsally ex- 
cept for small uniform pair of calcars; ventrally with 4 pairs of long 
tapering spurs plus the apical calcars, the external row much the larger. 
Foretarsi trimerous, segments 1 and 2 very short, segment 3 twice their 
combined length; distoventral angles of 1 and 2 lobate, of 3 acute. Mid- 
dle legs with meso femora unspined dorsally and ventrally except for 
a tooth on each lower genicular lobe. Mesotibiae dorsally with 3 to 4 
pairs spines plus apical calcars; ventrally with 3 pairs smaller spurs, 


Q2 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2. 1962 


spaced as dorsally, plus apical calcars. Mesotarsomeres 4, segments 2 
and 3 very short, 1 equal to 4 and each twice length of 2 and 3, disto- 
ventral angles acutely rounded in 1 and 4, acutely lobate in 2 and 3. 
that of 2 much the largest. Hind legs with semiappressed teeth on the 
dorsal ridge; external inferior keel with 16 to 18 large heavy teeth and 
internal inferior keel with 14-16 much smaller ones. Teeth in the fe- 
male much smaller. Caudal tibiae of male strongly arched; the ventral 
keel unspined except for median appressed subapical tooth and pair of 
small attingent calcars; the dorsal keels with 7 evenly paired spurs, 5 
pairs of which form the sand basket in the apical quarter plus the long 
apical calcars which are also part of the sand basket. Caudal tarso- 
meres—4, segments 2 and 3 very short, segment 4 twice the length of 
2 and 3, segment 1 about one and one-third times the length of 4; dis- 
toventral angles similar to those of mesotarsus. Genitalia with supra- 
anal plate roundly triangular, deflexed; cerci long, acuminate, hirsute: 
subgenital plate with rather strong forcipate arms. Ovipositor short, its 
length shghtly less than the pronotal length; dorsal valvulae terminat- 
ing in acute tooth; ventral valvulae bearing 4 uncinate apical hooks. 


Genotype: Macrobaenetes kelsoensis new genus and new species. 


The generic name refers to the large size of these sand treaders which 
distinguished them from Armrmobaenetes Hubbell which are of much 
smaller size, some species being rather minute. 

A key to males of this new genus, based on some of the more im- 
portant taxonomic features, is presented below. 


Key TO THE MALES OF Macrobaenetes NEW GENUS 


1. Size very large. Caudal femora with basal portion of the lower genic- 
ular lobes armed with 2 or 3, rarely 1 tooth; dorsal teeth confined to 
ridge; usually 16-18, sometimes 15-22 teeth on external inferior 
keel. Sides of mesonotum nontuberculate. . . .valgum (Strohecker ) 


Size large. Caudal femora with basal portions of lower genicular 
lobes armed with 1, rarely 2 teeth; dorsal teeth confined or not con- 
fined to dorsal ridge; 8-20 teeth on the external inferior keel. Sides 
of mesonotum tuberculate or nontuberculate ................. Y 


bo 


. Dorsal mesotibial spurs—3 pairs. Caudal femora with dorsal teeth 
confined to ridge; external inferior keel with 14-20, usually 16-18 
large closely spaced teeth. Sides of mesonotum nontuberculate 
Sica ce ESTAR oR Te FI Os ok eee He Be ene kelsoensis n. sp. 


New sand-treader camel crickets 93 


Dorsal mesotibial spurs—4 to 5 pairs. Caudal femora with dorsal 
teeth running in oblique rows exteriorly onto upper portion of outer 
pagina; external inferior keel with 9 to 18 teeth. Sides of mesonotum 
wile Glemtae TleelG Ang ie ek Ce ie. See nena ee rrene 3 


3. Usually five pairs of dorsal mesotibial spurs. Abdominal notites 
sparsely tuberculate; metanotum sparsely tuberculate. External in- 
ferior keel of caudal femora with usually 8-10, sometimes 8-13 large, 
muldelwaciacedmteetht eae ee en algodonensis n. sp. 


Usually 4 pairs of dorsal mesotibial spurs. Abdominal notites and 
metanotum with sides heavily tuberculate. External inferior keels 
of caudal femora with usually 13-14, sometimes 9 to 18 large, strong, 
lescmnndelyasmacedsteeth . 555s ser qe asec 1: slerrapintae n. sp. 


Macrobaenetes kelsoensis NEW SPECIES 
FIGuRE 1 


Differs from valgum (Strohecker ), algodonensis n. sp. and sierrapintae 
n. sp. by possessing only 3 pairs of dorsal mesotibial spurs, instead of 
4 or four and 1% or 5 pairs; from algodonesis n. sp. and sierrapintae 
n. sp. by the lack of dentate tuberculation on the flanks of the mesono- 
tum; by the confinement of the dorsal teeth to the dorsal ridge in the 
caudal femora and by the greater number of large strong teeth on the 
external inferior keel of the caudal femora; from valgum (Stroh.) by 
possessing only 1 instead of 2 or 3 teeth at the base of the inferior ge- 
nicular lobes of the caudal femora. These characters plus others in the 
spimation and spuration of the caudal tibiae offer ample means to the 
serious student for their separation. 


Description of Male: Size medium large, mesonotum with flanks non- 
tuberculate; metanotum and abdominal notites with flanks with few 
tubercles. Leg spmation as follows: forecoxal vertical keel unispinate; 
forefemora untoothed dorsally and ventrally except for a single long 
apical spine on the inner inferior lobe. Foretibiae dorsally unspined ex- 
cept for the apical calcars; ventrally with 4 pairs of spurs on the apical 
half, the outer 4 the largest, plus a larger pair of calcars. Foretarso- 
meres—3, segments 1 and 2 very short, 3 twice length of 1 and 2; dis- 
toventral angles of 1 and 2 lobate, of 3 normal. Middle legs with meso- 
coxal keel unspined; mesofemora unspined dorsally and ventrally on 
the keels with each inferior genicular lobe bearing a single spine. 
Mesotibiae dorsally with 3 pairs aciculate spurs, 1 pair in basal half, 2 
pairs in apical half, plus smaller apical calcars; ventrally with 3 ex- 


04 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


ternal spines and 2 internal spines plus calcars. Mesotarsomeres—4, 
segment 1 and 4 equal and each twice the length of 2 and 3 combined 
which are both very short; distoventral angles of 1 and 4 acute, of 2 
produced into a large, acute, spindle-shaped process with segment 3 
similarly shaped and smaller and interdigitating with it posteriorly. 
Hind legs typical. Caudal femora with dorsal teeth small and confined 
to the ridge; external inferior keel with 17-18 strong widely-spaced 
teeth plus a basal tooth or spine on the inferior genicular lobe; internal 
inferior keel with 13-15 very widely spaced much smaller teeth plus 
basal and apical teeth on the internal inferior genicular lobe. Caudal 
tibiae ventrally unspined except for the preapical spur plus the apical 
and almost attingent calcars; dorsally with 7 pairs of spurs plus apical 
calcars arranged in this fashion: first pair small, aciculate, at about the 
basal third and preceded by 2 outer and 6 internal small teeth; second 
pair spurs about the apical two-thirds, preceded by 6-7 external and 
6-8 internal larger teeth; third pair preceded by 1-2 external and 2 in- 
ternal teeth; fourth pair preceded by 0-1 and 0-0 internal teeth; the 
rest of the sand basket entire; the first 3 pairs of spurs smallest and 
semi-aciculate, the remainder long and narrowly spathulate for dig- 
ging and forming the sand basket. Caudal tarsomeres 4, first slightly 
longer than 4 which is twice the combined length of the very short 
second and third segments; distoventral angle of 1 acute, 2 large and 
spindle-shaped as typical in the genus, 3 smaller than 2 and inter- 
digitating with it posteriorly and 4 normal. 


Holotype Male: Kelso Dunes, San Bernardino County, California, elev. 
2500 feet, April 30, 1960. Ernest A. Tinkham; on bare hard packed 
sand ridges one-half mile inland from margin. Calliper measurements: 
body length 19.0; pronotum 4.8; caudal femora 14.1 x 5.8; caudal 
tibiae 12.3 mms. Holotype in the Tmkham Eremological Collection. 


Description of Female: Size medium, slightly smaller than the Holo- 
type, differing from the type in the followmg features: caudal tibiae 
straight and variations of spination as follows: forelegs with forefemora 
similar to Type; foretibiae ventrally with 4 large external and 3 in- 
ternal smaller spurs plus larger calcars; protarsomere as in Type. Mid- 
dle legs as in Type. Hind legs: caudal femora much smaller in length 
and depth with dentition much reduced: dorsal teeth of ridge much 
reduced in size and numbers and largely confined to a simgle row on 
the internal edge; dentition of lower keels greatly reduced, the external 
inferior keel hear ing 11-14 small scattered eoth plus a small basal tooth 
on the inferior genicular lobe; internal inferior keel with 12-14 mmute 


New sand-treader camel crickets 05 


and widely scattered teeth plus 1-2 small basal and 1 larger apical tooth 
on the inferior genicular lobes. Caudal tibiae straight, dentition and 
spuration less than in Holotype and as follows: external dorsals with 
5-6 minute teeth preceding the first spur at the basal third, 9-12 larger 
variable teeth preceding spur 2 at the apical third, 1-2 similar teeth 
preceding spur 3 and rest of spurs of sand basket entire; internal dorsals 
with 7-8 small teeth preceding spur 1 at the basal third, 9 variable 
larger teeth preceding spur 2 at the apical third, 1 tooth preceding spur 
3 and rest of sand basket entire. Caudal tarsomeres as in the Holotype. 


Genitalia: Supraanal plate roundly triangular; subgenital plate semi- 
circular. Ovipositor slightly less than the pronotal length; dorsal valvu- 
lae obliquely truncate with its apex armed with an acuminate spine; 
ventral valvulae with 4 uncinate hooks, the two at the extreme apex 
much the largest. 


Alloty pe Female: Same data as the type. Calliper measurements: body 
length 15.4; length to apex of ovipositor 17.8; caudal femora 11.2; 
caudal tibiae 9.8; ovipositor 3.7 mms. 


Paratype Males: 80, same data as the Holotype but collected on the fol- 
lowing nights; 1955: June 18,2 % ; 1957: June 14,1 3, June 30,2 3; 
SSE O 475 June 10) 13), July 12" 5 1959: May 9, 25 3; 
IMayeZ35 (515); 1900: April 30; 203, May 10,8 3 ; 1961: April 15, 
10 %. Range in measurements: body length 14.5-18.0; pronotum 4.2- 
4.8; caudal femora 11.8-16.0, caudal tibiae 9.2-13.0 mms. Paratypes to 
be distributed to the major orthopterological museums. 


Paratype Females: 60, same data as the Holotype but collected on the 
following nights: 1955: June 17,1 9; 1957: June 14,1 ?, June 30, 
Pen 58-7/Npril 195572 July tol); 1959: May, 952 9 May 23, 
oe 960 = April 305,209) May 148 951961: 15 2. 

All paratypes similar to their respective Types. 


Historical Ecology: Although I led a small group of young entomolo- 
gists to the Kelso Dunes for the first time on June 25-26, 1954, it was 
not until my second visit on June 17-18, 1955, that the presence of the 
giant sand-treader became known. The small sand-treader Ammo- 
baenetes n. sp., to be shortly described, is usually quite abundant in the 
soft sand of the marginal areas where Creosote, Sand Paper weed, Gal- 
leta Grass (Hilaria rigida) are common but the giant sand-treaders 


96 Bulletin So. Calif, Academy Sciences / Vol. 61, Pt. 2, 1962 


dwell on the hard packed sand ridges at least half a mile inland from 
the dune margins. On my third trip July 12-14, 1956, no Macro- 
baenetes had survived to that late date and only 2 Armmobaenetes were 
found. In the summer of 1957, I commenced my first summer of sand 
dune research under a three-year grant from the National Science 
Foundation and my dune research was greatly increased thereby. On 
June 14-16, I collected 25 “Ammos” and on June 30-July 1 took 2 
‘“Ammos” and 3 Macrobaenetes. On the sixth trip, Oct. 25-26, a fine 
drizzling rain, the only rain in 17 Kelso trips, made a strange night on 
the dunes. At this time the tiny nymphs of both genera were out and 
their burrows marked by small sand piles were much in evidence next 
morning on the sunlit dunes. Despite the winter rams of 1958 that 
brought a nice array of spring flowers out and made the margins of 
the Kelso Dunes like drifts of snow from the large white petals of the 
Dune Primrose, sand-treaders were not abundant, indicating that not 
winter rains but the late summer and early fall rains regulated the 
abundance of these creatures. Collections were made on April 19-20, 
1958, and later on when the heat of late summer had seared the plant 
life, I took 7 males and no females and 4 “Ammos” the night of June 
10. Still later, after midnight the night of July 1, I found 2 “Macros” 
and 2 “Ammos.” Visits were also made on Sept. 22-23 and Oct. 9-10 
of that year. In the spring of 1959 I commenced offermg my new 
course “Nature Study of the Desert” for the Extension Department of 
San Diego State College and began bringing large classes of teachers 
and laymen to the Kelso Dunes on an overnight study trip. Such large 
classes greatly augmented my collecting propensities but naturally de- 
stroyed the value of comparative collecting by one person. 1959 trips 
were made on May 9-10, May 23-24, October 17-18; 1960 trips were 
on April 30-May 1, May 14-15, Oct. 22-23; 1961 trips on April 15-16. 


Biology: The ova of Macrobaenetes kelsoensis ni. sp. 1s oval elliptical 
and measures 3.0 x 1.4 mms. in diameter. The exact compliment of 
ova per female is not known but is in the neighborhood of 50. Where 
the eggs are laid is not known for certain, but they are probably de- 
posited down in the stygian chambers deep in the damp sand for their 
chorion walls are thin and subject to desiccation, and the only place 
providing constant moisture for the ova would be in their chambers. 
They do not inhabit animal burrows like the much rarer Ceuthophilus 
fossor Hubbell. The advent of the late summer rains in August and 
September bring out the tiny young and by October these are about 
the size of a large “match head” although there is naturally some size 


New sand-treader camel crickets 07 


range. Their presence on the hard-packed sand ridges, half a mile in- 
land from the loose sand and vegetated margins, is conspicuously indi- 
cated by the small mounds of sand that in favorable locations resemble 
the colonies of certain terrestrial bees. With experience one can distin- 
guish between their mounds and those of scorpions, solpugids and spi- 
ders. I have taken some of these small young and by placing in small 
cartons with sand and feeding daily with a little lettuce have brought 
them through to maturity about the time those on the Kelso Dunes are 
maturing. These October nymphs are undoubtedly in their second 
stadia of their lives, there is another molt in January which brings them 
to almost half-grown size, and perhaps another molt in March before 
the final molt in April. The young dig their tunnels rather shallowly 
because the temperatures are cool during the winter months but the 
tunnels lengthen with the maturity of their occupants. The adults in 
May and June dig down at about 35-40 degrees of angulation with the 
surface and continue in a straight line downwards to pierce deeply in 
the damp sand layer which is usually down 8 or 9 inches. At the end 
of their tunnel, often two feet or more in length, they excavate a small 
chamber hardly bigger than themselves and here they rest during the 
day with their heads always pointing upwards and outwards. How 
they know, down in that blackness when evening is approaching is per- 
haps best explainable in the marvelous time mechanism possessed by 
wild creatures, but as dusk or darkness approaches, the sand-treaders 
are usually found at the mouths of their burrows waiting for darkness 
to encompass the dunes so that they can begin their purposeful peri- 
grinations. 

At night their wanderings are almost entirely in search for food 
which seems to be mainly the seeds of the Dune Grass (Orzyopsis hy- 
menoides ), bits of organic matter such as dried leaves, perhaps nibbles 
on green dune grass cut down by Kangaroo rats or other dead protein 
matter such as bits of dead insects or otherwise. No mating observations 
have ever been witnessed in nine years of study. 

The Kelso Dunes are always damp 6-8 inches down, sometimes only 
3-4 inches down, even during the driest years. It is believed the Kelso 
Dunes have their origin in subterranean or subammean springs em- 
anating from the delta region of the Mohave River some miles to the 
southwest in what is called the “Devil’s Playground” for even the high- 
est Kelso Sand Peak of 700 feet elevation and undoubtedly the highest 
in the United States, is always wet to the top. This fortuitous circum- 
stance is what creates the remarkable and rather indiginous nature of 
the biotae of the Kelso Dunes. 


98 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


Enemies: ‘The only known enemy is the Sand Dune Scorpion (Para- 
nurctonus mesaensis Stahnke). I have found complete and partial re- 
mains of these sand treaders in the scorpions’ chambers down in the 
sands. Their enemies have enemies, too, for the American Raven pur- 
posely hunts for the sites of the scorpion burrows and exhumes the 
creatures with their long heavy beaks, whack off their lethal extremi- 
ties and devour their juicy bodies. Scorpions also kill and eat their own 
kind, the larger preying on the smaller, so that the effect of scorpions 
on the sand treader population is negligible. The great factor controll- 
ing sand-treaders is heat and the advent of the torrid heat of late May 
and early June soon decimates the sand-treader population. In general 
the female sand treaders survive longer than the males because their 
bodies are packed with developing ova. 

This interesting new species of the new genus Macrobaenetes which 
is restricted to aeolian sand dunes of the Colorado Desert and its peri- 
pheral area, is named after that magnificent pile of quartz sand, whose 
high peak of 700 feet is unquestionably the highest in the United States. 


Orthopteran associates: The Kelso Dune fauna is considerable. Noc- 
turnal associates are the new Ammobaenetes, the large camel cricket 
Ceuthophilus fossor Hubbell and a new Jerusalem cricket that may 
represent a new genus as well as the following Decticids in good years 
(1.e. considerable winter precipitation), the large grey, black-winged 
Capnobotes fulginosis, the very rare Anoplodusa arizonensis, and a 
new species of Eremopedes. Diurnal associates are mostly acridids such 
as Coniana snow on the sands, Xeracris minimus im clumps of Petal- 
onyx Thurberi, Ligurotettix coquilletti on the stems of Creosote, Trim- 
erotropis p. pallidipennis an ubiquitous desert species as well as the 
mantids Litaneutria minor and Stagmomantis. Sand roaches of unde- 
termined species are also found on the dunes at night; these are being 
studied by Drs. Gurney and Friauf. 


Macrobaenetes valgum (STROHECKER) 
FIGURE 3 
Dathiniodes valgum Strohecker, 1960: 31-32, fig. 1. 
Strohecker’s decision to refer the present genus to Dathiniodes “largely 
on the basis of its tarsomeres” and not on the sum total of its char- 
acters as should be done for more accurate placement, has resulted im 
placing this creature in a genus that does not possess a “‘sand basket.” 
Thus, in choosing a single character, he has not only ignored a char- 
acter of great taxonomic importance, the “sand basket?’ but by so doing 


New sand-treader camel crickets 99 


has ignored zoogeographical concepts as well. In 1936, Hubbell defined 
the faunistics of his new genus Dathiniodes as dwelling “on high plains 
and eastern foothills of Rocky Mountains in Colorado, to southwestern 
Texas and Arizona” and I might add, as was also known to Strohecker, 
as living in the wet, cool, hard gypsum sand substrata of the White 
Sands. It is hardly conceivable or possible that a creature dwelling in 
such an environment and at a considerable elevation as defined above, 
could exist on the torrid sands of the Colorado Desert at elevations little 
above or even below that of sea level. 

Macrobaenetes valgum (Strohecker) differs from kelsoensis n. sp., 
algondonensis, n. sp. and sierrapintae n. sp. by possessing usually 2 or 
3 instead of 1 basal tooth on the inferior genicular lobes of the caudal 
femora. From kelsoensis n. sp. it is further separated by its large size 
and the dentition and spuration of the caudal femora and caudal tibiae; 
from algondonensis n. sp. and sierrapintea n. sp. by lacking tubercula- 
tion on the flanks of the mesonotum and by the greater number of teeth 
on the external inferior keel of the caudal femora and by the confine- 
ment of the dorsal teeth to the ridge in the caudal femora. 


Description of Male: Size very large and typical. Mesonotum usually 
smooth, metanotum and first five abdominal notites weakly tubercu- 
late, the tubercles confined to a sparse row on the flanks just cephalad 
of the posterior margin. Forelegs with forecoxal vertical keel unispin- 
nate; forefemora dorsally and ventrally unspined except for a slender 
spine on the internal inferior genicular lobe; foretibiae dorsally un- 
spined except for the calcars, ventrally with 4 pairs of spurs, outer row 
the largest, on the apical two-thirds, plus a pair of larger calcars. Pro- 
tarsomeres—3, segments 1 and 2 very short, segment 3 twice combined 
length of 1 and 2; distoventral angles of 1 and 2 acutely lobate, of 3 
acute. Middle legs; mesocoxal ridge unspined; mesofemora dorsally 
unspined, ventrally with a small spine arisimg from each of the in- 
ferior genicular lobes; mesotibiae with 4, sometimes 3 or 3 and 1, 
dorsal pairs of spurs plus large calcars, ventrally with 3 external and 2 
internal spurs plus larger calcars. Mesotarsomeres—4, segments 1 and 
4 equal and each twice the combined length of the very short second 
and third segments, segment 2 large and acutely spindle-shaped with 
the smaller segment 3 interdigitating posteriorly with it; distoventral 
angle of 1 and 4 acute, of 2 and 3 acutely lobate. Hind legs: caudal 
femora with dorsal teeth confined to ridge; external inferior keel with 
usually 16-18, sometimes 15-22, large strong, uniform, mostly evenly 
spaced teeth; internal inferior keel with 12-17 much smaller, much 


Se 


100 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


more widely spaced teeth; both lower genicular lobes with 2 or 3 basal 
teeth or spines. Caudal tibiae typically arched with 7 large or very 
large pairs of spurs plus an apical pair of very large calcars located as 
follows: first pair smallest, aciculate, located about the basal third and 
preceded by 2-7 external and 5-8 internal small teeth; second pair of 
spurs about the apical two-thirds, shghtly larger in size and preceded 
by 5-6 external and 7-8 internal larger teeth; 3rd pair larger still and 
preceded by 1-3 external, 1 internal teeth; rest of sand basket entire. 
Subgenital plate with forcipate arms. 


Males, 100, collected as follows: 4 miles west of Indio: 1952: 10 %. 
April 26; 1955: 4 6, April 10,5 @ , April 26, 3 3, May, O-eIOa7el0 
3, May 18; 1958: 1 § mymph, Feb. 21, 11 8, April 12,5 §, May 
3, 6. 3 miles west of 1000 Palms: 1958: 8 %, May 33 195925353, - 
May 5, 4 8, May 16,2 3, May 19,1 3, May 21; 1960: 18 &, April. 
Palm Springs Depot: 1952: 12 4, May 1. All collected by E. R. ‘Tink- 
ham. 


Description Female: Size medium large but smaller than the larger 
male. Form typical for Macrobaenetes females and characterized chief- 
ly by leg spination as follows: Forelegs with vertical procoxal keel uni- 
spimate; forefemora unarmed dorsally and ventrally with a spine 
emanating centrally from the internal inferior genicular lobe; fore- 
tibiae dorsally unspined with apical calcars; ventrally with 3 internal, 
4 external aciculate spurs in apical half plus pair of larger calcars, the 
external spurs much the largest and forming the tines of a fossorial 
basket. Protarsomeres—3, the first two segments very short, segment 3 
twice their combined length; distoventral angle of segments 1 and 2 
acutely lobate, of 3 normal, the segment bristling with short golden 
hairs. Middle legs with mesofemora dorsally unarmed and ventrally 
both keels unarmed but with both lower genicular lobes unispinate with 
a central tooth. Mesotibiae dorsally with 3 and 4% to 4 pairs of acicu- 
late spurs, 1 and 1% to 2 in basal half, 2 pairs apical half plus terminal 
calcars; ventrally with 2 pairs spurs at middle and apical third plus 
terminal calcars. Mesotarsomeres—4, the first and fourth segments 
equal and each twice combined length of the very short second and 
third; distoventral angles of 1 acute, of 2 and 3 acutely lobate of which 
2 is larger, of 4 almost normal. Hind leg with caudal femora with 
dorsal teeth of ridge minute and very sparse and chiefly indicated as a 
row on the inner edge; external inferior keel with 12-14 minute widely 
scattered teeth with 1-2 variable small to very small basal teeth on the 
inferior genicular lobe; internal inferior keel with 8-10 similar teeth 


New sand-treader camel crickets 101 


in basal half and with basal and apical tooth on the inferior genicular 
lobe. Caudal tibiae with 7 pairs of dorsal spines plus apical calcars, the 
first two pairs smaller and aciculate, the remainder larger and nar- 
rowly spathulate and arranged as follows: first pair at the basal quarter 
and preceded by 4-5 external and 6 internal minute teeth; 2nd pair 
just distad of center and preceded by 7-8 external and 4-5 internal 
larger teeth, 3rd pair about the apical third and preceded by 2 external 
and 1-2 internal and similar teeth; rest of sand basket entire and closely 
arranged to form the tines of the sand basket. Metatarsomeres—4, first 
segment larger than 4th which is twice the combined length of the very 
short 2 and 3 segments; their distoventral angles as in the mesotarso- 
meres. Ovipositor shorter than the pronotum, very obliquely truncate 
with terminal tooth; ventral valvulae with 4: teeth, the 2 apical un- 
cinate, the 2 basal mere serrations. 


Hypoallotype Female: 3 miles west of 1000 Palms, April 14, 1960, 
Ernest A. Tinkham; on the wind-swept drift sand in sand dune valley. 
Measurements: body length 19.8; length to apex of ovipositor 22.8; 
caudal femur 12.4; caudal tibia 10.0; ovipositor 4.2 mms. Hypoallo- 
type in the Tinkham Collection. A Hypoallotype is one described by a 
person other than the describer of the species. 


Hy poparaty pes: 149, with following data: 4 miles west of Indio: 1952: 
iow pile ©) 1955: 5) 9)Aprl 1053/9 April 26, 3.9 May 9; 1957: 
13 9 May 18; 1958: 2 9 nymphs Feb. 21,12 9 April12,1 9 May 
3. 3 miles west 1000 Palms: 1958: 10 9 May 3; 1959: 6 9 May 5, 
ieee Mayetion oo) May, 19%'5)'o May 2111960: 20.0) April id 1 
mile N. Palm Desert: 1959; 15 9 May 21. Palm Desert Depot: 1952: 
10 @ May 1. All collected by E. R. Tinkham. 3 miles W. 1000 Palms, 
20 2 April 14, 1960, Jacques Helfer. 

Hypoparatypes similar to the Hypoallotype. Range in measure- 
ments: body length 14,3-21.9; pronotum 3.5-5.0; caudal femora 11.0- 
13.8; caudal tibiae 8.8-10.8; ovipositor 4.4-4.0 mms. Hypoparatypes to 
be distributed to various museums. 


Distribution: M. valgum is found from the sand dunes just south of the 
Palm Springs Depot, 9 miles west of Palm Springs, California, east 
along the wind swept sand dune ridges to 2 miles west of Indio. The 
species apparently does not cross south of Highway 111, nor does it ex- 
tend as far east as where the Dillon Road joins Highway 66. 


Ecology: This species is largely regulated in its abundance by the win- 
ter rains which regulate the spring flowers of Coachella Valley. There 


102 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


are some spots favorable to permanent habitation where springs keep 
the sands damp, but in very dry years the species disappears over most 
of the sand areas. : 


Biology: No observations are available for the early stages in the fall. 
The Feb. 21, 1958, collection record is the earliest and indicates the 
species is more than half matured at this time. Temperature readings at 
this time were at 9:25 p.m. with the sand damp from a recent rain as 
follows: sand surface 57° and air temperature 62° Fahrenheit. On May 
3, 1958, at 3 miles west of 1000 Palms many burrows were noted. 
These went shallowly under the sand surface at about 2 inches depth 
and parallel to the surface for about one foot then aneled downward 
at about 40° to terminate in the damp sand substratum. Here the female 
was found resting facing outwards and upwards. On this date many 
males were found lying dead on the sands, due to lack of food, and 
although females were fairly common at this date, males were rare. 
The Sand Dune Scorpion sometimes prey on these sand treaders. 


Orthopteran Associates: Nocturnal orthopteran associates mclude the 
rare camel cricket, Cewthophilus fossor, the much rarer Stenopzlmatus 
fuscus found in wet springs around Cathedral City and the common 
sand roaches which are being studied by Drs. Friauf and Gurney. 
Diurnal associates include Conana snowi on the plant Coldenia plicata. 
Xeracris minimus on Petalonyx thurberi, Ligurotettix coquilletti on 
Creosote stems, the common Trimerotropis p. pallidipennis, Cibolacris 
parviceps aridus and Derotmema delicatula, the latter rare. 


Macrobaenetes algodonensis NEW SPECIES 
FIGURE 4 


M. algodonensis n. sp. is differentiated from kelscensis n. sp. by pos- 
sessing 5 instead of 3 pairs of dorsal mesotibial spurs, by the extension 
exterioradly of the dorsal teeth on the ridge of the caudal femora and 
by fewer teeth, quite widely spaced, on the external inferior keel of the 
caudal femora; from valgum (Strohecker) by possessing 1 instead of 2 
or 3 teeth on the basal portions of the lower genicular lobe of the caudal 
femora, the fewer teeth on the external inferior keel of the caudal fem- 
ora and by 5 instead of 3 or 4 pairs of dorsal mesotibial spurs; from 
sierrapintae n. sp. by the weaker tuberculation on the flanks of the 
metanotum and first five abdominal notites and by fewer and more 
widely spaced teeth on the external inferior keel of the caudal femora. 


New sand-treader camel crickets 103 


Description of Male: Size medium large, form typical. Flanks of the 
mesonotum bare, metanotum with less than 10 dentate tubercles, first 
four abdominal notites with a few tubercles in a single row posteriorly. 
Leg spination as follows: Forelegs with vertical keel of forecoxae uni- 
spimate; forefemora unspimmed dorsally and ventrally with the inferior 
genicular lobes each bearing a single spine or tooth, the internal one 
the larger; foretibiae unspined dorsally plus the apical calcars, ven- 
trally with 3 pairs of spurs in the apical half plus the calcars. Protarso- 
meres—3, first 2 segments very short, third about 2 and 1% times 1 and 
2; distoventral angles of 1 and 2 lobate, of 3 normal. Middle legs; 
mesocoxae unspined; mesofemora unspined dorsally, ventrally as in 
the profemora with an additional small tooth on the apical external 
keel; mesotibiae dorsally with 5 pairs of long aciculate spurs plus sim- 
ilar calcars, ventrally 2 pairs of spurs, one median, one apical plus 
apical calcars. Mesotarsomeres—4, first slightly longer than the fourth, 
the fourth about twice the combined length of the very short 2 and 3 
segments; distoventral angles somewhat acute in 1 and 4, greatly pro- 
duced in 2 and smaller in 3 which interdigitates posteriorly with it. 
Hind legs typical; caudal femora with dorsal teeth mostly confined to 
the dorsal ridge, some extending exterioradly in oblique rows onto the 
upper edge of the outer pagina; external inferior keel with 9-10 mostly 
large, widely spaced teeth plus a large tooth on the basal portion of the 
lower genicular lobe; internal inferior keel with 6-11 very widely 
spaced, minute teeth plus a basal tooth on the inferior genicular lobe. 
Caudal tibiae with 8 pairs of spurs including the calcars cf which the 
five apical pair plus the calcars are very long and narrowly spathulate 
to form the sand basket, the remaining three pairs aciculate and ar- 
ranged in this fashion: first pair small at the basal third preceded by 7 
small teeth on each keel, 2nd pair at the apical two-thirds and pre- 
ceded by 12-13 external and 8-10 internal similar teeth, third pair and 
first pair of the “sand basket” preceded by 1-2 external and 2-3 internal 
teeth, fourth pair preceded by 1 external and 1 internal tooth, rest of 
the sand basket without teeth between them. 


Holotype Male: Algodones Dunes, 15 miles NW of Yuma in California 
and 14 mi. E bridge on All American Canal, Apr. 25, 1958. Ernest R. 
Tinkham. Caliper Measurements: body length 11.8; pronotum 3.9; 
caudal femora 14.9; caudal tibiae 12.2 mms. for dried alcoholic ma- 
terial. Type in the Tmkham Eremological Collection. 


Description of Female: Size similar to Holotype from which it differs 
chiefly in spination of legs and genitalia. Forelegs with forefemora un- 


104. Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


spined dorsally and ventrally with the internal inferior genicular bear- 
ing an apical spine, remainder as in Holotype. Middle legs as in Holo- 
type. Hind legs with caudal femora shorter and less deep than in Holo- 
type, the dentition much reduced and as follows: dorsal teeth on ridge 
of caudal femora minute and largely confined to a row on the internal 
edge; external inferior keel with 4 medium-sized teeth widely sep- 
arated on the middle portions of the keel plus a basal tooth on the lower 
genicular lobe; internal inferior keel unarmed with basal and apical 
small teeth on the lower genicular lobe. Caudal tibiae dorsally with 8 
pairs of spurs characterized as follows: externally first spur at basal 
quarter preceded by 6 small teeth, second spur just distad of center and 
preceded by 10-11 larger teeth; third spur just distad of the apical 
third and preceded by 1-2 similar teeth; rest of spurs entire and form- 
ing the outer tines of the sand basket; internal dorsal keel with first spur 
at basal quarter preceded by 9-10 small teeth, second spur situated in 
the middle and preceded by 6-7 larger teeth, spur three at about apical 
third and preceded by 4 similar teeth and rest of sand basket entire. 
Caudal tarsomeres as in Holotype. Ovipositor typical of the genus. 
shorter than the pronotum; the dorsal valvulae obliquely truncate and 
terminating in a spine; ventral valvulae with 4 apical uncinate hooks, 
the two at extreme tip conspicuous, the two basal minute. 


Allotype Female: Same data as the Holotype. Calliper measuremenis: 
body length 14.3; pronotum 3.9; caudal femora 11.0; caudal tibiae 
9.8; ovipositor 2.7. Allotype in the Tinkham Collection. 


Paratype Males: 115 with data as follows: 19, April 28, 1952, same 
location as Holotype; 13, April 25, 1958, same location as Holotype; 
62, April 29, 1961, 2 miles west of Glamis, Calif.; 10, May 13, 1961, 
same location; 8, 4 miles west Glamis, April 20, 1961S emerS;, 
1960, 4 mis. W. Ogilby, Calif., Roth and Irwin. Range in measure- 
ment as follows for series: 1952 series, body length 11.2-15.9; pro- 
notum 3.0-4.2; caudal femora 10.7-15.0; caudal tibiae 8.8-13.5 mms. 
2 miles W. Glamis series: body length 10.0-15.3; pronotum 3.0-3.9; 
caudal femora 10.6-13.2; caudal tibiae 8.9-11.2 mms, 4 mis. W. Glamis 
series: body length 16.5-19.2; pronotum 4.2-4.6; caudal femora 14.6- 
17.0; caudal abiae 12.5-14.5 mms. Ogilby series: body length 10.5- 
13.4; pronotum 2.7-3.0; caudal femora 11.0-10.9; caudal tibiae 9.1-9.2 
mms. 

Paratypes closely similar to the Holotype with slight variation in the 
dentition of the caudal tibiae but within the species range. 


New Sand-treader camel crickets 105 


Paratype Females: 140 with data as follows: 9, April 28, 1952, same 
data as Allotype; 10, April 25, 1958, same location as Allotype; 54, 
April 29, 1961, 2 mis. W. Glamis; 41, May 13, 1961, 2 mis. W. Glamis; 
26, 4 mis. W. Ogilby, June 5, 1960, Roth and Irwin. Range in measure- 
ments for series as follows: 1952 series: body length, 13.2-22.8; pro- 
notum 3.2-4.2; caudal femora 9.2-13.0; caudal tibiae 7.8-10.0; ovi- 
positor 2.4-4.1 mms. 2 mis. W. Glamis series: body length 11.0-15.0; 
pronotum 3.4-3.5; caudal femora 10.7-11.2; caudal tibiae 8.8-9.0; ovi- 
positor 2.8-2.6 mms. Ogilby series: body length 12.8-17.0; pronotum 
3.1-3.8; caudal femora 9.7-11.2; caudal tibiae 7.4-9.3; ovipositor 2.2- 
3.8 mms. 

Female Paratypes, closely similar to the Allotype but with consid- 
erable size variation depending upon the aridity of the season. 

Male and Female Paratypes to be deposited in the major museums 
and the Tinkham Collection. 


Ecology: The dune fauna in recent years with little precipitation is 
much less than in such springs as 1953, when on the night of April 28, 
the cold dunes, wet with a late evening cold rain, were covered with 
dune creatures such as sand treaders, scorpions, solpugids, tenebrionid 
beetles of several genera and species as well as geckos, all in consider- 
able profusion and immobile on the cold sands. Never before or since 
have I witnessed such an array of life. 

The 1958 and 1960, and 1961 series of sand-treaders average much 
smaller than the 1953 series. It was interesting in 1961 to find sand- 
treaders at 4 miles west of Glamis (on the west edge of the dunes) to 
be much larger than those 2 saute west of Glamis along the main ridge 
of the Algodones. 


Biology: Very little is known oe healy about the life history of this 
new species but generally its life history is similar to that of M. kel- 
soensis n. sp. and the other species. Some adults persist through until 
mid-June, rarely to late June. 

Orthopteran Associates: Nocturnal associates include sand roaches, 
being studied and the large camel cricket Ceuthophilus fossor; diurnal 
orthopteran associates include such acridids as Xeracris minimus, 
Coniana snowi and the everpresent Trimerotropis p. pallidipennis. 


Macrobaenetes sierrapintae NEW SPECIES 
FIGURE 2 


Differs from kelsoensis n. sp .and valgum (Strohecker) by the heavy 
tuberculation on the sides of the meso-, metanotum and first five ab- 


106 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


y 


y/, 


del £ AR Fizkha tx 


Figure 1. Macrobaenetes kelsoensis n. g. et n. sp. Left hind leg of Holotype. 
Kelso Dunes, San Bernardino County, California. 


Figure 2. Macrobaenetes sierrapintae n. sp. Left hind leg of Holotype. Sierra 
Pinta Dunes, 65 miles south of Mexicali, Baja California, Mexico. 


New sand-treader camel crickets 107 


y / CAI, EIR. Med A, 
Figure 3. Macrobaenetes valgum (Strohecker). Left hind leg of male from dunes 
3 miles west of 1000 Palms, Coachella Valley, Riverside County, California. 


Figure 4. Macrobaenetes algodonensis n. sp. Left hind leg of Holotype Male. 
Algodones Dunes, 15 miles NW of Yuma, Imperial County, California. 


108 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


dominal notites; by the larger dorsal teeth of the caudal femora ex- 
tending obliquely in rows onto the upper portions of the outer pagina; 
by the dentition on the external inferior keel of the caudal femora and 
the spination and dentition on the caudal tibiae. From Kelsoensis it is 
further distinguished by possessing 4 or more pairs, instead of three, 
dorsal mesotibial spurs and from valgus by having only one instead of 
two or three teeth on the basal portions of both external and internal 
lower genicular lobes of the caudal femora. From algodonensis n. sp. 
it is separated by the greater dentition on the external inferior keel of 
the caudal femora; the dentition and spuration of the caudal tibiae as 
well as by the heavy tuberculation on the flanks of the metanotum and 
first five abdominal notites which in algodonensis is sparse. 


Description of Male: Size large, flanks of pronotum bare, mesonotum 
posteriorly with few scattered tubercles; metanotum and first five ab- 
dominal notites heavily dentate tuberculate. Leg spination as follows: 
Forelegs with vertical procoxal ridge unispinate; profemora unspined 
dorsally and ventrally with both lower genicular lobes unispinate, the 
internal one larger. Protibiae dorsally with 2 internal, 1 external spur 
basally plus apical calcars; ventrally with 3 pairs of aciculate spurs in 
apical half plus a pair of larger similar calcars. Protarsomeres—3, first 2 
segments very short, the third twice the combined length of 1 and 2; 
distoventral angles of 1 and 2 lobate, cf three normal. Middle legs with 
mesocoxal ridge unspined; mesofemora as in the profemora. Meso- 
tibiae dorsally with 4 and 1% pairs of aciculate spurs, 2 pair basal half, 
2 pair apical half and one intermediate spur on the external keel; ven- 
trally as in the protibiae. Mesotarsomeres—4, first and 4th segments 
about equal and each twice the combined length of the very short sec- 
ond and third segments; distoventral angle of 1 lobate, of 2 produced 
into a long acute spindle-shaped process with segment 3 similar but 
smaller and interdigitating with it posteriorly, and segment 4 normal. 
Hind legs with caudal femora very deep, the external inferior keel 
gently arcuate; dorsal ridge bearing obliquely running rows of short 
semi-appressed teeth extending exterioradly onto the upper portions of 
the face of the outer pagina especially in the apical third; imferior 
genicular lobes each with a basal tooth, the mner lobe also with an 
apical tooth; external inferior keels with 13 to 14 mostly large, strong, 
well spaced teeth; internal inferior keel with 12-13 much scattered and 
much reduced teeth. Caudal tibiae strongly arched with 8 external 
and 8 internal dorsal spurs of which 6 pairs in the apical region are 
long, narrowly spathulate spurs and form the sand basket, the rest are 


New sand-treader camel crickets 109 


smaller and more aciculate, the basal 2 pairs located as follows: first 
pair about the basal third and preceded by 4-5 external and 4-7 in- 
ternal small teeth; second pair about the %rds position and preceded 
by 7-9 external and 5-7 internal larger teeth; the third pair of spurs 
preceded by 3 external and 2-3 internal similar teeth; pair 4 by 0-1 
external and 1-0 internal teeth, the remainder of the sand basket entire. 
Caudotarsomeres—4, segment 1 longest and longer than the first meso- 
tarsomere, segments 2 and 3 very short, segment 4 about twice the 
combined length of 2 and 3; lobation of the distoventral angle as de- 
scribed in the mesotarsomeres. 


Holotype Male: Sierra Pinta Dunes, 65 miles south of Mexicali on 
Highway No. 5, Baja California, Mexico, April 3, 1958, Ernest R. Tink- 
ham. Calhper measurements: body length 14.8; pronotum 4.4; caudal 
femur 15.0 x 5.8; caudal tibiae 12.1 mms. Holotype in the Tinkham 
Eremological Collection. 


Description of Female: Closely similar in size to Holotype but differing 
chiefly in dentition of hind legs and genitalia. Leg spination as follows: 
Fore and middle legs as in Holotype. Hind legs with caudal femora 
less massive and caudal tibiae straight. Caudal femora with minute 
dorsal teeth and largely confined to a row on the internal keel; external 
keel with 14-15 minute teeth plus a larger basal tooth on the inferior 
genicular lobe; internal inferior keel unarmed, the lower genicular lobe 
with 1-2 basal and 1 apical larger teeth. Caudal tibiae with only 7 well- 
arranged pairs of spurs plus apical calcars, the external dorsal keel with 
the basal pair situated at the basal quarter and preceded by 5-6 small 
teeth, spur 2 just basad of the apical third and preceded by 10-11 larger 
teeth, spur 3 at the apical quarter and preceded by 1 tooth, the rest of 
the sand basket spurs entire or lacking teeth between them; internal 
dorsal keel with all spurs opposite and matching those on the external 
keel, spur 1 preceded by 6 small teeth, spur 2 preceded by 10-12 larger 
teeth, spur 3 preceded by 1 similar tooth and rest of sand basket entire. 
Caudal tarsomeres as in Holotype. Ovipositor shorter than the pro- 
notum, the dorsal valvulae squarely and obliquely truncate with ter- 
minal spine; ventral valvulae with 5 teeth of which the 2 apical are 
uncinate, the median small and the two basal mere serrations. 


Allotype Female: Same data as the Holotype. Measurements as fol- 
lows: body length 13.8; length to tip ovipositor 16.8; pronotum 4.7; 
caudal femur 13.3 x 4.5; caudal tibia 11.6; ovipositor 4.0 mms. Allo- 
type in the Tinkham Collection. 


110 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


Paratype Males: 82, same location as Holotype but collected as follows: 
10 8, April 11, 1953; 29 ¢, April 3, 1958; 18 %, May 13, 19615 all 
E. R. Tinkham. 25 ¢, April 15, 1960, Pearson, San Diego State College. 
Paratypes closely similar to the Holotype with slight deviation in denti- 
tion of caudal tibia falling within the species range. 


Paratype Females: 104; same locality as Holotype but collected as fol- 
lows: 31 9, April 11, 1953; 26 2, April 3, 1958; 22 9 , May 13, 1961, 
all E. R. Tmkham, 25 9, April 15, 1960 (Pearson; Michigan Cln). 


Paratypes closely similar to Allotype. 
Paratypes to be deposited in the major Museums and Tinkham Cln. 


Ecology: With no meteorological stations or records available from 
northeastern Baja California, it is difficult to determine good or bad 
years as far as precipitation is concerned, but usually the west coastal 
plain of the Gulf of California is exceedingly dry and drier than even 
the Californian portion of the Colorado Desert. In 1953, the species was 
fairly common in the swales lying between the bare sand ridges, these 
swales being flat bottomed and containing semistabilized hummocks of 
Atriplex polycarpa and Creosote (Larrea divaricata). On April 3, 
1958, in the same swale I counted 25 specimens around one large hum- 
mock of approximately 10 x 15 feet dimensions. In 1961, the sand 
ridges bordering this swale had grown considerably in height but the 
swale bottom was unchanged. Despite a winter with practically no 
precipitation in the northern Californian portion of the Colorado Des- 
ert, Macrobaenetes sierrapintae was not rare, although there were 
rumors of a fall storm in 1960 crossing the area but not verified. 


Biology: Nothing in particular is known about the more intimate life 
history of this species. The sands are much drier than the Algodones or 
Kelso Dunes and yet this distinct and attractive species survives under 
much more xeric conditions. 


Orthopteran Associates: These are very few. The only other nocturnal 
orthopteran is a large species of Ceuthophilus of the fossor group and 
sand roaches which are being studies by Dr. Gurney and Dr. Friauf. 
The only diurnal orthopteran is the acridid Anconia integra which 
feeds on Atriplex. 


LITERATURE CITED 


HUBBELL, THEODORE H. 
1936. A monographic revision of the genus Ceuthophilus. Univ. Fla. Publ., 2 (1): 
1-551, numerous figures. 


New sand-treader camel crickets all 


STROHECHER, H. FE 

1960. Several new species of North American Orthoptera. Pan Pac. Ent., 36 (1): 
31-35, 5 text figs. 

TINKHAM, ERNEST R. 


1947. New Species, records and faunistic notes concerning Orthoptera in Arizona. 
Amer. Midland Nat., 38 (1): 127-149, 4 plates with many figs. 


A NEW MEGAAHIPPUS FROM THE BARSTOW FORMATION 
SAN BERNARDINO COUNTY, CALIFORNIA 
Ricuarp H. TEDFoRD 
University of California, Riverside 
and 
RaymMonp M. ALF 
Webb Schoo] of California, Claremont 


INTRODUCTION 


The fossil beds of the Barstow syncline have been worked by amateur 
and professional paleontologists since 1911. It was the good fortune of 
the Peccary Society of the Webb School of California in 1955 to find 
an upper molar of an anchitherine horse and later in 1957 to find a 
palate of the same type of animal. 

These discoveries represent the first record of the genus Megahippus 
from the Pacific coast and the first reported from deposits of Barstovian 
age. 

We would like to thank Dr. M. C. McKenna of the American Mu- 
seum of Natural History, and Dr. T: Downs of the Los Angeles County 
Museum for critically reviewing the manuscript. The holotype was 
carefully prepared by Mr. Sanford Gifford of Claremont, and the line- 
drawing restoration painstakingly rendered by Mr. Robert Fakundiny 
of Riverside. 

The abbrevation UCMP refers to material in the University of Cali- 
fornia, Museum of Paleontology collections; UCR to collections at the 
University of California, Riverside; and RAM to material in the Ray- 
mond Alf Museum, Webb School of California, Claremont. 

All measurements are in millimeters. 


SYSTEMATIC DESCRIPTIONS 
Class MAMMALIA 
Order PERISSODACTYLA 
Family Equidae Gray, 1821 
Subfamily Anchitheriinae Osborn, 1910 
Genus Megahippus McGrew, 1938 
(Genotype: Hypohippus matthewi Barbour, 1914) 


Megahippus mckennai NEW SPECIES 
Figures 1 AND 2 
Etymology: Named in honor of Dr. Malcolm C. McKenna of the Amer- 
ican Museum, who has been a constant inspiration in encouraging our 
paleontological work. 


113 


114, Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


Holotype: Palate with shattered, but complete, well worn cheek tooth 
dentition, left I1/, right 12/, and both canines, RAM 6500. 


Type locality: Barstow syncline, UCR locality 3695, western side of 
“Fuller’s Earth Canyon” just north of major north bend approximately 
in southwest corner of N.W. 14, sec. 14, T. 11 N., R. 2 W,, Opal Moun- 
tam Quadrangle, 1:62,500, U.S.G.S. Edition 1955; 10,900 ft. N. 24° 
W. of road intersection 2825: Brown sandstones overlying grey silt- 
stones. Megahippus palate from the base of the sandstones, Strati- 
graphic position near the base of the upper member of Barstow forma- 
tion (Lewis, 1960). 


Referred specimens: Well worn left M1/ or M2/, UCR 3696-1, lack- 
ing both the labial and lingual borders, UCR locality 3696, outcrops on 
south side of badland amphitheatre north of Fossil Canyon, approxi- 
mately in center of N.W. 14, sec. 15, 1.11 N., R. 2 W, Opal Mountain 
Quadrangle, 1:62,500 U.S.G.S. Edition 1955, 15,300 ft. N. 35° W. of 
road intersection 2825: Greenish and tan siltstones and sandstones near 
top of the upper member of the Barstow formation (Lewis, 1960). 

Left lower jaw fragment with M/1 and M/2, UCMP 21215, fig- 
ured by Merriam (1919, fig. 28) from UCMP locality 2060 north- 
western exposures of the Barstow formation in the north limb of the 
Barstow syncline at a stratigraphic position roughly equivalent to the 
holotype of M. mckennai. 

Merriam (1919:473) also mentions a fragmentary upper molar, 
UCMP 21214 of “a large species of Hypohippus” from the Barstow 
syncline, UCMP locality 2056, Barstow formation stratigraphically 
slightly below the occurrence of the holotype of M. mckennai. This 
specimen was not figured or further described and it is only provision- 
ally referred here. 

Fragmentary limb bones of a large anchitherme are known from 
several horizons in the upper part of the Barstow formation of the Bar- 
stow syncline. Some of these are described by Merriam (1910). They 
compare closely in size and general morphology with the sample of 
Hy pohippus nr. affinis described by Henshaw from ‘Tonopah, Nevada 
(Henshaw, 1942). 


Age: late Barstovian. 


Diagnosis: Approximately 25 percent smaller than WM. matthew1; |mn- 
gual cingula on premolars not as well developed as in M. matthew; 
mesostyles with sharper crests than in M. matthew. 


New Megahippus from California 13s 


DEscRIPTION AND COMPARISONS 


Palate: Only the sides of the skull anterior to the orbits and the ventral 
rim of the right orbit are present in the holotype. The Barstow species 
was compared with figures of Hypohippus osborni (in Osborn, 1918) 
and 7. affinis (in McGrew, 1938) and directly with specimens of the 
Tonopah H. nr. affinis. 

The premaxilla is short and narrow with a pronounced constriction 
in front of the P1/ alveolus. The I3/-C diastema is shorter than in H. 
affinis and H. nr.affinis and about the length in H. osborni. The C-P1/ 
diastema is similar in length to that in H. nr. affinis, but much shorter 
than in H. affinis or H. osborni. 

The infraorbital foramen les in the ventral margin of the facial 
fossa above the parastyle of P4/. Part of the deep facial fossa is visible 
on the right side. It extends posteriorly from a point just above and 
behind the infra-orbital foramen as a deep pocket into the anterior end 
of the jugal and adjacent maxillary bones. The portion of the facial 
fossa visible is more like that in Archaeohippus (both A. ultimus and a 
specimen referred to A. mourningi from the Punchbowl formation, 
Cajon Pass, California) than described in H. affinis and H. osborni. 
Hypohippus ur. affinis, however, shows a deep malar pocket as in 
Megahippus mckennai. 

Crushing and distortion of the palate makes it difficult to discern the 
true posterior border of the palatal portion of the palatine bone. It 
seems to lie opposite the hypocone of M1/. Likewise, the exact position 
of the posterior palatine foramen is in some doubt although the groove 
which extends from this foramen anteriorly along the palatal surface 
can be clearly seen opposite M1/ on the right side of the holotype. 


Dentition: Incisors. The 11/ is the largest incisor with a massive, high 
crown and prominent lingual cingulum. A deep median pit is present 
on the occlusal surface which apparently opened laterally in early 
wear. The I2/ is decidedly smaller than I1/, but like the latter it has 
a massive high crown and large labially opening median pit on the oc- 
clusal surface. This tooth lacks a lingual cmgulum at the stage of wear 
of the holotype. The I3/ is unknown, but its alveolus indicates that it 
was smaller than I2/. 

The large and high crowned incisors are unlike those of any species 
of Hypohippus. They agree with the large and procumbent lower in- 
cisors held diagnostic of Megahippus and thus constitute one of the 
principal reasons for assignment of the Barstow form to that genus. 


116 Bulletin So. Calif, Academy Sciences / Vol. 61, Pt. 2, 1962 


Figure 1. Megahippus mckennai, new species, palatal view of holotype, RAM 
6500, approximately X0.5. Specimen whitened with ammonium chloride before 
photographing. : 


New Megahippus from California 


Figure 2. Megahippus mckennai, new species, incisors and left cheek tooth series 
in occlusal and lateral view, holotype, RAM 6500, X0.5. Teeth restored. 


1 
' 
‘ 
' 
‘ 
i] 
‘ 
' 
' 
' 


17, 


118 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


Canines. Large teeth with recurved compressed conical crowns bear- 
ing lmgual cingula. 

Premolars. P1/ is a birooted or fused birooted tooth. The roots appear 
relatively larger than in M. matthewi indicating that this tooth may 
not have been as reduced as in the latter species. The P1/ is ovoid in 
occlusal outline, flattened lingually, and bears low anterolabial and 
well developed lmgual cingula. The longitudinal valley branches pos- 
teriorly at this state of wear. There is a low anterolabial cinglum on 
the P1/. 

The premolars increase in size posteriorly with P4/ being the largest 
molariform tooth. As in MW. matthewi P2/ and P3/ have continuous 
Imgual cingula, somewhat better developed on P2/ than on P3/. P4/ 
lacks a cingulum on the lingual faces of the protocone and hypocone. 
There is a cingulum on the anterior face of the protocone in all the 
premolars. An homologous cingulum is conspicuously better developed 
on the molars. This cmgulum is as well developed on the molars as on 
the premolars in Hypohippus. The protocone P2/ is nearly circular 
in occlusal outline at the stage of wear of the holotype; it has a flattened 
anterior wall in P3/ and P4/. The protoconule is poorly differentiated. 
The hypocone is smaller than the protocone except in P2/ and its pos- 
terior wall is flattened. A crochet is present on the right and left P3/ 
and left P4/. There is a trace of this fold (angulation of the metaloph 
border at the proper position) on P2/ and the right P4/. This is prob- 
ably a very variable feature, but its frequency in the holotype suggests 
that the presence of the crochet on some of the teeth may be a constant 
feature. This fold is far more variable in Hy pohippus and usually pres- 
ent only as a spur of enamel, not a dentine filled fold as in Megahippus 
mckenna. The premolar mesostyles are sharper crests than in M, 
matthewi, more as in H. nr. affinis from Tonopah. 

Molars. The M1/ is the largest molar, the molars gradually decrease 
in size posteriorly. The M1/ and M2/ are rectangular mm occlusal out- 
line; the M3/ is wider anteriorly than posteriorly. In morphology the 
molars are much like the premolars. They differ in having weaker 
parastyles, stronger cingula anterior to the protocones, and in lacking 
continuous lingual cingula (continuous across the protocone but not 
the hypocone in M3/). The crochet is strong on M3/. In that tooth it 
blocks the floor of the protoconal valley. A crochet is present on M2/ of 
the holotype and represented on the M1/ by a thickening of the enamel 
(left M1/) or angulation of the metaloph border (right M1/). The re- 
ferred M1/ or M2/, UCR 3693-1, bears a well developed crochet. 
Enough of the base of the hypostyle remains on M2/ to indicate that it 


New Megahippus from California 119 


was a crescent-shaped cusp connected by a ridge to the hypocone. The 
ends of the crescentic hypostyle join the posterior cingulum enclosing 
an enamel-lined pit which is still visible at this stage of wear. The hypo- 
cone and hypostyle on M3/ are reduced over the condition in the an- 
terior molars. The hypostyle is represented by a ridge which connects 
the hypocone with the posterior cmgulum. 


Discussion: The new species is clearly a member of the genus Mega- 
hippus as defined by McGrew (1938:315). It differs from the Claren- 
donian M. matthewi principally in being smaller sized. The lesser de- 
velopment of internal cingula on the premolars and sharper mesostyles 
in the Barstovian form may or may not serve as valid differences when 
both species are better known. The P1/ may be slightly better devel- 
oped in the Barstow species. It seems clear from the close relationships 
of these animals that they very likely represent two members of a single 
phyletic line. If true, the dominant morphological trend in that phylum 
was simple size increase from the Barstovian to Clarendonian. 

The discovery of the Barstow species carries this lineage back into 
the later Barstovian, but does not appreciably clarify the exact ancestry 
of the genus. There are many resemblances in dentition between 
Megahippus and its contemporary Hypohippus. These resemblances 
can be interpreted as implying either the derivation of the larger, spe- 
cialized Megahippus directly from species that would be recognized as 
belonging to the genus Hypohippus or the derivation of both genera 
from a common anchitherine ancestor in the earlier Miocene. The 
genus Anchitherium, long regarded as ancestral to Hypohippus, may 
also have yielded Megahippus as an independent derivative. 

Roughly contemporaneous with Megahippus mckennai is the well 
known Hy pohippus osborni from Pawnee Creek (probably the Kenne- 
saw fauna), Colorado, and H. nr. affinis from ‘Tonopah, Nevada. The 
oldest well known species of Hypohippus is H. equinus from the Bar- 
stovo-Hemingfordian Deep River fauna, Montana. Of about the same 
age as the latter are specimens referred to H. osborni from various 
Great Basin localities. Matthew (1924:72) mentions isolated teeth 
from the Hemingfordian Sheep Creek fauna which agree with the Snake 
Creek H. pertinax. These undescribed specimens are probably some of 
the earliest known representatives of the genus. It seems clear that in 
the early Hemingfordian or late Arikareean Hy pohippus merges with 
Anchitherium, yet thus far this critical stage in the evolution of Hypo- 
hippus lacks documentation. 

Hy pohippus equinus is generalized enough in structure to serve as a 


120 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


common ancestor for both the later Hypohippus species and Megahip- 
pus. It occurs too late in time to be directly ancestral to Megahippus 
unless one entertains the possibility of rapid hypertrophy of the denti- 
tion in the course of evolution of this group in Barstovian time. 

Isolated teeth and jaw fragments referred to H. osborni from the 
Virgin Valley, Sucker Creek, Skull Springs, Beatty Buttes and Mascall 
faunas are approximately contemporaneous with H. equinus. Most of 
these teeth are slightly larger than those of the holotype of H. equinus. 
and somewhat smaller than H. osborni. They show a reduced P/1 as 
in H. pertinax and H. osborni, and in addition show a tendency to de- 
velop a lingual cingulum on the protocone of the upper cheek teeth. A 
distinct crochet appears on some of the upper cheek teeth from Virgin 
Valley. Unfortunately no specimens showing the upper or lower in- 
cisors are known from these Barstovo-Hemingfordian sites. When 
Hemingfordian Hypohippus species become better known we may find 
suitable ancestral types for the Barstovian Megahippus. 

When we consider the hypothesis that Megahippus has an inde- 
pendent origin in Anchitherium we find that the North American 
Arikareean species of Anchitherium show no special affmity with 
Megahippus. None of the new or Old World species of Anchitherium 
whose skulls are known show the deeply pocketed facial fossae typical 
of Megahippus. However, if we examine the Old World species we do 
find forms whose dental features approach Megahippus. This is par- 
ticularly true of the Asiatic species Anchitherium gobiense Colbert, 
1939, which differs from the European A. aurelianense and approaches 
Megahippus in such features as increased size, enlargement of I/1-2 
and reduction of I/3, reduction of P/1, and lack of ribs on the labial 
surface of the ectoloph between the styles. Unfortunately the two oc- 
currences of A. gobiense, the Chokrak formation of the northern Cau- 
casus (Tortonian, Borissiak, 1945) and the Tung Gur formation of 
Mongolia (Sarmatian), are too late in time to represent animals con- 
ceivably ancestral to Megahippus. Anchitherium gobiense probably 
represents an Asiatic line of Anchitherium showing dental trends par- 
allel to that of Megahippus just as its contemporary Paranchitherium 
karpinskii Borissak, 1937, parallels Parahippus. Nevertheless it is 
worthy of note that the closest approach to the dental peculiarities of 
Megahippus lie in the Asiatic anchitherines. 


TABLE 1 


Measurements of the Palate and Dentition of Megahippus mckenna: 
The lengths of the upper molariform teeth were measured across the 


New Megahippus from California 124 


occlusal surface from the protoconule to the hypostyle. Their widths 
were measured at the base of the crown from the paracone to the proto- 
cone. The lengths and widths of the incisors, canine and P1/ are maxi- 
mum diameters. 


Approximate length of palate from posterior 


palatine border to tip of premaxillary 165 
13/-C diastema 10.2 
C-P1/ diastema ca 25 
Width across premaxillary outside 13/ 4.4.1 
Left I1/ - Length Aol 
Width ca 17 
Right 12/ Length IDG 
Width ca 13.5 
Left Canine Length ca 9 
Width 7.8 
Left P1/ Length ES 
Width I 
Left P2/ Length srl 
Width 29.7 
Left P3/ Length 26.4 
Width 3553 
Left P4/ Length 27.4 
Width 38.2 
Left M1/ Length DEY 
Width So) 
Left M2/ Length 2553 
Width ca 39 
Right M3/ Length 91.3 
Width, paracone to protocone Silo7/ 
Width, metacone to hypocone Dies 
Left P2/-P4/ Length 81.5 


Left M1/-M3/ Length ca 77 


122 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 2, 1962 


LITERATURE CITED 
BARBOUR, T. H. 


1914. A new fossil horse, Hypohippus matthewi. Nebraska Geol. Surv., 4:169-173, 
1 fig. 2 pls. 


BORISSIAK, A. 
1945. On the Equidae from the middle Miocene of Northern Caucasus. Trudy 
Paleontologicheskogo Instituta Akademii Nauk SSSR, 13:1-52, 14 figs. 3 pls. 


COLBERT, E. H. 
1939. A new anchitheriine horse from the Tung Gur formation of Mongolia. 
Amer. Mus. Novitates, No. 1019, 9 pages, 3 figs. 


HENSHAW, P. C. 
1942. A Tertiary mammalian fauna from the San Antonio Mountains near Tono- 
pah, Nevada. Carnegie Inst. Wash., Pub. 530, p. 77-168, 7 figs. 11 pls. 


LEWIS, G. E. 
1960. Miocene vertebrates of the Mojave Desert. Bull. Geol. Soc. Amer., 71:1916. 


MATTHEW, W. D. 
1924. A third contribution to the Snake Creek fauna. Bull. Amer. Mus. Nat. Hist.. 
50:59-210. 63 figs. 


McGREW, P O. 
1938. The Burge fauna, a lower Pliocene mammalian assemblage from Nebraska. 
Univ. Calif. Publ., Bull., Dept. Geol., 11:437a-437c, 438-585, 12 figs. 


OSBORN, H. E 


1918. Equidae of the Oligocene, Miocene, and Pliocene of North America. Mem. 
Amer. Mus. Nat. Hist., n. ser., 2:1-217, 173 figs., 54 pls. 


REPORT OF AN ECHIUROID WORM HAMINGIA ARCTICA DAN- 
IELSSEN AND KOREN FROM THE BEAUFORT SEA. While the icebreaker 
USS Burton Island was enroute to establish a research station on floe ice in the 
Beaufort Sea, five bottom samples were taken. A complete echiuroid worm of the 
species Hamingia arctica Danielssen and Koren was brought up in an orange peel 
bucket by John Tibbs on September 6, 1960. This represents the first report of this 
species in normal substrate in the Beaufort Sea, although the find was not unex- 
pected. MacGinitie (1955) reports that H. arctica was washed onto the shore at 
Point Barrow, Alaska after a storm. There are other reports of this species from 
arctic and antarctic seas. 

Hamingia arctica has been found from a depth of 20 to 440 meters and a re- 
lated species, Hamingia sibogae has been reported by Sluiter (1902) from 4391 
meters in the Banda Sea. These specimens have been found in clay or mud. The 
specimen found by Tibbs was recovered from 110 meters in clayey silt. 

This specimen agrees with the description of Wesenberg-Lund (1934) in both 
internal and external characteristics including the observation that in her ma- 
terial, the papillae described by Danielssen and Koren (1881) were lacking. 
The only exception to her account is that the proboscis was somewhat shorter in 
relation to the rest of the body. This difference is considered to be not significant. 

The specimen was fixed in 7% formalin sea water buffered with hexamethyl- 
enamine and preserved in 70% ethanol. Measurements of this preserved speci- 
men are: 


Length of trunk 35mm. 
Widest diameter of trunk 18mm. 
Length of proboscis 22mm. 
Length of bifurcated part of proboscis 22mm. 


These studies were aided by a contract between the Office of Naval Research, 
Department of the Navy, and the University of Southern California, NR 107-567. 
We also gratefully acknowledge the use of the laboratory facilities of the Allan 
Hancock Foundation.—Carolyn Brahm and John L. Mohr, Biology Department, 
University of Southern California, Los Angeles 7. 


LITERATURE CITED 


DANIELSSEN, D. C., and J. KOREN 
1881. Gephyrea. Den Norske Nordhavs-exped. (1876-1878) 4:1-58. 


MacGINITIE, G. E. 
1955. Distribution and ecology of the marine invertebrates of Point Barrow, 
Alaska. Smithson. Misc. Coll. 128:1-201. 


SLUITER, C. P 

1902. Die Sipunculiden und Echiuriden der Siboga-Expedition, nebst zusam- 
menstellung der Ueberdies aus dem indischen Archipel bekannten Arten. 
Siboga-exped. 25:1-53. 


WESENBERG-LUND, E. 


1934, The Scoresby Sound committee’s second East Greenland expedition in 1932 
to King Christian [X’s Land. Gephyreans and Annelids. Medd. Gronland 
104(14) : 1-38. 


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Southern California 
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LOS ANGELES, CALIFORNIA 


CONTENTS 


Suspected Melanophore Movement in Fishes beyond the Larval 
Stage. Melba C. Caldwell and David K. Caldwell ........... 
Taxonomic Changes and Descriptions of New Tingidae (Hemip- 
tera). Carl J. Drake and Florence A. Ruhoff ..............- 
Fossil Arthropods of California. 24. Some Unusual Fossil Arthro- 
pods from the Calico Mountains Nodules. W. Dwight Pierce 
PTLOG OTIS TET OFT, S72) oyte esc! aussie ees pec eae wien s biacdi ties « 
On the Recognition of a Second Species of the Genus Pelagophycus. 
PERPLIZI TD OUDSOTD a8 Meh AD vig Sac = ally ale alae cis dies) aiaie S16 4:6) (5 8s 
Blood Parasites of Mammals of the Californian Sierra Nevada 
Foothills, with Special Reference to 7rypanosoma cruzi Chagas 
and Hepatozoon leptosoma Sp. N. Sherwin F Wood ......... 
A New Species of Chigger, Genus Euschoengastia (Acarina, Trom- 
biculidae), with Notes on Other Species of Chiggers from the 
Santa Ana Mountains, California. Richard B. Loomis and 
Marilyn Burmell ..........2.0 20s cece eee rete ee ee es 
A New Ambrysus from South America (Hemiptera, Naucoridae). 
UME ENELICT Stas ier ee ae) seas) 20! alah n sla aielelnieiar vhs lacs! ¢ atele s 


coe ee ae eres ae caeee rere seesr eee seen o 


Issued October 24, 1962 


BULLETIN en THE 


VoL. 61 Juiy-SEPTEMBER, 1962 Part 3 


185 


Southern California 
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VoL. 61 JuLy-SEPTEMBER, 1962 Part 3 


SUSPECTED MELANOPHORE MOVEMENT IN FISHES 
BEYOND THE LARVAL STAGE? 
Mesa C. CALDWELL? AND Davin h. CALDWELL® 
Los Angeles County Museum 


In his summary of the migration of melanophores, Parker (1948: 256) 
stated that melanophore movement “‘is limited in amount and restricted 
to the very early stages in the life of these cells” and that “locomotor 
activity ...is apparently characteristic of chromatophores only in 
their very early stages of growth” Parker (/oc. cit.) summarized the 
conclusions of Franz (1935) by stating that “after they [chromato- 
phores] have become members of a general pattern, even in larval in- 
dividuals, they retain their places in that pattern and their original 
locomotor powers seem to subside almost completely?’ However, it has 
recently been strongly indicated (Caldwell, M. C., in press; Caldwell, 
D. K., in press) that melanophores continue to move well beyond the 
larval stage in some marine fishes that undergo metamorphosis with 
an environmental change. This migration was shown for melanophores 
in the second dorsal fin of an Atlantic goatfish, Mullus auratus Jordan 
and Gilbert, up to a standard length of 47 mm., and for the soft dorsal, 
soft anal, and caudal fins of the Atlantic short bigeye, Pseudopria- 
canthus altus (Gill), to a length of 73 mm. In view of these findings 
the generally accepted assumptions concerning melanophores in fishes 
must be restricted to observed species. 

In the development of the color patterns of certain marine fishes 
(e.g., M. auratus, P. altus, Chaetodon striatus Linnaeus) the fins are 
particularly susceptible to drastic color pattern changes at the time of 
change of habitat. The change of pattern, however, is not restricted 
to the fins. A change in body color pattern, as well as that of the verti- 


1Portions of this study were aided by a grant to the senior author from The 
Society of the Sigma Xi RESA Research Fund. 


“Also, Department of Zoology, University of California, Los Angeles. 
3Also, Research Associate, Florida State Museum and Collaborator in Ichthyol- 


ogy, Institute of Jamaica. 


129 


130 = Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


cal fins, was noted by Gosline and Brock (1960:200, 201, Fig. 217 h 
& i) for the Pacific butterfly fish Chaetodon lunula (Lacépede). The 
changes are probably as much the rule as the exception among tropical 
shallow-water marine fishes with a pelagic early stage. These changes 
in color pattern should not be confused with that so frequently seen 
in fishes responding to shifting changes in their environment, in which 
the distribution of the pigment within the melanophores changes, often 
very rapidly (Odiorne, 1957). 

The first question raised in regard to a change in color pattern is 
whether the melanophores arise in situ at the time of change of pat- 
tern. Although only preserved fish were examined, the definite im- 
pression gained in our observations is that the melanophores move and 
that there is not a successive generation of melanin aggregates (Fox, 
1957; Blackburn, 1950: 155) and degeneration of melanophores 
(Odiorne, 1936: 28). Examinations of series of metamorphosing M. 
auratus and P. altus clearly indicate progressive migration of melano- 
phores from the base of the fins to a geneticaally fixed area on the fin. 
The path of melanophore migration is out along the fin rays, although 
some melanophores cross the interray membrane of the fin before com- 
ing to rest (Caldwell, M. C., 7 press; Caldwell, D. K., in press). Enough 
of these im proper location form the adult color patterns of the fins 
typical of the species. 

Ageregations of melanin-containing cells in regions of encysted 
unidentified skin parasites (Fig. 1) in large preserved specimens 
of Mulloidichthys xanthogrammus (Gilbert), one of the Pacific goat- 
fishes, raises the possibility of the melanophores’ ability to migrate even 
when the fish has become adult. The tissue surrounding parasitized 
areas shows what appears to be a streaming of elongated melanin- 
containing cells either into or away from the site of the cyst. Direct 
observation of the parasitized areas should be made in vivo during the 
formation of the aggregations to determine the direction of the flow. 
If the cells arise in the infected area, and stream outward, they are 
probably atypical cells caused by the parasite or melanin-containing 
macrophages. If they migrate into the area, the melanophores could 
possess a defense role similar to that of the leucocytes which they so 
closely resemble histologically. This would necessitate a review of the 
origin and role of the macromelanophores found in melanotic tissue 
(Gordon, 1957) as it should be established that they are not initially 
present as defense mechanisms. 

A parasitized area on the lower jaw in a region in which melano- 
phores normally do not exist contained no melanin-containing cells at 


Melanophore movement in fishes 131 


the site of infection. If melanophores arise in situ, they should be ex- 
pected here as well as in other infected parts of the body. With one 
exception, in all areas of M. xanthogrammus where there exists a 
generalized spotting of melanophores and there was a parasite, there 
was an accompanying aggregation of melanin-containing cells at the 
site. The exception was an embedded parasite, smaller than the others, 
that may well have been of too recent implantation to have stimulated 
a reaction. 


Figure 1. Encysted unidentified parasite in the 
caudal fin interray membrane of a Pacific 
goatfish, Mulloidichthys xanthogrammus, 
showing associated heavy aggregation of mel- 
anin-containing cells. Normal scattering of 
melanophores (retouched in the photograph) 
can be seen at the extreme top, left and bottom 
of the photograph. (About X 10.) 


The reaction of these cells is probably due to a chemical rather than 
to a physical stimulus. One of the parasitized goatfish was found to have 
two embedded and overgrown sand grains within the skin, neither of 
which showed an aggregation of these cells, although they were in 
an area normally having scattered melanophores. 

The site of implantation of one of the parasites was dissected and 
pigmentation was found to extend into the muscle tissue surrounding 
the parasite. 


132 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


This aggregation of melanin-containing cells around encysted para- 
sites is probably not unusual. Hoffman (1960:439) refers to the cysts 
as being “pigmented (usually with melanin, black . . .;’ but he makes 
no mention of the term melanophores or of movement of pigment into 
the infected area. 


ACKNOWLEDGMENTS 


We wish to thank the following for commenting on earlier versions of 
the manuscript: Dr. Joseph M. Odiorne of the New York University 
School of Medicine, Dr. Denis L. Fox of Scripps Institution of Ocean- 
ography, and Dr. Otto H. Scherbaum of the Department of Zoology 
of the University of California, Los Angeles. 


LITERATURE CITED 

BLACKBURN, M. 

1950. The Tasmanian whitebait, Lovettia seali (Johnston) and the whitebait 
fishery. Australian J. Marine and Freshwater Res., 1:155. 

CALDWELL, DAVID k. 

In press. Development and distribution of the short bigeye, Pseudopriacanthus 
altus (Gill), in the western North Atlantic. U. S. Fish and Wildlife Serv.. 
Fish, Bull., 62(203): in press. 

CALDWELL, MELBA C. 

In press. Development and distribution of larval and juvenile fishes of the family 
Mullidae in the western North Atlantic. U. S. Fish and Wildlife Serv., Fish. 
Bull., 62, in press. 

FOX, DENIS L. 

1957. The pigments of fishes. Jn Brown, Margaret E., Editor, The physiology of 
fishes. New York: Academic Press, Inc., Vol. 2, pp. 367-385. 

FRANZ, V. 

1935. Struktur und Mechanismus der Melanophoren im Farbenkleid der Teleos- 
tier. Zeitschrift Zellforschung, 23:150-197. (Not seen.) 

GORDON, MYRON 

1957. Physiological genetics of fishes. Jn Brown, Margaret E., Editor, The physi- 
ology of fishes. New York: Academic Press, Inc., Vol. 2, pp. 431-501. 

GOSLINE, WILLIAM A., anp VERNON E. BROCK 

1960. Handbook of Hawaiian fishes. Honolulu: University of Hawaii Press, 372 
pp. 277 figs. 

HOFFMAN, GLENN L. 

1960. Synopsis of strigeoidea (trematoda) of fishes and their life cycles. U. S. Fish 
and Wildlife Serv., Fish. Bull., 60 (175) :439-469. 

ODIORNE, JOSEPH M. 

1936. The degeneration of melanophores in Fundulus. J. Exp. Zool. 74:7-39. 

1957. Color changes. In Brown, Margaret E., Editor, The physiology of fishes. 
New York: Academic Press, Inc., Vol. 2, pp. 387-401. 

PARKER, GEORGE H. 

1948. Animal colour changes and their neurohumours. Cambridge: Cambridge 
University Press, 377 pp., 126 figs. 


TAXONOMIC CHANGES AND DESCRIPTIONS 
OF NEW TINGIDAE 
(HEMIPTERA ) 

Carx J. DRAKE AND FLORENCE A. RUHOFF 


Smithsonian Institution! 


This paper describes a new genus and three new species of Tingidae. 
The taxonomic changes include the suppression of several trivial names 
of species as synonyms and the creation of either new names or resur- 
rection of available names for homonyms that have been theretofore 
overlooked. The location of the holotype is recorded beneath the de- 
scriptions of each new species. We are indebted to Elmor Stromberg 
and Lisa Biganzoli, both of Washington, D.C., for the fine illustrations. 
This study was supported in part by National Science Foundation 
Grant 18721. 
New SyNoNnyMs 


Diplocysta nubila Drake is suppressed as a synonym of Penottus mon- 
ticollis (Walker ) ; Stephanitis oschanini Vasilev as synonym of S. pyri 
(Fabricius): Jeleonemia lanceolata (Walker ) as synonym of T tricolor 
(Mayr); Tingis bengalana Drake as synonym of Dasytingis rudis 
Drake and Poor; Tingis himalayae Drake as synonym of T: buddleiae 
Drake; Urentius abutilinus Priesner and Alfieri as synonym of 
U. eunonymus Distant; and U. sentis Distant as synonym U. hystri- 
cellus (Richter ). 


Naitingis, NEW GENUS 


Obovate or oblong, dorsal and ventral surfaces moderately clothed with 
very short golden pubescence. Head very short, armed with the usual 
dorsal spines; bucculae wide, areolate, closed in front; eyes moderately 
large; antenniferous tubercles short, thick, bluntly rounded in front, 
concave within. Antennae long, slender, segment I and II short, III 
very long, and IV moderately long, only slightly swollen. Rostrum 
long, reaching to end of sulcus; sternal laminae of rostral sulcus low, 
areolate, widely separated from each other, open at base. Ostiolar sulcus 
of metathoracic scent glands present on each metapleuron. 

Pronotum moderately convex, punctate, tricarinate; collar distinct, 
areolate, truncate in front, without hood; paranota narrow, erect, com- 
posed of one row of small cells; outer margins of paranota and elytra 
without hairs or bristles. Elytra slightly wider and longer than abdo- 
men, divided into the usual division; costal area narrow, composed of 


1United States National Museum (USNM). 


133 


134. Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


one row of areolae, sometimes biseriate in hollow of costal area; dis- 
coidal area extending beyond middle of elytron, on same level as the 
sutural area; sutural areas overlapping each other so that their apices 
he jointly rounded in resting position. Metathoracic wings longer than 
abdomen. Legs moderately long, femora slightly swollen. 


Type Species, Tropidocheila mayne 
Schouteden, Africa 
This new genus is founded for the reception of Tropidocheila maynei 
and var. biseriata Schouteden, T) nyanzae Schouteden and Tingis 
blukwana Drake. Biseriata represents a variety with a partial second 
row of areolae in hollow of costal area. 

Naitingis, n. gen., may be separated from Tingis and its subgenera 
by the lack of pronotal hood, very narrow paranota, and dorsal cloth- 
ing of very short golden hairs, The exterior margins of the paranota 
and elytra are without hairs or bristles. 


Engynoma isolata, NEW NAME 
Tingis insularis Hacker 1929, Mem. Queensland Mus., vol. 9, p. 330. 
pl. 34, fig. 10. 


Engynoma insularis: Drake 1942, Jour. Washington Acad. Sci., vol. 
S27 O07. 


Since the trivial names of Tingis insularis (Horvath) (1902) and 
Tingis insularis Hacker (1929) are homonyms, we are here proposing 
the specific name of isolata for the species described by Hacker. 


Tingis strictula (Puton) 
Monanthia platychila strictula Puton 1878, Ann. Soc, Ent. France. 
ser. 5, vol. 8, p. Ixvi. 
Tingis strictula; Horvath 1906, Ann. Mus. Nat. Hungarici, vol. 4, 
Sante 
Tingis montana Lindberg 1932, Soc. Sci. Fennica Comm. Biol., vol. 3, 
p. 42. (New Synonymy ) 


A comparison of a paratype of 7. montana Lindberg from Spain with 
a specimen of 7? strictula (Puton) (det. Puton) from Oran, Algeria, 
shows that these two trivial names were given to the same species, the 
latter having priority by many years. The illustration by Lindberg 
(loc. cit. p. 42, pl. 2, fig. 5) is Tingis cardui (Linnaeus) and not 7: mon- 
tana as labeled. 


Tingidae taxonomy 135 
Tingis stachydis (Fieber ) 
Monanthia maculata Herrich-Schaeffer 1838, Wanzen. Insekt., vol. 4, 
pp. 51, 56, pl. 123, fig. 389, a-c. 
Monanthia stachydis Fieber 1844, Ent. Mon., p. 73, pl. 6, figs. 13-15. 


Tingis maculata: Horvath 1906, Ann. Mus. Nat. Hungarici, vol. 4, pp. 
69, 84. 


The transference of Monanthia maculata Herrich-Schaeffer to the 
genus Tingis by Horvath (1906) made the specific names of this species 
and that of Tingis maculata Herrich-Schaeffer homonyms. As the first 
revisors, we are here resurrecting Monanthia stachydis Fieber as the 
available trivial name for the former species, and are citing the refer- 
ences needed to verify the homonym and validity of stachydis as the 
valid trivial name. 


Galeatus maculatus (Herrich-Schaeffer ) 


Tingis maculata Herrich-Schaeffer 1838, Wanzen. Insekt., vol. 4, p. 68, 
pl. 126, fig. 393. 


Tingis subglobosa Herrich-Schaeffer 1838, Wanzen. Insekt., vol. 4, 
p. 68.—Fieber 1844, Ent. Mon., p. 106, pl. 9, figs. 17-20. 


Galeatus maculatus: Stal 1874, Ofv. Vet. Forh., p. 48. 


As explained beneath the above caption, Jingis maculata Herrich- 
Schaeffer and Monanthia maculata Herrich-Schaeffer (now in Tingis ) 
are homonyms. Stal (1874) correctly transferred 7’ maculata to the 
genus Galeatus. 

Galeatus spinifrons (Fallén) 


Tingis spinifrons Fallén 1807, Monographia cimicum Sveciae, p. 38. 

Galeatus spinifrons: Horvath 1906, Ann, Mus. Nat. Hungarici, vol. 4, 
19, OS: 

Copium artemisifolium Shinji 1938, Zool. Mag. Japan, vol. 50, p. 316. 
—Takeya 1951, Kurume Univ. Jour. (Nat. Sci.), vol. 4, p. 19. (New 
Synonymy ) 

The type of C. artemisifolium Shinji from Honshti Island (Morioka 
Island, Prov. Rikuchu), Japan, is apparently lost. Since the original 
description of artemisifolium fits that of Galeatus spinifrons and since 
it breeds on the leaves of Artemisia vulgaris, we are here treating the 
two trivial names as synonyms, spinifrons having the priority by many 
years. In a paper in press elsewhere, the authors have placed other 
species in synonymy with spinifrons. 


136 = Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


Leptodictya elitha, NEW SPECIES 
Large, very broad. Head blackish fuscous with dorsal spines testaceous; 
bucculae dark fuscous. Pronotum blackish fuscous with hood, collar. 
paranota, carinae, and apex of hind projection testaceous. Elytra pale 
testaceous with exterior veins, six obliquely transverse bands (each 
band consists of one depressed vein and adjoiming rows of cells) in 
costal area blackish fuscous; subcostal area with veinlets brown; dis- 
coidal area with boundary veins and two adventitious veins brown to 
fuscous; sutural area with veimlets blackish fuscous. Antenna fuscous- 
brown with first two segments blackish. Legs ight brown with tarsi 
blackish. Body beneath black, with entire abdomen or only genital 
segments reddish brown. Length 4.70 mm., width (elytra) 2.70 mm. 

Head very short, armed with five, very long, sharp spines; eyes 
moderately large, dark fuscous; bucculae long, wide, five areolae deep 
in widest part, closed in front. Antennae long, slender, clothed with 
whitish pubescense, measurements: segment I, 0.25 mm.; II, 0.12 
mm.; III, 3.50 mm.; IV, 0.75 mm. Labium testaceous, extending to 
base of sulcus; laminae of sulcus very wide, mostly biseriate, with 
large V-shaped opening at base, without laminae on prosternum. 

Pronotum moderately convex, coarsely punctate, tricarinate; me- 
dian carina composed of one complete row of moderately large areo- 
lae, plus two extra cells just behind the hood; lateral carinae less raised 
than median carina, uniseriate, slightly constricted behind middle of 
pronotal disc; hood moderately large, strongly compressed laterally. 
about twice as long as high, feebly produced in front, extending back- 
wards between calli on fore part of pronotal disc; paranotum fairly 
wide, outer part reflexed so as to leave two rows of areolae on underside 
and then between two and three rows on upper side, only the outer 
row of latter resting on the dorsal surface of pronotum; posterior proc- 
ess triangular, areolate. Ostiole and ostiolar canal prominent on each 
metapleuron, the side of the channel raised, vertical and slightly tilted 
posteriorly. 

Elytra sharply widened at base, much wider and longer than abdo- 
men, sutural areas only partly overlapping so that their apices, in 
resting position, are separated from each other; costal area very wide 
with areolae irregular in size, arrangement, and not in regular rows, 
ten to eleven cells deep in widest part; subcostal area very narrow. 
vertical, composed of two rows of small areolae, discoidal area extend- 
ing beyond middle of elytron, acutely angulate at base and apex, seven 
areolae deep in middle; sutural area large, with areolae as variable in 


Tingidae taxonomy 137 


size and as confused in arrangement as in costal area. Hind pair of 
wings slightly longer than abdomen. 

Holotype (male) and allotype (female), macropterous, Queros, Rio 
Cosnipata, Department Cuszo, Peru, l.iv. 1952, EF Woytkowski, in 
Drake Coll. (USNM). 

The markings of the elytra are prominent and very similar in ap- 
pearance to those in L. approximata. It differs from the latter species 
by its larger size, slightly more elevated pronotal carinae, longer ce- 
phalic spines, and especially by the longer and distinctly laterally 
compressed hood. These two species have the sternal laminae of the 
rostral sulcus biseriate. 


Amblystira angella, NEW SPECIES 
FicureE 1 


Small, oblong, deep black, shiny, with hollow part of costal area brown- 
ish testaceous; eyes moderately large, black; body beneath black, shiny, 
sparsely clothed with pale pubescence; laminae of rostral sulcus black, 
dull. Labium brownish with apex black. Antenna pale brown with 
fourth segment black and shiny, the pubescent hairs sparse on third 
segment, much more numerous and longer on fourth segment. Legs 
pale brown with tarsi black, pubescence sparse and pale, Length 2.60 
mm., width (elytra) 1.07 mm. 

Head very short, with an irregular, median, longitudinal furrow 
on vertex, armed with three, short, blunt anterior tubercles and two 
longer, closely appressed, ridgelike occipital spines; buccal laminae 
wide, areolate, black, closed in front. Labium extending to middle of 
mesosternum; laminae of sulcus low, widely separated from each 
other, composed of one row of tiny cells, present on all three sternal 
divisions, closed at base. Antenniferous tubercles very short, thick, 
blunt, concave within. Antenna moderately long, slender, measure- 
ments: segment I, 0.10 mm.; II, 0.09 mm.; III, 0.70 mm.; IV, 
0.38 mm. 

Pronotum moderately convex, coarsely punctate, tricarinate, each 
carina composed of a single row of very small areolae, lateral carinae 
less raised than median and slightly divergent anteriorly; collar raised, 
composed of three transverse rows of tiny punctures; calli small, 
impressed, impunctate; paranotum narrow, slightly reflexed, slightly 
wider at humeral angle, with two to three rows of tiny punctures, 
distinctly obtusely angulate opposite humeral angle. Metathoracic 


138 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


Figure 1. Amblystira angella, n. sp. 


scent glands with a prominent, nearly vertical sulcus on each meta- 
pleuron. 

Elytra with outer margins (also those of paranota) mimutely ser- 
rate, longer than abdomen; costal area narrow, composed of one row 
of small rounded cells on basal half, thence posteriorly in hollowed 
part wider with large areolae and bi- to triseriate; subcostal area wide, 
five areolae deep in widest part (six opposite apex of discoidal area) ; 


Tingidae taxonomy 139 


discoidal area reaching to middle of elytron, widened posteriorly, 
obliquely truncate at apex, there six areolae deep, acutely angulate 
at base; sutural areas overlapping each other at rest. Hind pair of 
wings nearly as long as elytra, pale fuscous. 

Holotype, macropterous female, (fig. 1), Monson Valley, Tingo 
Maria, Peru, 29.11.1954, E. I. Schlinger and E. S. Ross, in Calif. Acad. 
Sciences, San Francisco. 

This shiny black species can be separated at once from all other 
members of the genus by having the paranota distinctly angulate 
opposite the humeral angles. As in other members of the genus, the 
clavus is narrow and concealed beneath the backward projection of 
the pronotum when elytra are in resting position. 


Haedus oios, NEW SPECIES 
FIGURE 2 


Small, oblong, widest across apices of elytra, tapering anteriorly with 
a large hollowed sinus behind the middle of outer margin of each 
elytron; testaceous with head, pronotal disc and the discoidal, subcostal, 
and sutural areas of elytra brownish, also with a wide transverse band 
across apices of elytra dark fuscous; body beneath pale brown, shiny. 
Antenna pale testaceous with fourth segment slightly brownish. Legs 
pale testaceous with tip of tarsi blackish. Rostrum testaceous with tip 
blackish. Buccal and sternal laminae of rostral sulcus testaceous, An- 
tenniferous tubercles and cephalic spines pale testaceous. Length 2.65 
mm., width (across middle and apices of elytra) 0.82 mm., and 0.95 
mm., respectively. 

Head short, armed with five very long, subporrect spines; bucculae 
wide, mostly four areolae deep, closed in front. Labium extending to 
middle of mesosternum; sternal laminae of sulcus uniseriate, com- 
posed of one row of areolae, channel narrow and open at base. Antenna 
very slender, sparsely clothed with short setal hairs, with longer hairs 
on fourth segment; measurements: segment I, 0.10 mm.; IT, 0.08 mm.; 
IIT, 1.20 mm.; IV, 0.32 mm. Antenniferous tubercles very long, trans- 
versely compressed, feebly curved inwardly, extending slightly be- 
yond apices of second sternal segments. Orifice and ostiolar channel 
not visible on metapleuron. 

Pronotum with sides narrowed anteriorly, moderately convex, retic- 
uately punctate, tricarmate; median carina much higher than lateral 
pair, raised anteriorly, biseriate behind hood, uniseriate with larger 
rectangular areolae behind middle of pronotal disc, there with areolae 


140 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


% ae ¢ 
? 
ee 
F dinkeRecasee 
PUN OXCS fe 
| BR ay 
7 per BOCK Wt 
ae x t 
: \ 
4 ‘ = 


Figure 2. Haedus oios, n. sp. 


Tingidae taxonomy 141 


shorter than high; lateral carinae low, indistinctly areolate, nearly 
parallel; hood small, tectiform, angulately produced in front; para- 
notum narrow, long, slightly reflexed, mostly uniseriate, wider and 
biseriate opposite calli and collar. Legs very slender, indistinctly pu- 
bescent. 

Elytra with outer marginal veins (also that of paranota) minutely 
serrate, apices partly overlapping in resting position; costal area fairly 
wide, mostly biseriate; discoidal area mostly biseriate, nearly vertical, 
extending a little beyond middle of elytron, widest behind middle, 
there four areolae deep, with boundary vein separating it from subcos- 
tal area obtusely angulately raised, highest behind middle, there and at 
apex with black spots; sutural area with areolae subequal in size to 
those in discoidal area; hypocostal laminae uniseriate, the areolae small 
and rounded. Abdomen, as seen from beneath, rather densely clothed 
with short, pale hairs along the upper margins of each side. Male para- 
meres strongly curved inwardly. 

Holotype, macropterous male, Salisbury, Southern Rhedesia, col- 
lected by C. M. Smithers, in Drake Collection (USNM). 

This species can be separated from other members of the genus by 
the strongly constricted elytron behind the middle and the form of 
body and color markings as shown in the illustration of the type. 


Dictyonota maroccana Ribaut 


Dictyonota maroccana Ribaut 1939, Bull. Soc. Sci. Nat. Maroc, vol. 19, 
p. 186. 


An examination of the type (lectotype) of D. maroccana Ribaut, 
brachypterous female, Djebel M’Goun Canyon Tessaut, Morocco, 
1.1X.1915, shows that it belongs to the subgenus Alcletha Kirkaldy, 
and that it is a synonym of aridula Jakovlev. Maroccana differs from 
the latter in having the first antennal segment slightly longer and the 
antenniferous tubercle more narrowed anteriorly and pointed at the 
apex. The other structural characters are similar to each other. 


Dictyonota tricornis (Schrank ) 

Acanthia tricornis Schrank 1801, Fauna Boica, p. 67. 

Dictyonota dictyesthes Garbiglietti 1869, Bull. Soc. Entom. Italiana, 
vol. 1, p. 274. —Drake and Ruhoff 1960, Proc. U.S. Natl. Mus., vol. 
112, p. 7. (New Synonymy) 

Dictyonota tricornis: Horvath 1906, Ann, Mus. Nat. Hungarici, vol. 4, 
p. 41. 


142 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


Dictyonota ifranensis Vidal 1951, Bull. Soc. Sci. Nat. Maroc, vol. 31, 
p- 63. (New Synonymy ) 

Dictyonota tricornis americana Parshley 1916, Psyche, vol. 23, p. 164. 
(New Synonymy ) 


An examination of the holotype of Dictyonota ifranensis Vidal, 
macropterous female, Ifrane, Morocco, VI.1938, J. P. Vidal, shows 
that it is inseparable from and a synonym of D. tricornis (Schrank) 
(det. Horvath). An examination of many specimens of tricornis from 
Europe shows that it is very variable in size and in the areolation of 
the paranota and elytra, especially costal area. D. tricornis americana 
Parshley is also suppressed here as a synonym of tricornis, as well as 


dictyesthes Garbighetti. 


FOSSIL ARTHROPODS OF CALIFORNIA. 24. SOME UNUSUAL 
FOSSIL ARTHROPODS FROM THE CALICO 
MOUNTAINS NODULES. 


W. Dwicut PIERCE 
Los Angeles County Museum 
and 
JoHNn Grpron, Sr. 
Campbell, California 


The unusual silicified arthropods recovered from Miocene petrolif- 
erous nodules, described in this article, were collected by Gibron at 
U. S. Geological Survey Site #19057, the original discovery site of 
Dr. Allen M. Bassett, reported on by Dr. Allison R. Palmer (1957). 
This site is located in S.W. 1/4, Section 24, R.1.E., T-10.N., Calico Moun- 
tains, San Bernardino County, California, at 2420 feet altitude. We 
believe that this is the oldest fossil site yet reported on in this mountain 
range. 

The specimens were all extracted from the nodules by use of hydro- 
chloric acid by Gibron and Mrs. Gibron (Julia). Critical study has 
been made by Pierce, who also prepared the figures. 


Class ARACHNIDA 
Order ARANEI 
Family Dictynidae 
Genus Argenna Thorell 


Argenna fossilis PETRUNKEVITCH 1957 
FicurEs 1A AND 1B 


A beautiful, crystalline spider; specimen 5275, (Gibron #H) was ob- 
tained from the type site of the Petrunkevitch species, and although 
larger than the type probably represents an older instar, It is mounted 
in balsam. 

Total length 2.16 mm., cephalothorax 0.96 mm., abdomen 1.20 mm. 
The leg pattern (Table I) shows minor differences from those given 
in the original description. (See Palmer, 1957:246-248. ) 


TABLE [| 
Leg dimensions in millimeters of specimen No. 5275 
Patella 
and 

Leg Femur Tibia Metatarsus Tarsus Total 
J 0.68 0.68 0.40 0.36 212 
II 0.60 0.64 0.40 0.32 1.96 
III 0.60 0.64: 0.48 0.36 2.08 
IV 0.68 0.76 0.40 0.28 Dil, 


143 


144. 


Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


Fossil arthropods from California 145 


The specimen is crystal clear except as mentioned below. 

The nervous system is pigmented with black masses almost filling 
the center of the cephalothorax, from which radiate nerves reaching 
the whole length of the appendages, and a large mass in the abdomen. 

The cribellum is transverse, and the four spinnerets seem to be in 
a row, the outer ones larger with anal lobe below (Fig. 1b). 


Order CHELONETHIDA Thorell 
Suborder HETEROsPHYRONIDA J. C. Chamberlin 1929 
Family Chthoniidae Hansen 1894 


A silicified pseudoscorpion cheliceral claw was extracted from a nodule 
from the same site (19057) noted above. 

In view of the absence of any indication of poison apparatus in either 
finger, and because the chela has no pedicel, but rather a socket for 
attachment, it is assigned as follows: 

Subfamily Chthoniinae Daday 1887 
Tribe Lechytini J. C. Chamberlin 1929 
Genus Lechytia Balzan 1891 


FIGURES 2A AND 2B 


Chamberlin (1929), in his monograph of this suborder, figured the 
right chela of Lechytia pacifica (Banks) (Roncus ), which most closely 
resembles this specimen in proportions and general shape, although 
differmg in minor details. Banks (1893) very briefly described the 
hand of his species from Washington state, and Chamberlin records it 
also from California. 

Length of chela 2.32 mm.; width 0.72 mm.; movable finger length 
1.20 mm.; width 0.24 mm. 

No setae are visible. The inner margin of each finger is armed with 
small quadrate teeth, and similar teeth occur on the outer margin of 
the movable finger, in the position often occupied by spmnarets. 

A peculiar organ (Fig. 2c), looking like a tree growing out of a 


Facing page: 

Figure 1a, Side view of Argenna fossilis; size 2.16 mm. Figure 1b. Enlarged side 
view of end of abdomen of A. fossilis, showing spinnerets. Figure 2a. Cheliceral 
claw of pseudoscorpion, outer side; length 2.32 mm. Figure 2b. Inner side of base 
of cheliceral claw of pseudoscorpion, showing articulation. Figure 2c. Enlargement 
of unusual organ on finger of claw of pseudoscorpion. Figure 3. Posterior portion of 
an Entomobrya; length of specimen 1.6 mm. Figure 4. Palaeosminthurus juliae. 
new species of symphypleonid; length 2.4 mm. Figure 5a. Dorsal view of anterior 
portion of Trogiid corrodentian; length of specimen 1.6 mm. Figure 5b. Face and 
leg fragments of Trogiid corrodentian. 


146 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


circular depression, occurs near the distal fourth of the movable finger. 
The attachment of the chela is in a socket on the upper inner side. 
a little beyond the base. This claw differs from L. pacifica, in which 
the attachment is basal. 
This seems to be the first fossil evidence of the pseudoscorpions in 
America, although there are European records. 


Class INSECTA 
Order CoLLEMBOLA Lubbock 1873 
Suborder ARTHROPLEONA Borner 1901 
Family Entomobryidae (Tomésvary 1883) Gisin 1944 
Genus Entomobrya Rondani 1861 


Pierce (1960) described a fossil Collembolan, Entomobrya (Entomo- 
brya) kirkbyae from the other side of the Calico Mountains in N.W. 
Y% Section 19, R.2.E.,T10.N., a little over one mile away but in a much 
later geological horizon of the Miocene. It measured only 1.50 mm.. 
for complete insect with extended springer. 

A second record (Fig. 3) is now added, of a posterior portion of an 
Entomobrya without the complete springer. The preserved portion 
measures 1.6 mm., and therefore the entire insect would be consider- 
ably larger than E. kirkbyae, but still could be a later instar of that 
species. 

Suborder SymMPHYPLEONA Borner 1901 
The unusual specimen now to be described is in some respects inter- 
mediate between the two suborders, and in absence of a collophore 
would seem to stand by itself. The condensed form of the body other- 
wise places it in this suborder but a number of characters require 
separation to form a new family. 


Family Palaeosminthuridae, New Family 
Head with pigmented eyes beyond middle and lateral to insertion of 
antennae, and with four clear lensed ocelli on vertex. Prothorax re- 
duced, but bearing the only functional legs. Mesothorax greatly en- 
larged; metathorax dorsally distinct. Middle and hind legs greatly 
reduced. First abdominal indicated but without collophore. Anal 
papilla and furcula distinct. 


Genus Palaeosminthurus, New Genus 
Fore legs elongate; middle and hind legs rudimentary; collophore lack- 
ing; small oval spiracles on mesothorax; metathorax and first abdomi- 
nal. 


Fossil arthropods from California 147 


Palaeosminthurus juliae, NEW SPECIES 


FIGURE 4 


Named in honor of Julia, Mrs. John Gibron, Sr., in recognition of her constant 
collaboration with her husband in the work of collection and separation of the 
fossils. 


Although this interesting insect is crystallized and semi-transparent, 
the white surface pigmentation remains. 

Length, 2.4:mm.; height, 2 mm. 

Head on flexible neck, directed downward, hypognathous. Antennae 
located in front of the middle. Only two joints of one antenna remain. 
Pigmented eyes outlined by a clear oval line occur lateral to the bases 
of the antennae. On the vertex are two large oval, and two smaller 
crescent shaped ocelli with clear convex lenses. These are placed in 
the area where the so-called eye patches of separate ocelli occur in other 
Collembola. An ensiform mandible is indistinctly visible, slender, 
acute. Other mouth parts are indistinct in a foamy mass of crystal. 

The prothorax is membranous, small, with a curved cervical plate 
from base of head to anterior corner of coxae. The front legs are long 
(one is complete except for claws), with coxa, trochanter, femur and 
tibiotarsus in the proportions 30:20:50: 60. 

Mesothorax massive, humped high above prothorax, strongly con- 
vex, definitely defined by deep suture behind, but ventrally meets the 
prothorax and abdomen in a point. Faintly the prescutum, scutum 
and scutellum are defined, and on scutellum at the side is an elongate 
oval opening which seems to be a spiracle, and on one side there is a 
sinuous scar above this. The rudimentary second legs are indicated by 
two segments, upon the side of the body below the first spiracle. 

Metathorax dorsally consists of scutellum, but laterally this is pre- 
ceded by a narrow scutum, which bears a transverse opening or spira- 
cle, and below this are the rudiments of the third leg. 

First segment of abdomen is dorsally distinctly outlined, bears an 
opening or spiracle opposite that of metathorax, and the lateral bound- 
ary is faintly outlined. There is no trace of a collophore, nor is there 
any space for such an organ, as the mesothorax, metathorax, and first 
abdominal come to a single ventral point adjoining the prothorax. 

The second, third and fourth abdominal segments are one large un- 
differentiated mass, except that from the venter of the fourth springs 
the manubrium of the springer. 

This organ is soft, two-segmented, and lacks the dens and mucro in 
the present specimen. 


148 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


The anal papilla, or fifth and sixth segments from side view, has 
a dorsal and ventral process, with a central lobe (sixth). 

The presence of eyes and ocelli, absence of collophore, reduction of 
middle and hind legs, and great development of meso- and metathorax, 
set this insect off as very distinct from any insect described in either 
suborder of springtails. 


Superorder CorRRODENTIA Burmeister 1839 
Order CoPEOGNATHA Enderlein 1903 
Suborder TRoGIOMORPHA 


Family Trogiidae 


FIGURES 5A AND 5B 


A fragment of a crystallized procid is represented by Specimen 5282, 
collected and extracted by Gibron. 

This is the second record for this order, from the Miocene of the 
Calico Mountains (for the first, see Pierce, 1960:45-46.). 

Length of specimen 1.6 mm.; width of head 0.72 mm. 

The head is turned down beneath the thorax, but the broad face is 
complete with bulging eyes, broad clypeus and labrium. The thorax 
is definitely three segmented, mesothorax largest. An oval disc on 
metathorax may be a wing pad. 


Order EPHEMERIDA Leach 1817 
Family Baetidae Ulmer 1920 


FiGcuREs 6, 7, AND 8 
In article No. 23 of the Series W.D.P. recorded a fossil mayfly and her 
egg mass found in Section 19, R.2.E., T10.N., Calico Mountains. 

We are now able to report on three crystallized mayfly adults, Speci- 
mens 5096, 5286, 5307; all found by Gibron at U.S.G.S. site No. 
19057, noted above. 

Fossil mayflies date back to the Permian Period, but the record is 
based almost entirely on winged adults crushed and lying in shales. 
In these new specimens we have two crystallized adults, and one 
subimago, with only fragments of the wings, and it is hardly yet advis- 
able to name them. The small size seems to limit them to the family 
Baetidae. Only the genera Caenis, Baetis, Pseudocloeon and Cloeon 
are recorded as holding species with adult size under 3 mm. 

Organisms of decay are in all three specimens, indicating death be- 
fore preservation. This is natural as the usual life of a mayfly is but a 


day. 


Fossil arthropods from California 149 


Figure 6. Adult mayfly, No. 5286; length 1.96 mm. Figure 7. Subimago maylly, 
No. 5096; length 1.92 mm. Figure 8. Adult mayfly, No. 5307; length 3.04 mm. 
Figure 9. Adult chalcid wasp; length 1.88 mm. Figure 10a. Adult ichneumonid 
wasp; length 6.0 mm. Figure 10b. Side view of head of ichneumonid wasp. Figure 
11. Lepidopterous larva; length 1.03 mm. 


150 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


Specimen 5096 (Fig. 7) is a subimago, measuring 1.92 mm. Crys- 
tallization is not clear cut. There are round bodies in the intestinal 
tract. 

Specimen 5286 (Fig. 6a and 6b), is a mature insect, measuring 
1.96 mm. in length without its cauda; with three legs and antenna 
visible. There is considerable brown pigmentation of the thoracic and 
abdominal sclerites. Viewed from an unnatural angle, the posterior 
margin of the head appears to be arched posteriorly, neither straight 
nor emarginate, and this would seem to remove it from any of the 
genera mentioned above. The antennae (Fig. 6b), have a short, trans- 
verse first jomt, a longer but transverse second joint, and an elongate 
third joint. The legs are short. 

Specimen 5307 (Fig. 8) is a larger insect, measuring 3.04 mm., 
with only the bases of the caudal filaments present; and only the coxal 
bases of the legs. In side view it has a typical mayfly appearance with 
large meso-metathorax; small prothorax. Crystallization is not clear 
cut due to decay materials which obscure many outlines. 


Order HyMENOPTERA 
Superfamily Chalcidoidea Ashmead 


FIGURE 9 
A minute crystallized Chalcid adult, Specimen 5289 (Gibron H35) 
was recovered by Gibron, from a nodule from Site 19057, U.S.G.S. 
Length 1.88 mm., specimen pigmented dorsally. Fragments of 
wings, mouth parts, and legs are undecipherable. 


Superfamily Ichneumonoidea 


Figures 104 AND 10B 
An excellent specimen of ichneumonid wasp, Specimen 5087 was re- 
covered from a nodule found at Site 19057, U.S.G.S. 

Length 6.0 mm, Crystallized, but with reddish brown pigmentation 
on head, thorax and three abdominal segments. The head (Fig. 10b) 
has three round ocelli, but mouth parts are not distinct, and only the 
bases of antennae are present. The legs are long and slender. 


Order LEPIDOPTERA 


FicurE 11 


A crystallized first instar larva, Specimen 5079 (Gibron #6) is an 
interesting addition to the series collected at Site 19057, U.S.G.S. 


Fossil arthropods from California 151 


Size 1.03 mm. This larva has three pairs of legs, and also evidences 
of four pairs of prolegs, and two caudal cerci. It is presented 1o repre- 
sent the order until certain moths found by Mrs. Kirkby can be studied. 


LITERATURE CITED 
BANKS, NATHAN 


1893. New Chernetidae from the United States. Can. Ent., 25:64-67. 
CHAMBERLIN, J. C. 


1929. Synoptic classification of false scorpions. Ann. Mag. Nat. Hist., ser. 10, 
4:50-80. 


PALMER, ALLISON R. 

1957. Miocene arthropods from the Mojave Desert, California. U.S. Geol. Surv. 
Professional Paper 294-G; pp. 237-280, figs. 89-101, plates 30-34. 

PIERCE, W. DWIGHT 


1960. Fossil arthropods of California. No. 23. Silicified insects in Miocene nodules 
from the Calico Mountains. Bull. So. Calif. Acad. Sci., 59 (1):40-49, plates 
14-17. 


ON THE RECOGNITION OF A SECOND SPECIES 
OF THE GENUS PELAGOPHYCUS' 
E. YALE Dawson 


University of Southern California, Los Angeles? 


The Elk Kelp or Bull Kelp, Pelagophycus, is one of the best known 
seaweeds of southern California and northwestern Baja California 
where it grows abundantly in depths of 40-100 feet along the seaward 
margins of the Macrocystis beds. The very long stipe with its massive 
terminal pneumatocyst and antlers is a common sight on the beaches. 
It has been known and described in the scientific literature since 1822, 
and Setchell (1896, 1908, 1912) has given the extensive historical 
and descriptive accounts of it that have established the currently ac- 
cepted name Pelagophycus porra (Leman) Setchell. 

In May, 1961, Dr. Wheeler J. North, diving at the north tip of 
Catalina Island for a cruise of the Scripps Institution of Oceanography, 
noticed and photographed specimens of a peculiar “dwarf” Pelagophy- 
cus in which the stipes, of what appeared to be fully adult plants, were 
only about 1 meter long, and the 5-7 meter-long blades lay along the 
bottom in the manner of Laminaria farlowi. Inasmuch as neither he 
nor other experienced divers had ever seen such Pelagophycus plants 
elsewhere among the Channel Islands or along the mainland, he called 
my attention to them and proposed to make special collections for 
study. His observation brought to mind a group of juvenile, short-stiped 
Pelagophycus specimens that I had dredged at depths of 48-90 feet 
at Whites Cove, Catalina Island in 1948-49. 

Dr. North subsequently searched on five dives along the west side 
of Catalina Island from Catalina Harbor to Eagle Rock without find- 
ing specimens of the Pelagophycus again, but upon my suggestion of 
White’s Point on the east side, he succeeded in locating a colony near 
Emerald Cove from which in July, 1961, he obtained a complete series 
of specimens from juveniles to the largest adults present. These were 
growing on a bottom of sand grading into silt at depths of 50-100+ 
feet. The greatest concentration was at about 90 feet. Although adult 


1This study was conducted incidentally to other researches on Pacific marine 
algae under support of contract Nonr-3096(00) Amend. 2 between the Office of 
Naval Research and the Beaudette Foundation. Reproduction in whole or in part 
is permitted for any purposes of the United States government. 


2Allan Hancock Foundation. 


153 


154. Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


ALINS 


Figure 1, Pelagophycus giganteus. Juvenile plants collected at Catalina Island by 
W. North, 7/25/61 at a depth of 90 ft., showing progressive development of stipe 
and pneumatocyst, and division of blade. E represents a plant with 22 cm. stipe. 
Note early development of haptera from lower part of stipe. These plants are 
essentially equivalent in development to their counterparts in Fig. 2. All X 0.47. 


Second species of Bull Kelp 15 


Or 


Figure 2. Pelagophycus porra. Juvenile plants collected at La Jolla by B. Best, 

7/29/61 at a depth of 90 ft., showing progressive development of stipe and pneu- | 
matocyst, and division of blade. E represents a plant with 77 cm. stipe. These | 
plants are essentially equivalent in development with their counterparts in Fig. 1. 
All X 0.47. 


150 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


plants there were not so short-stiped as those observed and photo- 
graphed on the west side of the island in April (10 ft. vs. 3 ft.), they 
nevertheless showed distinct differences from the characteristic main- 
land plants of Pelagophycus porra, and a comparison of the two forms 
has led to a curious discovery. 

A review of the Pelagophycus literature revealed that the Swedish 
botanist Areschoug’s (1876) fragmentary type specimens of Nereo- 
cystis gigantea, a collection by Gustav Eisen from Santa Catalina 
Island, is, indeed, this dwarfish plant that shows such different char- 
acters from the mainland Pelagophycus porra. The original description 
clearly shows that Areschoug had a plant with long-elliptical vesicle 
(5-6 cm. long, 3.5 cm. wide) and a doubly furcate “petiole” above 
the vesicle. The type has been reexamined for me by Dr. Sten AhIner. 
curator of the Naturhistorska Riksmuseet, Stockholm. It agrees in all 
essential details with some of the Catalina Island topotypes recently 
collected by North. Five years later, in 1881, Areschoug set up the 
new genus Pelagophycus to receive Eisen’s plant, but in distinguishing 
it from Nereocystis he did not recognize, working in his laboratory im 
Sweden, the differences between his type specimen from Catalina 
Island and the large plants with spherical vesicles that were known 
on the southern California mainland. Neither did Setchell, many years 
later at the University of California, recognize these differences, for 
his experiences were only with mainland plants at San Pedro (Fig. 3). 
and Areschoug’s imperfect type was in Sweden. Instead, Setchell’s 
discovery of a much earlier description by Leman (1822) of the Cali- 
fornia Elk Kelp led him to assume the Areschoug plant, the Leman 
plant and his own to be a single species which he called Pelagophycus 
porra according to Areschoug’s genus but Leman’s earlier specific 
epithet. It is now clear, however, that Areschoug’s type actually rep- 
resents a different species from the common coastal Pelagophycus 
porra and is a plant of apparently restricted insular distribution. This 
species, which must be known as Pelagophycus giganteus (Areschoug ) 
Areschoug, is distinguished from P porra according to the following 
comparison: 


P giganteus P. porra 
HOLD-_ haptera developing from haptera confined to a short length of 
FAST progressively higher lev- lowermost stipe, not extensive. 10-15 


els on base of stipe, be- (20) cm. broad. usually attached to a 
coming extensive. to 60 rock. 

cm. broad or more, spread- 

ing into sand and shell. 


STIPE 


VESICLE 


ANTLERS 
BLADES 


Second species of Bull Kelp 157 


P. giganteus 


short, 10-35 cm. long in 
young plants, reaching a 
maximum of about 2.5-3 
meters. 

elliptical, to 12 cm. long, 
9 cm. wide. 

flattened. 

4 only in number, com- 
paratively large in young 
plants, much longer than 
the stipe, lying undu- 
lately along the bottom. 


P. porra 
long, 1-3 meters in young specimens, to 
20-27 meters long in adults. 


essentially spherical, 12-20 cm. in diam- 
eter. 

essentially cylindrical. 

usually 8, but sometimes 12-20, not es- 
pecially large in young plants, usually 
shorter than the stipe. 


Except for specimens from the vicinity of San Pedro and La Jolla, 
California, actual herbarium records of Pelagophycus are few. This is 
largely due to the excessive size of the plant which makes preparation 


Figure 3. Pelagophycus porra. A complete plant of medium size collected and 
photographed by W. A. Setchell at San Pedro, Calif., in 1895. Note the numerous 


blades, small holdfast and spherical vesicle giving rise to cylindrical “antlers.” 


158 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


difficult. Accordingly, our knowledge of the range is based mainly on 
observations. 

From north to south the distribution of Pelagophycus porra is repre- 
sented as follows: 

Entrance to Tomales Bay, Central California. Drifted specimen only. 

Santa Cruz Island, at 15-16 fms., between Smugglers Cove and 
Yellow Banks anchorage. This C. L. Hubbs field note is verified by 
personal communication with Dr. Thomas Sharp and Robert Fritzen 
who report individuals 30-40 ft. long, almost certainly of P porra. 
These men and Dr. North searched Anacapa Island but found no 
Pelagophycus. 


Figure 4. Pelagophycus giganteus. A medium size plant (left) and an adult plant 
(right) collected at Catalina Island by W. North. Note the large spreading hold- 
fasts, the relatively short stipes and the elliptical vesicles giving rise to flattened, 
twice-furcate “antlers.” 


Second species of Bull Kelp 159 


San Pedro, California, Setchell 1146, Dec. 1895, 12 fms. (Herb. 
WG.) 

La Jolla, California. Several herbarium specimens in Herb. Han- 
cock and Herb. U.C. Many observations by W. J. North from La Jolla 
to Point Loma at depths of 40-100 ft. 

San Diego, California. Several specimens from San Diego Bay and 
Coronado Island in Herb. U.C. An extensive bed at Point Loma exam- 
ined by Dr. North. 

Off Tijuana Slough, Baja California, at depth of 120 ft. midway 
between Point Loma and Islas Coronados. Observed by J. R. Stewart, 
early 1957. 

Islas Coronados, Baja California. Abundant at about 70 ft. on a bank 
about 6 miles southeast of Isla Coronado del Sur. Observed by J. R. 
Stewart and considered to be the largest colony known in Baja Cali- 
fornia. 

Bahia de Todos Santos, Baja California. Specimen in Herb. Farlow. 

Punta San Jose, Baja California. Observations by Dr. Michael Neu- 
shul, Nov. 18, 1957: “many plants up to 15 m. long at a depth of 18 m. 
About half of them had a peculiar bend in the stipe [at about the point 
of transition from solid stipe to hollow vesicle }?” 

Punta San Jose to Isla San Martin. Observed on several occasions 
from shipboard by Charles Martin. Some plants assuredly attached. 

Islas San Benitos. Drifted specimens observed by the writer. 

Apparently absent at Bahia Tortuga where North searched exten- 
sively between depths of 25 and 80 ft. 

Pelagophycus giganteus is known positively only from Santa Cata- 
lina Island: the Gustav Eisen: type; Dawson 5609, 5599, 8055, Silva 
4229, all from 8-15 fms. at White Cove; Silva 4261, cast ashore White 
Cove; Dawson 4794, floating near Avalon; North 7/25/61, Emerald 
Cove, 50-100 ft. (Fig. 1, 4) ; North, May 1961 (color photograph only ) 
north tip, west side, while setting anchor. 

Young Pelagophycus plants now thought by North possibly to be 
P. giganteus were observed at the outer edge of the Macrocystis bed at 
the northeast end of San Clemente Island, Jan. 28, 1957. ““The Pelago- 
phycus was noted at the bottom of a cliff estimated at 20 ft. high; the 
divers remained at the top of the cliff which was at a depth of about 
100 ft:’ Neushul reports similar observations of Pelagophycus at San 
Clemente Island on the same date: “Northwest Harbor at 50 ft.; west 
side of island at 78 ft’? A fragmentary specimen, probably of this 
species but lacking stipe and pneumatocyst, from Wilson Cove, San 


160 ~=Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


Clemente Island is in Herb. U.C.: Silva 4715, Feb. 22, 1949, pulled 
up on anchor from 12 m. 


LITERATURE CITED 


ARESCHOUG, J. E. 


1876. De tribus Laminarieis et de Stephanocystide osmundaceae (Turn.) Trev. ob- 
servationes praecursoriae. Botaniska Notiser, 3:65-73. 


1881. Beskrifning pa ett nytt algslagte Pelagophycus, hérande till Laminarieernas 
familj. Botaniska Notiser, 2:49-50. 
LEMAN, D.S. 


1822. Laminaria, in Levrault, Dictionnaire des sciences naturelles, 25:189. 
SETCHELL, W. A. 

1896. The Elk Kelp. Erythea, 4(12):179-184. 

1908. Nereocystis and Pelagophycus. Bot. Gaz. 45:125-134. 


1912. The kelps of the United States and Alaska. Jn Fertilizer resources of the 
United States. 62 Coner., 2 Sess.. Sen. Doc., 190:130-178. 


BLOOD PARASITES OF MAMMALS OF THE CALIFORNIAN 
SIERRA NEVADA FOOTHILLS, WITH SPECIAL REFERENCE 
TO TRYPANOSOMA CRUZI CHAGAS AND 
HEPATOZOON LEPTOSOMA SP.N. 

SHERWIN FE Woop 
Los Angeles City College 


INTRODUCTION 


Blood films were made from 509 native mammals during the summers 
of 1950, 1951, and 1952 at the San Joaquin Experimental Range, 
O’Neals, Madera Co., California, while the writer engaged in the 
study of the ecology of parasitic diseases, especially Trypanosoma cruzi 
Chagas. Four more samples were received in 1953, bringing the total 
to 513. This sampling program was greatly facilitated by concurrent 
studies of other investigators on the mammals and reptiles in areas 
where permanent drift traps were maintained. The prime purpose of 
this paper is to identify and enumerate the blood protozoa. 

The writer is indebted to the Pacific Southwest Forest and Range 
Experiment Station, Berkeley, for use of facilities at the San Joaquin 
Experimental Range, O’Neals, California; to Dr. E. L. Bibersteim, 
University of California, Davis, for identification of the spirochetes; 
to Dr. F D. Wood for suggestions in preparation of the figures; and to 
Dr. G. H. Ball, University of California, Los Angeles, for supervision 
of and helpful suggestions in preparation of the manuscript. 

Previous publications on trypanosomes in conenose bugs (Wood, 
1950), contaminative effect of conenose bugs (Wood, 1951a), trypa- 
nosome development in bat bedbugs (Wood, 1951b), annoyance by 
house invading conenose bugs (Wood, 1951c), occurrence of blood 
parasites in 215 range mammals (Wood, 1952a), bug and mammal 
transport (Wood and Wood, 1952), natural occurrence of Trypa- 
nosoma cruzi (Wood, 1952b), and prevalence of ectoparasites (Au- 
gustson and Wood, 1953) have dealt with related studies in this area. 
Additional details of the general environment, wildlife habitat and 
related investigations can be found in Hutchison and Kotok (1942) 
and Reppert and Green (1958). 


MATERIALS AND MrEetrHops 


Most live animals were transported in small wire cages or live traps 
to the headquarters laboratory building where a fresh blood sample 
and one blood smear were taken from each animal. The fresh blood 


161 


162 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


was examined immediately and if spirochetes, trypanosomes, or 
Plasmodium microgametes were seen, additional smears were made. 

All terrestrial mammals were marked by a standard toe clipping 
system of the station and released in the original area of capture. No 
marked land animals were recaptured. Bats were marked with alumi- 
num numbered bands from the U. S. Fish and Wildlife Service and 
some individuals were recaptured at irregular intervals, usually in new 
roosts. 

Rodents were run into wire cylinders from transport cages or live 
traps and blood taken from the clipped tail or ears before marking. 
Microtus and Peromyscus were sampled from ear veins. Dead animals 
were sampled from the heart in addition to tissue touch preparations 
from internal organs. Some samples were taken at the field plots where 
ecological studies were under way. Since this is a dry dusty area for 
most of the year, the smears show the usual contaminants of field 
prepared slides. 

Slides were stained with Jenner-Giemsa, labelled and stored for 
future study. The 1950 slides were mounted in Piccolyte and all others 
in Grubler’s Euparal Green. 

Rapid examination of the summer 1950 slides revealed 30 infec- 
tions. The recognition of pathological blood patterns and abnormal 
cells led the writer to use differential blood counts in searching for 
parasites in all other slides. A minimum of 200 leukocytes was counted 
on all slides reported here. This technique resulted in detection of 
many more parasites as noted below although some increase is at- 
tributable to seasonal sampling. 

The per cent of host cells infected and the number of parasites for 
each 100 leukocytes were recorded for each differential count. Since 
age of infection was unknown, great variation in numbers of leuko- 
cytes was expected. Differentiation of neutrophils is the same as previ- 
ously used for Peromyscus californicus (Wood, 1937). 


OBSERVATIONS 


One hundred twenty-five bacterial, protozoal, and helminthological 
blood infections were found in 513 mammals representing 19 genera 
and 22 species, a general infection rate of 24.3%. Thirty infections, 
or 13.9%, were recognized in 1950 from 215 mammals. Seventy-seven 
infections, or 34.19%, were detected in 225 in 1951 and 19, or 26%. 
were found in 73 examined in 1952 (69) and 1953 (4). 

‘Table 1 lists the number of uninfected and infected hosts. Chroma- 


Blood parasites of mammals 


14aq]18 lana} snosAwo19g 
jaquips snyojnoiupu snasAwu018g 

1y4oqg 440g snosAwo1ag 
aDsod 1A SNIIULO{IIDI $NJOLIIY 


{SOFT 
juepoy 


Y "sod ‘dan 
TOVUTAA 


snjo4npyy puv snosAwosag 104; ATeuruINs UOTdezUt 9yIse1ed pooyq [euoseas 


@ AIA I, 


PIULOF TED “YIOT YON 18 P2222]100 


snpnva1su0] s1j0jDsawu sXu0JUOposy pay 
1442q]19 tana] Snas{wmos9g 
ljaquips snipjnoiupiu snos{wo1ad 
myAog 1jhog snosXwmosag 
SNJDULOUL SNJDULOUL SnYyJDUsOL3ag 
sn4yI0 snaIUsofypa snyywUso1ag 
10jbaajs sadiosn{ DuL0jJ0a AJ 
avpsod1ADUL SNIIULO{I]DI $NJOLII J] 
snutpipudsaq snosn{ snoisaidq , 
snoyfiond snpyjod snozo.jup 


° 


OS 


Om 


wnipousnigd %|a 


eIUIOFITLD ‘sTean.Q ‘esuey [eyueutsedxgq umbeof ueg oy} wiory syeutueur Jo saytsered poolg 


J Hidv EL 


164 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


tin remnants representing nuclear fragments of Hepatozoon or Try pa- 
nosoma were found in the monocytes on smears from 4 ¢@ and 4 2 
Microtus californicus mariposae R. Rellogg,4 ¢ and 1 2 Peromyscus 
boylu boyli (Baird) and 3 ¢ Peromyscus truei gilberti (J. A. Allen). 
This means these rodents were infected at the time sampled although 
no intact parasites or trypanosome “shadows” could be found. The 
nuclear-like structure of these larger granular masses of irregular size 
but generally spherical shape is distinct from the occasional small 
azurophil granules seen in normal monocytes. 

Double infections with Hepatozoon citellicola and Trypanosoma 
microti were found in 3 ¢@ and 3 2 Microtus californicus. Try pano- 
soma microti and microfilariae were found in 1 2 Microtus. Hepa- 
tozoon muris with Trypanosoma cruzi was found in 1 ¢ Peromyscus 
true. Hepatozoon muris and Trypanosoma peromysci were found in 
1 ¢ and1 2 Peromyscus truei. Borrelia and Haemobartonella were 
found in 1 @ and Haemobartonella with Hepatozoon muris in an- 
other & Peromyscus truel. 

Special effort was made to trap Peromyscus since this rodent was 
found previously to be susceptible to infection with Trypanosoma cruzi 
from California sources (FE D. Wood, 1934, S. F Wood, 1937). General 
results for these rodents are summarized in Table 2 and indicate an 
infection rate of 20.4%. 

Figures 1 and 2 of Plate I are the first illustrations of Trypanosoma 
cruzi from a naturally infected native host in California. The typical 
regressive C form blood trypanosome (Pl. I, Fig. 1) was found in a 
375 sq.mm. smear from 1 ¢ Peromyscus truei gilberti trapped 8-IV-52 
from the garage shed near the horse barn in the headquarters area. 
The trypanosome shows a deep basophilia characteristic of active 
forms resident in the blood for some time. 

Slides from a previously reported Peromyscus truei gilberti revealed 
five typical blood forms in one thick drop and the one intact parasite 
of Pl. I, Fig. 2 (Wood, 1952b). This transitional regressive parasite 
shows increasing vacuolization and volutinization associated with the 
Se of a erouniedl body form between figs. 2 and 3 of Pl. I of 

Wood (1953). The lighter, more dispersed basophilia is a reflection 
of the increased volume of the parasite preparatory to fission. The 
flagellum and undulating membrane begin resorption as the kineto- 
plast complex becomes more rounded and the parasite folds upon itself 
assuming the rounded leishmaniform shape. These parasites probably 
drift into capillary networks where, if host resistance is overcome, they 
complete regression to the leishmaniform stage and by cell division 


Blood parasites of mammals 105 


10 pe { 


Plate I. All figures were drawn with the aid of a camera lucida. Fig. 7. Typical 
regressive C form Trypanosoma cruzi from ¢ Peromyscus truei gilberti, 8-1V-52. 
Fig. 2. Circulating regressive Trypanosoma cruzi from Q Peromyscus truei gil- 
berti, 6-VII-50. Fig. 3. C form Trypanosoma vespertilionis from tissue contact lung 
preparation of ¢ Antrozous pallidus pacificus, 22-VII1-50. Fig. 4. Trypanosoma 
peromysci from 92 Peromyscus b. boylii, 30-X11-51. Fig. 5. Trypanosoma microti 
from ¢' Microtus californicus mariposae, 26-V1-51. Fig. 6. Trypanosoma neotomae 
from 6 Neotoma fuscipes streatori, 8-1V-52. 


give rise to more progressively differentiating parasites which re- 
populate the blood stream as does Trypanosoma zapi (Davis, 1952). 
Recent studies by Régo (1956) report developmental forms of Trypa- 
nosoma cruzi in the circulating blood of white mice. This is probably 
the way long standing light blood parasitemias are maintained al- 
though limited foci of reticulotropic or myotropic forms are also 
present. 


166 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


Microtus neutrophils averaged 2.7 (6.6%) segemented forms (0.5 
to 7), 27.6 (68.5%) type (a) cells (8 to 53.5), 8.8 (21.7%) type (b) 
cells (0.5 to 23), and 1.3 (3.2%) unsegmented forms (0 to 4.5) for 
each 100 leukocytes from 23 infected voles. Six Microtus californicus 
revealing infections with Hepatozoon citellicola (P\. II, Figs. 9 and 10) 
during the differential counts showed 7.1% (3.2 to 15.5) infection of 
monocytes. Three of these voles also revealed 81.1% (66.6 to 100) 
infection of unsegmented neutrophils or 14.2% (1.8 to 37.7) of all 
classes of neutrophils. Trypanosome and hepatozoan infections stimu- 
late a shift to the younger type (b) and unsegmented neutrophils. In 
six double infections with Trypanosoma microti (Pl. I, Fig. 5) and 
Hepatozoon, trypanosomes averaged 111.6 (0.5 to 328.5), hepato- 
zoans (2 infections) 0.5, one parasite in a monocyte (5.8% ) and one 
in an unsegmented neutrophil (100% ). For seven voles with T. microti 
only, trypanosomes averaged 38.7 (1 to 111.5). 

Monocytes averaged 23.7% (10 to 35) for 14 infected Peromyscus. 
Neutrophils for these same mice averaged 0.6 (2.2%) segmented 
forms (0 to 3), 15.9 (59.3%) type (a) cells (4.5 10413) Be in(202176)) 
type (b) cells (1 to 21), and 3.2 (11.9% ) unsegmented forms (0 to 11) 
for each 100 leukocytes. In one Peromyscus boylii with Hepatozoon 
muris (Pl. II, Fig. 11) three parasites were found in monocytes 
(9.69%). In four mice with Trypanosoma peromysci (Pl. I, Fig. 4), 
trypanosomes averaged 280.7 (32 to 805). 

In three Peromyscus maniculatus gambelii (Baird), Hepatozoon 
leptosoma (PI. II, Figs. 13 and 14) averaged 3.6 (0.5 to 8) with 6.5% 
(2.3 to 10) in monocytes and 45.8% in unsegmented neutrophils (two 
mice). 

In Peromyscus truei, one mouse with Hepatozoon muris (Pl. II, 
Fig. 12) and Trypanosoma cruzi (Pl. I, Fig. 1) revealed 1.5 hepato- 
zoans in monocytes (7.8%) and two mice with H. muris and T! pero- 
mysci revealed 0.5 hepatozoans, 0 and 1, in monocytes (6.2%) and 
8.7 trypanosomes, 0 and 17.5. Two mice with 7! peromysci only aver- 
aged 24.2 trypanosomes (0 and 48.5) for each 100 leukocytes. 

The following mammals were negative for blocd parasites: 2 <. 
1 @ Spermophilus beecheyi fisheri, 14 Dipodomys heermanni tularen- 
sis, 1 & Eutamias m. merriami,1 o,1 2 Lepus c. californicus,1 3 
Lynx rufus californicus,2 3,4 9 Myotis yumanensis sociabilis, 1 
Pipistrellus hesperus merriami, 1 3,2 9 Procyon lotor psora, 1 ? 
Scapanus 1. latimanus, 1 3.7 2 Sylvilagus audubonu vallicola, 
2 3 Tadarida brasiliensis mexicana, and 13 3.7 9 Thomomys bottae 
mewa. 


Blood parasites of mammals 167 


o 9 i 14 


Plate II. Fig. 7, Microgametocyte of Plasmodium sp. from <¢ Antrozous pallidus 
pacificus, 9-VI1-51. Fig. 8. Macrogametocyte of Plasmodium sp. from ¢ Antro- 
zous pallidus pacificus, 9-VII-51. Fig. 9. Hepatozoon citellicola in monocyte from 
3 Microtus californicus mariposae, 19-IV-51. Fig. 10. Hepatozoon citellicola in 
unsegmented neutrophil from © Microtus californicus mariposae, 19-IV-51. Fig. 
11. Hepatozoon muris in a monocyte from < Peromyscus b. boylii, 30-XII-51. 
Fig. 12. Hepatozoon muris from a monocyte from < Peromyscus truei gilbert, 
8-IV-52. Fig. 13. Hepatozoon leptosoma sp. n., freed by rupture from a monocyte 
from @ Peromyscus maniculatus gambelii, 8-IV-52. Fig. 14. Hepatozoon lepto- 
soma sp. n., in a monocyte from @ Peromyscus maniculatus gambelii, 8-1V-52. 


168 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


Hepatozoon citellicola (Wellman and Wherry, 1910) 


Gametocytes (Pl. II, Figs. 9 and 10) on one stained slide from a 
California vole, Microtus californicus, averaged 9.3 gw long (8.1 to 
10.3) by 3.9 w wide (3.3 to 4.5) for 25 specimens chosen at random. 
The gametocyte nuclei of these parasites averaged 4.2 x long (3.9 to 
Ob iby-3.3 jewides(31013:9)): 

This sausage-shaped parasite exhibits a distinct limiting plasma 
membrane which pulls away from the host cell cytoplasm in some 
cells. The lightly basophilic cytoplasm appears irregularly vacuolated 
towards each end of the cytosome. The average number of meta- 
chromatic granules was 9.8 (6 to 14) for the 25 parasites, These are 
similar to volutin granules of trypanosomes and occur in the nuclear 
area or scattered through the cytoplasm. Most parasites le parallel to 
the main axis of orientation of the monocyte nucleus but occasionally 
the organism les within the crescentic cavity of the horseshoe shaped 
structure. 

The parasite nucleus is irregularly oval or rounded in shape tending 
to be shghtly elongate. The peripheral chromatin (basichromatin ) 
appears in irregular clumps or bands separated by compact or broad 
irregular parachromatin (oxychromatin) spaces and surrounding a 
less dense central core of nucleoplasm. 

Hepatozoon citellicolla is parasitic in the monocytes and unseg- 
mented neutrophils of the blood of Microtus californicus mariposae, 
San Joaquin Experimental Range, O’Neals, California. There is no 
way at present of distinguishing this parasite from that described by 
Wellman and Wherry (1910) from the California ground squirrel. 
Spermophilus beecheyi. Since the same species of ground squirrel is 
commonly associated with the foothill mammals studied here, their 
name is used in preference to Hepatozoon microti (Coles, 1914) until 
more data is available on the life cycle. 

The host cells of the blood of the vole are the unsegmented neutro- 
phils and monocytes. The cytoplasm of the unsegmented neutrophil is 
moderately basophilic, lightly vacuolated, and with none or a few 
distinct neutrophilic granules. 

The ring shaped neutrophil nucleus shows a compact chromatin 
structure similar to the smaller monocytes and larger lymphocytes. 
Where the central aperture has enlarged and the circular, band-form 
nucleus has become more labile, the parasite may extend mto the 
immature neutrophils have not yet developed their distinctive granu- 
lation. Older cells with more irregular banded nuclei show the diag- 


Blood parasites of mammals 169 


nostic granules clearly. One segmented type (a) neutrophil showed 
Hepatozoon remnants in the densely granulated cytoplasm similar 
to those seen in monocytes. 

The moderately basophilic monocyte cytoplasm has a lightly vacuo- 
lated and granular appearance. Granules are fine or coarse. The finely 
granular appearance is due to the presence of normal azurophil 
spherules similar to that found in the blood of other rodents. The 
coarse granules, numbering 22 to 27 in the cells counted, are break- 
down products of the nuclei of hepatozoans or trypanosomes as shown 
by the similar structure and staining in the nuclei of these parasites. 
The general absence of a perinuclear halo, variable shaped nuclear 
structure, and extensive less basophilic cytoplasm distinguish this cell 
from the lymphocyte. There are medium sized lymphocytes in Micro- 
tus blood with looser chromatin organization suggesting a transition 
form to the monocyte. 

The monocyte nucleus has broad or narrow dense chromatin masses 
with narrow or broad irregular parachromatin spaces. In younger 
cells it is rounded or slightly crescentic. Older parasitized cells show 
an elongate C shaped structure in which occasionally the lobes are 
folded back over each other. The labile nucleus in one instance was 
folded S like along one side of the cytosome. 

Specimens are deposited at the Department of Zoology, University 
of California, Los Angeles. 


Hepatozoon muris (Balfour, 1905) 


Gametocytes (Pl. H, Figs. 11 and 12) on one slide from Boyle’s 
white-footed mouse, Peromyscus boyliu, averaged 8.2 wu long (7.8 to 
9.3) by 3.5 w wide (3 to 4.2) for 15 specimens chosen at random. 
The gametocyte nuclei of these parasites averaged 3.5 ww long (2.7 to 
4.2) by 2.8 w wide (2.1 to 3.3). Measurement of seven parasites 
from four slides of the Gilbert white-footed mouse, Peromyscus truet, 
averaged 8 yw long by 3.4 w wide for the cytosome and 4 mu by 3.1 
for the nuclei. 

Terminal cytoplasmic caps within the parasites’ plasma membrane 
of a substance staining similarly to the protoplasmic metachromatic 
granules or nuclear chromatin distinguished this parasite from all 
others studied here. The slightly shorter and broader cytosome oc- 
casionally reveals from two to six free metachromatic granules. The 
similarity in staining reaction of the substance of the cytoplasmic caps 
to that of the metachromatic granules and nuclear chromatin suggests 


170 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


origin from these cytosomal components. In some parasites, the caps 
form a distinct broad band, 1 to 2 u thick at the center, and tapering 
away to the plasma membrane halfway from the end of the cytosome 
to the edge of the central nucleus. Parasites freed by rupture of the 
host cell and those extending into the host cell protoplasm without 
overlying the nucleus show this structure most clearly. Although 
appearing homogeneous in side view in some cells, it is more granular 
appearing in others. This narrower parasite appears more often in 
varying positions in the host cell. It was found along the convex sur- 
face of the host nucleus, within the elongate U shaped lobes, or ex- 
tending across the cytosome at an angle to the nucleus. 

The parasite nucleus is oval or rectangular with rounded corners. 
The chromatin is peripherally concentrated in broad bands with 
irregularly shaped less extensive parachromatin spaces. Some nuclei 
appear nearly solid from one surface. 

Hepatozoon muris is parasitic in the monocytes of the blood of Pero- 
myscus boyliu boylu and P. truei gilberti from the San Joaquin Ex- 
perimental Range, O’Neals, California. This parasite is probably the 
same as the parasite described by Balfour (1905) from mononuclear 
leukocytes of Mus decumanus and whose life history was reported by 
Miller (1908). Although the cytoplasmic caps were indicated by Bal- 
four (1905), Porter (1908) and Mackerras (1959), they were not 
mentioned by Miller (1908). Hoogstraal (1957) noted these “caps” 
in Hepatozoon balfourt. 

The monocyte cytosome exhibits a lightly basophilic reticulated 
cytoplasm often suggesting an alveolar pattern. There is sometimes a 
perinuclear halo. 

The monocyte nucleus shows broad homogenous, dense_basi- 
chromatin masses separated with a few irregular, parachromatin spaces 
detracting little from the general homogeneity of the structure. The 
oval or slightly indented nucleus of normal cells becomes U shaped 
in response to parasitism. 

Specimens are deposited at the Department of Zoology, University 
of California, Los Angeles. 


DraGnNosiIs 
Hepatozoon leptosoma, NEW SPECIES 
Gametocytes (Pl. I, Figs. 13 and 14) on one slide from a Gambel 
white-footed mouse, Peromyscus maniculatus, averaged 10.5 w long 
(9.6 to 11.5) by 2.1 w wide (1.8 to 2.7) for 15 specimens chosen at 


Blood parasites of mammals 171 


random. The gametocyte nuclei of these parasites averaged 4.6 ws long 
(2.4 to 6) by 1.7 w wide (1.5 to 2.4). 

The elongated, crescent shaped parasite often occupies a position 
within the concavity of the horseshoe-shaped nucleus of the monocyte 
(Pl. II, Fig. 14). It is sharply delimited by a distinctly stiff plasma 
membrane. It is narrowly elongate and crescentic with slightly baso- 
philic, more homogeneous cytoplasm containing from five to twelve 
metachromatic granules in some specimens. Although the commonest 
orientation of the parasites is enclosed in the arms of the U shaped 
nucleus of monocytes, some parasites lie under the plasma membrane 
of the unsegmented neutrophils on the concave or convex side of the 
host cell nucleus. In one instance, a parasite extended through the 
opening of the neutrophil nucleus. In some parasites with terminally 
displaced nuclei, a clearer large vacuolated area occupied the opposite 
end of the cytosome. 

In surface view, the oblong parasite nucleus consists of broad 
chromatin bands with narrow irregular parachromatin spaces (PI. II, 
Fig. 13). In some nuclei the bulk of the chromatin was distributed 
peripherally presenting a tube effect with a central nucleoplasmic 
core. In other parasites, the nuclear chromatin appeared barrel shaped 
with parachromatin scattered through the nucleoplasmic core. In one 
instance, the chromatin and parachromatin mass appeared cup-shaped. 

Hepatozoon leptosoma is parasitic in the monocytes and unseg- 
mented neutrophils of the blood of Peromyscus maniculatus gambelii 
at the San Joaquin Experimental Range, O’Neals, California. 

The host monocyte is similar to those described for Peromyscus calt- 
fornicus insignis (Wood, 1937). Most nuclei of monocytes have an 
elongate C shape enclosing the parasite. There are also numbers of 
unsegmented neutrophils with heavy ring shaped nuclei of broad basi- 
chromatin bands and narrow oxychromatin spaces crowding the para- 
site to the edge of the cytosome. 

Hepatozoon leptosoma may be distinguished from other California 
rodent hepatozoans by size, narrow elongate form and preferred host. 

Type specimens are deposited in the Department of ‘Zoology, Uni- 
versity of California, Los Angeles. 


XENODIAGNOSIS 
Xenodiagnosis was carried out for 186 mammals in 1950, 99 in 1951 
and 71 in 1952 and 1953. The 356 mammals included 253 rodents. 
91 bats, 8 rabbits, 2 raccoons and 2 domestic cats. This involved the 
feeding of 1,275 conenose bugs, mostly laboratory raised, including 


172 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


896 Triatoma p. protracta (Uhler), 321 T: rubida uhleri (Neiva), 49 
Paratriatoma hirsuta Barber, and 9 Triatoma recurva (longipes ) Stal. 
Most of these bugs were 1st and 2nd instar nymphs at time of initial 
blood meal. 

The 12 infections in Triatoma p. protracta were from three Gilbert 
white-footed mice, Peromyscus truei gilberti, from the range head- 
quarters area. Six 1st instar nymphs fed on 1 2 trapped 6-VII-50 in 
an old wood pile, three of four 1st fed on 1 @ trapped 13-VII-50 near 
buildings, and three 2nd fed on 1¢ trapped 8-IV-52 from the open 
garage shed near the horse barn showed metacyclic Trypanosoma cruzi 
in voluntary fecal droplets released from the 94th to 104th days after 
the original infective meal. 


DIscussION 


Miller (1908) used the term mononuclear for lymphocytes of the rat 
but listed the infected cells as large lymphocytes. Harris (1960) pomts 
out that only monocytes and neutrophils are phagocytic. Miller (1908) 
states that Hepatozoon is engulfed by the host cell with no effect on it 
but Balfour (1905), Porter (1908) and Laird (1959) infer active pen- 
etration of the host cell or nucleus by the parasite. If Hepatozoon ac- 
tively penetrates the host cell, it could be found in any adaptable blood 
cell. If we follow Harris (1960) and Miller (1908) as to engulfment. 
then the host cells are neutrophils or monocytes. Hepatozoon canis is 
reported from neutrophils (Laird, 1959). If the Hepatozoon gameto- 
cyte is mature when released by the internal tissue cells, engulfment 
would seem logical. In some species the red blood corpuscles are in- 
vaded and “young” gametocytes are found so this would indicate active 
penetration (Hoogstraal, 1957, Wellman and Wherry, 1910). In 
heavy Hepatozoon citellicola infections of Microtus, both monocytes 
and neutrophils contain parasites but not the lymphocytes. Coles 
(1914) reported karyolytic action on the monocyte nucleus in Hepato- 
zoon microti which was not noted here. The above observations suggest 
engulfment of Hepatozoon citellicola by the host cells in view of known 
function (Harris, 1960). Research on the cellular reactions to Hepato- 
zoon might add new insight to the functions of leukocytes. 

Analysis of the data as to season reflects the relationship of environ- 
mental temperature to intensity of blood parasitemias. This is a definite 
clue to use for retrieving material for additional study with the excep- 
tion of those mammals restricted to warmer activity periods as bats. 
Summer samples numbered 378, spring 76, and winter 59 with 74. 
23, and 20 infections, respectively, for rates of 19.5, 31.2 and 33.8 


Blood parasites of mammals 173 


per cent. If we deduct the bats from the summer total, the infection rate 
drops to 17.1%. 

Of the 162 bats examined, 37 or 22.8% were infected. These mam- 
mals offer a special study of the relationship of Trypanosoma cruzi and 
T. vespertilionis which are closely allied species as shown in PI. I, Figs. 1 
and 3. The life cycle of the Plasmodium (PI. II, Figs. 7 and 8) from 
Antrozous deserves intensive study since this bat can be maintained in 
captivity. 

The use of the differential leukocyte count is well known as a diag- 
nostic aid in clinical hematology. It was used here as an aid in enu- 
merating and finding blood parasites, and to standardize coverage of the 
smear. The same problem exists here for differentiation of lymphocytes 
and monocytes as occurs in human blood (Fey, 1958, Harris, 1960). 
Until detailed studies of normal blood of native rodents are made, it 
is not possible to relate accurately changes in the blood picture to in- 
fection. Therefore, the writer has used normal figures for Peromyscus 
californicus and Mus musculus for the comparisons indicated below. 

The structure of the rodent monocyte host cell is similar to that of 
human blood in size of the cytosome, extent and staining of the cyto- 
plasm and the usual-absence of a perimuclear halo. The dense banded 
and blotched pattern of basichromatin with little oxychromatin is more 
distinctive of lymphocyte nuclei than monocytes where the amount 
of basichromatin and oxychromatin is more equal. The extensive 
cytoplasm of these leukocytes of Peromyscus and Microtus and the 
variable nuclear shape in an irregular S, C or U form agrees with 
the differentiation for white mice (Fey, 1958). There is also con- 
siderable numerical variability in normal white mice (Snell, 1941). 
Detailed study of monocytes and lymphocytes led to the recognition 
of the parasite remnants from normal cytoplasmic granulation. Both 
hepatozoans and trypanosomes contribute to these remnants. Similar 
chromatin remnants have been reported in polymorphonuclear neu- 
trophils for Hepatozoon canis (Laird, 1959). 

Differential leukocyte counts on 12 normal Peromyscus californicus 
revealed 27% neutrophils, 7% eosinophils, 3% basophils, 9% mono- 
cytes, and 54% lymphocytes since rodent lymphoid cells with wide 
zones of cytoplasm are now considered monocytes (Fey, 1958, Harris, 
1960, Wood, 1937). Normal differential counts for white mice of 
various strains averaged 18% neutrophils, 2% eosinophils, 11% 
monocytes, and 69% lymphocytes (Snell, 1941). Nino (1929) study- 
ing the blood picture of white mice infected with virulent Trypanosoma 
cruzi found an initial lymphocytosis followed by a neutrophil leuko- 


174. Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


cytosis 25 to 35 days after infection. The high neutrophil count then 
persisted in numerical balance with the lymphocyte count until death. 
Since Nino did not separate monocytes and the age of the wild infec- 
tions here is unknown, it is not possible to compare the two sets of data. 

Tabulated differential leukocyte counts here revealed a monocytosis 
with marked reduction of neutrophils for three Peromyscus boyli with 
Trypanosoma peromysci and four P. truei, one with T: cruzi, two with 
T. peromysci, and one with T. peryomysci and Hepatozoon muris. A 
monocytosis only was found in one P boylii with Hepatozoon muris 
and one P. truei with T: cruzi. A neutrophil leukocytosis was found in 
one P. maniculatus with Hepatozoon leptosoma and two P. truei, one 
with 7. cruzi and one with T. peromysci but both also harboring Hep- 
atozoon muris. 

Differential counts for the California vole, Microtus, revealed a 
lymphocytosis with reduced neutrophils in one animal with Try pano- 
soma microti, A monocytosis only occurred in three voles with 7! mi- 
croti and one with Hepatozoon citellicola. A monocytosis with reduced 
neutrophils was found in one vole with 7! microti and two with T. mi- 
croti and H. citellicola. A neutrophil leukocytosis was found in two 
voles with 7. microti, seven with H. citellicola and two with both infec- 
tions. 

Galliard et al. (1959) demonstrated persistent low grade parasit- 
emias in white mice with chronic infections of Zrypanosoma cruzi 
long after experimental infection of the same mice with Borrelia dut- 
toni and B. crociduri. Therefore, the presence of spirochetes in the blood 
of Peromyscus truei is probably an important factor in maintenance of 
T. cruzi infections in native rodents of California. 


SUMMARY 


Blood infections of mammals with Haemobartonella (12), Borrelia 
(7), Trypanosoma cruzi (3), T. microti (17), T’ neotomae (2), 
T: peromysci (10), T: vespertilionis (16), Plasmodium (19), Hepato- 
zoon (36) and microfilaria (3) are recorded from central California. 
Hepatozoon citellicola is reported from Microtus californicus muriposae 
and Hepatozoon muris from Peromyscus boylu boylu and Peromyscus 
truei gilberti. Hepatozoon leptosoma sp. n. is described from Pero- 
myscus maniculatus gambeli. 

Differential leukocyte counts used in finding parasites revealed a 
neutrophil leukocytosis, lymphocytosis and monocytosis in infections 
with trypanosomes and hepatozoans. 


Blood parasites of mammals 175 


LITERATURE CITED 


AUGUSTSON, G. FE anp WOOD, S. F 

1953. Notes on California mammal ectoparasites from the Sierra Nevada foot- 
hills of Madera County. Bull. So. Calif. Acad. Sct., 52:48-56. 

BALFOUR, A. 

1905. A haemogregarina of mammals, H. jaculi (H. balfouri Laveran). Jour. 
Trop. Med., 8:241-244. 

COLES, A. C. 

1914. Blood parasites found in mammals, birds, and fishes in England. Parasitol., 
7:17-61. 

DAVIS, B. S. 

1952. Studies on the trypanosomes of some California mammals. Univ. Calif. 
Publ. Zool., 57:145-250. 

FEY, EF 

1958. Das weisse blutbild normaler mause des stammes agnes bluhn (AB.) Folia 
Haematologica, 75:451-470. 

GALLIARD, H., LAPIERRE, J. anp ROUSSET, J. J. 

1959. Atténuation de l’infection a Trypanosoma cruzi chez la souris blanche par 
différentes souches de Borrelia. Bulletin de la Société de Pathologie Exotique, 
52:272-276. 

HARRIS, H. 

1960. Mobilization of defensive cells in inflammatory tissue. Bact. Reviews, 24: 
3-15. 

HOOGSTRAAL, H. 

1957. Hepatozoon balfouri (Laveran) in Egyptian jerboas (Jaculus spp.) and ex- 
perimental infection of the camel tick, Hyalomma dromedarii Koch. J. Proto- 
zool., Suppl., 4:6. 

HUTCHISON, C. B. ann KOTOR, EI. 

1942. The San Joaquin Experimental Range. Calif. Agr. Expt. Sta. Bull., 663: 1- 
145. 

LAIRD, M. 

1959. Malayan Protozoa 2. Hepatozoon Miller (Sporozoa: Coccidia), with an un- 
usual host record for H. canis (James). J. Protozool., 6:316-319. 

MACKERRAS, M. J. 

1959. The haematozoa of Australian mammals. Aust. J. Zool., 7:105-135. 

MILLER, W. W. 

1908. Hepatozoon perniciosum (n. g., n. sp.) a haemogregarine pathogenic for 
white rats, with a descr-ption of the sexual cycle in the intermediate host, a 
mite (Lelaps echidninus Berlese). U.S. Hyg. Lab. Bull., 46:1-51. 

NINO, EF L. 

1929. Contribucién al estudio de la enfermedad de Chagas o tripanosomiasis 
Americana en la Republica Argentina. Buenos Aires, Imprenta de la Univer- 
sidad. 237 pp. 

PORTER, A. 

1908. Leucocytozoon musculi sp. n. a parasitic protozoan from the blood of white 
mice. Proc. Zool. Soc. London, 3:703-716. 

REGO, S. F. pe M. 

1956. Sobre encontro de formas tissulares do Trypanosoma cruzi Chagas 1909 no 


176 = Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


sangue circulante do comundongo branco (Mus musculus). Folia Clinica et 
Biologica S. Paulo, 26:17-45. 

REPPERT, J. N. anp GREEN, L. R. 

1958. Annotated bibliography of publications from the San Joaquin Experimental 
Range. Calif. Forest & Range Expt. Sta., Berkeley, Tech. Paper No. 27. pp. 
1-32. 

SNELL, G. D. 

1941. Biology of the laboratory mouse. Dover Publications, Inc. N.Y., 497 pp. 


WELLMAN, F. C. anp WHERRY, WB. 

1910. Some new internal parasites of the California ground squirrel (Otospermo- 
philus beecheyi). Parasitol., 3:417-422. 

WOOD, E D. 

1934. Natural and experimental infection of Triatoma protracta Uhler and mam- 
mals in California with American human trypanosomiasis. Am. Jour. Trop. 
Med., 14:497-517. 


WOOD, S. E 

1937. Cytological variations in the blood and blood-forming organs of white-footed 
mice experimentally infected with Trypanosoma cruzi. Univ. Calif. Publ. 
Zool., 41:389-418. 

1950. The distribution of California insect vectors harboring 7rypanosoma cruzi 
Chagas. Bull. So. Calif. Acad. Sci.. 49:98-100. 

1951a. Importance of feeding and defecation times of insect vectors in transmis- 
sion of Chagas’ disease. Jour. Econ, Ent., 44:52-54. 

1951b. Development of Trypanosoma cruzi in the bat bedbug. J. Parasitol. 37: 
330-331. 

1951c. Bug annoyance in the Sierra Nevada foothills of California. Bull. So. Calif. 
Acad. Sci., 50:106-112. 

1952a. Mammal blood parasite records from southwestern United States and 
Mexico. J. Parasitol., 38:85-86. 

1952b. Trypanosoma cruzi revealed in California mice by xenodiagnosis. Pan- 
Pacific Entomologist, 28:147-153. 

1953. Hematologic differentiation of the intramuscular developmental forms of 
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WOOD, S. EF anp WOOD, FE D. 
1952. A water cooler for transporting heat sensitive animals, especially insects. 
Bull. So. Calif. Acad. Sci.,51:108-111. 


A NEW SPECIES OF CHIGGER, GENUS EUSCHOENGASTIA 
(ACARINA, TROMBICULIDAE), WITH NOTES 
ON OTHER SPECIES OF CHIGGERS FROM 
THE SANTA ANA MOUNTAINS, 
CALIFORNIA 


RicHarp B, Loomis aND MARILYN BUNNELL! 
Long Beach State College 


Examination of available specimens of several species of mammals, 
from the Santa Ana Mountains and adjacent areas in Orange and 
Riverside counties, California, has revealed larvae of eleven species of 
chiggers. One species in the Genus Euschoengastia is described as new, 
and range extensions of more than 200 miles to the southeast from 
previous records are reported for five species. New host records are 
listed, along with notes on the attachment sites and their seasonal 
occurrence. Only the characteristics which are at variance with other 
published information are described. Selected scutal measurements 
are included for each species. 

The Santa Ana Mountains are located in both Orange and Riverside 
counties with the county line nearly bisecting the range. The moun- 
tains are ten miles east of Santa Ana and some twenty miles from the 
Pacific Ocean. The range covers nearly 400 square miles and has an 
elevation of 5600 feet at its highest point. 

The climate is affected by the position of the mountains, with south- 
westerly winds driving the rain-bearing clouds at a right angle to the 
length of the mountains. The wet season begins in December and ends 
sometime in April. Over 90 per cent of the rainfall occurs during these 
months. The rest of the year is considered the dry season. For addi- 
tional information, especially on the biota, the reader should refer to 
Pequegnat (1951). 


ACCOUNTS OF THE SPECIES 

The terminology used throughout this paper is that of Wharton, eg al. 
(1951), with few modifications. All of the measurements are in ml- 
crons. All of the larvae were studied by means of a phase contrast 
microscope, with specimens mounted on slides in polyvinyl alcohol 
L-P medium. Readers are referred to Brennan and Jones (1959) and 

1Contribution No. 1 from the Biological Sciences Department. Acknowledg- 
ment is made to the National Institutes of Health, Research Grant E-3407, for 


assistance. 


177 


178 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


Vercammen-Grandjean (1960) for certain taxonomic changes and for 
keys including the species listed below. 


Odontacarus linsdalei (Brennan and Jones) 


S) pecimens examined. Total. 17 larvae, as follows: 


RIVERSIDE CO., 244 mi. SW Lake Mathews, Dipodomys agilis, 16 
April 1957. 

Remarks. The above locality represents a range extension of 285 
airlme miles to the southeast from the type locality in Monterey 
County, California. Some variation was noted in the dorsal body setal 
formula, 8-8-8-10-12-10-6-2-2 in above, as compared to the holotype 
with 10-10-8-10-10-8-6-2-2. Averages of scutal measurements of 4 
specimens are as follows: AW-55, PW-75, SB-25, ASB-26, PSB-18, 
PNP ANIMES Se Alli 37 oP l-3 9" S272. 


Neotrombicula californica (Ewing) 


Specimens examined. Total, 59 larvae, as follows: 


ORANGE CO., Modjeska Canyon, Neotoma fuscipes, 19 Jan. 1957 
(19), Peromyscus californicus, 19 Jan. 1957 (2), 12 March 1957 (3). 
RIVERSIDE CO., 234 mi. S, 4 mi. W Corona, Tin Mine Canyon, 
Neotoma fuscipes, 17 March 1956 (26); 3 mi. SW Lake Mathews, 
Neotoma lepida, 23 Feb. 1957 (7). 

Remarks, Larvae were found attached to the external auditory canal 
and the proximal areas of the pimna. Averages of scutal measurements 
of 4 specimens from Orange Co. are as follows: AW-66, PW-80, SB-25. 
ASB-34, PSB-25, AP-28, AM-38, AL-43, PL-43, S-83. 


Neotrombicula dinehartae (Brennan and Wharton ) 


Specimens examined. lotal. 18 larvae, as follows: 


ORANGE CO., 1 mi. E O'Neill Park, Trabuco Canyon, Peromyscus 
californicus, 18 Nov. 1956. 

Remarks. This locality represents a range extension of 285 airline 
miles to the southeast from the type locality in Monterey County. 
Chiggers were found attached to the external auditory canal with a 
few scattered on the ear pinna. Averages of scutal measurements of 4 
specimens are as follows: AW-65, PW-89, SB-23, ASB-36, PSB-23., 
AP-32. AME39: AL-55, PL-62.°S-78- 


New species of chigger 179 


Miyatrombicula scottae (Brennan) 


Specimens examined, Total, 8 larvae, as follows: 


ORANGE CO., 1 mi. E O’Neill Park, Trabuco Canyon, Peromyscus 
californicus, 18 Nov. 1956. 

Remarks. This record represents a range extension of 285 airline 
miles to the southeast from Monterey County. Averages of scutal meas- 
urements of 4 specimens are as follows: AW-39, PW-44, SB-12, ASB- 
20, PSB-26, AP-22, AM-24, AL-21, PL-31, S-32. Most of these meas- 
urements are slightly smaller than those given for the holotype. 


Euschoengastia frondifera Gould 


(Figures D anp E) 
Euschoengastia frondifera Gould, 1956: 53. 


Specimens examined. Total, 34 larvae, as follows: 


RIVERSIDE CO., 8 mi. SSE Lake Mathews, Neotoma lepida, 22 Dec. 
1956 (12); 3 mi. SW Lake Mathews, Neotoma lepida, 23 Feb. 1957 
(21). ORANGE CO., Modjeska Canyon, Peromyscus californicus, 14 
Milage Ia7 Ct). 

Remarks. See remarks under FE. otophila for comparison of the two 
species. Averages of scutal measurements from 7 types (Los Angeles 
Co.) and from 4 specimens from Riverside Co., in that order, are as 
follows: AW-62, 64, PW-80, 79, SB-30, 31, ASB-22, 18, PSB-16, 12, 
AP-14, 13, AM-26, 22, AL-27, 30, PL-44, 43, S-31, 34. Larvae were 
found attached to the external auditory canal. 


Euschoengastia otophila, NEW SPECIES 


(Figures A-C, F anp G) 


Ty pe data. Holotype and 30 paratypes from 1 mile east of O’Neill Park, 
Trabuco Canyon, Orange County, California, host Peromyscus calt- 
fornicus, field number RL561118-1, collected 18 November 1956, by 
R. B. Loomis. Additional paratypes from the same locality are listed 
below. The holotype and two paratypes will be deposited in the Rocky 
Mountain Laboratory, Hamilton, Montana. Paratypes, now in the 
collection of R. B. Loomis, will be distributed to the United States Na- 
tional Museum, the University of Kansas and to other appropriate 
institutions and individuals. 

Diagnosis. Resembling Euschoengastia micheneri Gould, E. lanceo- 


180 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


PLATE I. Numbers adjacent to the nude setae represent measurements in mi- 
crons. Euschoengastia otophila new species. Figure A. Scutum and eyes. Figure B. 
Dorsal seta of the first posthumeral row. Figure C. Posterior dorsal seta. Figure F. 
Lee I, showing nude setae. Figure G. Leg II, showing nude setae. Euschoengastia 
frondifera Gould. Figure D. Dorsal seta of the first posthumeral row. Figure Jd 


Posterior dorsal seta. 


New species of chigger 181 


lata Brennan and Beck, E. frondifera Gould, E. nifi Brennan and 
Jones, and E. romola Brennan and Jones in having broadly expanded 
dorsal setae, trifurcate palpal claw and lacking tibiala III. Differs from 
E. micheneri and E. romola in having subterminala I; from E. lanceo- 
lata and E. micheneri by nude or forked lateral setae on palpal tibia; 
from E. nihi by presence of genuala HI; and from E. frondifera by 
difference in shape of dorsal setae, dorsal setal formula 2-10-10-; and 
longer humeral setae (40). 

Description of holotype. (With differences noted in paratypes in 
parentheses. ) 

Body: Small, nearly round. Color white in life, Two pairs of eyes 
widely separated and ocular plate lacking. (Body length and width 
varies from 246/265 to 379/333, average 260/270, in 10 specimens. ) 

Gnathosoma: Cheliceral base with few scattered puncta posteriorly. 
Blade slender, typical, with tricuspid cap. Palpal femoral and genual 
setae with numerous branches, dorsal and ventral palpal tibial seta 
nude or forked. Palpal tibial claw trifurcate. Tarsus with 7 branched 
setae and moderate tarsala (8). Galeal seta with 4 to 5 branches. 

Scutum: More than twice as wide as long with scattered puncta 
medially. Anterior and posterior margins of scutum sinuous; lateral 
margins concave, diverging posteriorly. Sensillae clavate, short 
stemmed and bases slightly posterior to bases of PL’s. AL’s subequal 
and much shorter than PL’s. AM slightly shorter than AL’s. Measure- 
ments of holotype as follows: AW-59, PW-76, SB-26, ASB-24, PSB-11, 
AP-12, AM-23, AL-28, PL-41, S-31. Measurements of 4 types, aver- 
ages and extremes, are as follows: AW-57 (55-59), PW-75 (74-77), 
SB-26 (23-29), ASB-21 (18-24), PSB-11 (11-12), AP-13 (12-14), 
AM-21 (20-23), AL-27 (26-28), PL-41 (39-43), S-30 (30-31). 

Body setae: (see Figs. B and C) Approximately 50 broad dorsal 
setae with numerous heavy setules. Dorsal setae of first row measuring 
44 becoming progressively shorter (36m) and thicker posteriorly. One 
pair of broad humeral setae (40m). Dorsal setal formula of holotype 
2-10-10-10-8-6-4 (dorsal formula varying from 2-10-10-10-8-6-4 to 
2-11-10-8-6-4-2). Ventral setae numbering from 34 to 40; small (23u) 
and not broadly expanded. Last two posterior rows resemble dorsal 
setae. Sternal setae (2-2) measuring 38m and 29 respectively. 

Legs: Coxae lightly punctate. ‘Tarsala II longer than tarsala I, ratio 
1.4 to 1.7. Tibialae I and II bluntly tipped. Arrangement of branched 
setae similar to E. frondifera Gould, 1956. (See Figs. F and G). Nude 
sensory setae (and measurements) as follows: Leg I, with 2 genualae 
(11-13), microgenuala; 2 tibialae (11-13), microtibiala; mid- 


182. = Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


dorsal tarsala (11-12), microtarsala, subterminala, parasubterminala 
and pretarsala. Leg H, with genuala (6-7), 2 tibialae (6-8), long 
mid-dorsal tarsala (17-19), proximal anterior microtarsala, and pre- 
tarsala. Leg IIT, with genuala (7-8) (tibiala absent). 

Specimens examined. Total, 62 larvae, as follows: Holotype and 61 
paratypes, all from ORANGE CO., 1 mi. E O’Neill Park, Trabuco 
Canyon, Peromyscus californicus, 18 Nov. 1956 (30); Modjeska Can- 
yon, Peromyscus californicus, 19 Jan. 1957 (18); 12 March 1957 (6); 
and 14 May 1957 (8). 

Remarks. This species closely resembles E. frondifera but is sepa- 
rated by the following characters. The dorsal setae are not flattened 
dorsoventrally but are broad with numerous thickened setules. The 
dorsal formula is variable in both species but E. otophila maintains a 
2-10-10- as compared to E. frondifera with a 2-14-14. Humeral setae 
are much longer, 40u as to 31 in E. frondifera. The lateral palpal 
tibial seta is usually forked and occasionally nude whereas in E. fron- 
difera it is usually nude and occasionally forked. One specimen of 
E. frondifera was found with a series of E. otophila from Modjeska 
Canyon. 

Peromyscus californicus had heavy infestations of these larvae on 
the pinae. Two mice had more than 100 chiggers on each of their ears. 


Euschoengastia terrestris Gould 


Specimen examined. Total, 1 larva, from RIVERSIDE CO., 24% mi. 
SW Lake Mathews, Dipodomys agilis, 16 April 1957. 

Remarks, This locality represents a range extension of 225 airline 
miles to the southeast of the localities in Fresno, Monterey, San Mateo, 
and Napa counties as reported by Gould (1956:56). Scutal measure- 
ments of the single specimen are: AW-54, PW-66, SB-20, ASB-21, 
PSB-21, AP-23, AM-25, AL-24, PL-36, S- lacking. The larva was 


removed from the external auditory canal. 


Euschoengastia californica (Ewing) 


Specimens examined, Total, 150 larvae, as follows: ORANGE CO.., 
1 mi. E O’Neill Park, Trabuco Canyon, Peromyscus californicus, 18 
Nov. 1956 (23); Modjeska Canyon, Neotoma fuscipes (11), Pero- 
myscus californicus (22), 19 Jan. 1957; Peromyscus californicus, 12 
March 1957 (39); and 4 May 1957 (8). RIVERSIDE CO., 234 mui. 
S, 4 mi. W Corona, Neotoma fuscipes, 17 March 1956 (5); 8 mi. 


New species of chigger 183 


SSE Lake Mathews, Neotoma lepida, 22 Dec. 1956 (19); Neotoma 
lepida (21) and Peromyscus maniculatus (23), 23 Feb. 1957. 

Remarks. Averages of scutal measurements of 4 specimens from 
Orange Co. are as follows: AW-43, PW-55, SB-18, ASB-26, PSB-13, 
AP-13, AM-32, AL-50, PL-47, S-31. The hosts were found with heavy 
infestations in the external auditory canals and the chiggers were 
grouped in clusters on the mner part of the ear pinna. 


Euschoengastia radfordi Brennan and Jones 


Specimens examined. Total, 6 larvae, as follows: 


ORANGE CO., Modjeska Canyon, Neotoma fuscipes, 19 Jan. 1957 
(5). RIVERSIDE CO., 3 mi. SW Lake Mathews, Neotoma lepida, 
23 Feb. 1957 (1). 

Remarks. Averages of scutal measurements of 4 specimens are as 
follows: AW-52, PW-75, SB-25, ASB-25, PSB-11, AP-20, AM-28, 
AL-47, PL-36, S-29. 


Euschoengastia criceticola Brennan 


Specimens examined. Total, 23 larvae, as follows: 


ORANGE CO., 1 mi. E O’Neill Park, Trabuco Canyon, Peromyscus 
californicus, 18 Nov. 1956 (9); Modjeska Canyon, Peromyscus calt- 
fornicus, 19 Jan. 1957 (2); Peromyscus californicus, 12 March 1957 
(1). RIVERSIDE CO., 234 mi. W Corona, ‘Tin Mine Canyon, Veotoma 
fuscipes, 17 March 1956 (8). 8 mi. SSE Lake Mathews, Neotoma 
lepida, 22 Dec. 1956 (3). 

Remarks. This species is widespread throughout western United 
States, from California eastward to Kansas. Averages of scutal meas- 
urements of 4 specimens from Orange Co. are as follows: AW-50, 
PW -61, SB-21, ASB-23, PSB-7, AP-11, AM-26, AL-30, PL-42, S-32. 
The larvae were attached to the external auditory canal and the adja- 
cent area of the ear pina. 


Pseudoschoengastia occidentalis Brennan 


Specimen examined. ‘Total, 1 larva, as follows: 


RIVERSIDE CO., 3 mi. SW Lake Mathews, Peromyscus maniculatus, 
23 Feb. 1957. 

Remarks. The specimen from Riverside County represents a range 
extension of 285 airline miles from previous localities in Plumas and 
Monterey counties, California (Gould, 1956). Scutal measurements 


184 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


of the single specimen are: AW-37, PW-57, SB-13, ASB-18, PSB-16, 
AP-27, AM-19, AL-15, PL-26, S- lacking. 


LITERATURE CITED 


BRENNAN, JAMES M. ann E. kh. JONES 

1959. Keys to the chiggers of North America with synonymic notes and descrip- 
tions of two new genera (Acarina: Trombiculidae). Ann. Ent. Soc. Amer., 
52(1):7-16. 

GOULD, DOUGLAS J. 

1956. The larval trombiculid mites of California. (Acarina: Trombiculidae). 
Univ. Calif. Publ. Ent., 11(1):1-116, pls. 1-26. 


PEQUEGNAT, WILLIS E. 
1951. The biota of Santa Ana Mountains. J. Ent. & Zool.. 42 (3-4) :1-84. 


VERCAMMEN-GRANDJEAN, P. H. 

1960. Introduction a un essai de classification rationnelle des larves de Trombicu- 
linae Ewing, 1944 (Acarina: Trombiculidae). Acarologia, 2(4):469-471, 1 
table. 

WHARTON, G. W. D. W. JENKINS, J. M. BRENNAN. H. S. FULLER, G. M. 

KOHLS anp C. B. PHILIP 


1951. The terminology and classification of trombiculid mites (Acarina: Trom- 
biculidae). J. Parasitol., 37:13-31. 


A NEW AMBRYSUS FROM SOUTH AMERICA 
(HEMIPTERA, NAUCORIDAE) 
Ira La Rivers 


University of Nevada, Reno 


The distinctive genus Ambrysus, occurring from north-central United 
States south to central Argentina, contains approximately 75 species 
of aquatic, vegetation-crawling bugs. The following new species adds 
another entity from the largely untapped South American reservoir 
of these distinctive insects. It may be known as: 


Ambrysus stali, NEW SPECIES 


General appearance: size medium for the genus, rather narrow, 9.0- 
9.25 mm. long, 5.0 mm. wide. Dorsum more-or-less unicolorous dark 
brown, often blackish on head, pronotum and scutellum. 

Head: Very deeply set into pronotum in center, its postero-centrum 
squarely-set into pronotum. Shiny, Eyes completely flush with head 
surface, convergent anteriorly, black, distinctly darker than head. 
Head ratios are: (1) Total length to width (including eyes), 50::67 
(75%); (2) Anterior distance between eyes to posterior distance, 
28::43 (65%); (3) Anterior distance between eyes to inner eye 
length, 28::30 (93%); (4) Posterior distance between eyes to great- 
est length of head posterior to this line, 43::15 (35%). 

Pronotum: Ratios are: (1) Width between anterior angles to width 
between posterior angles, 55::102 (54%); (2) Median length to great- 
est width, 36::100 (36%); (3) Distance between anterior and pos- 
terior angles on the same side to perpendicular distance between an- 
terior angle and baseline of pronotum, 55::52 (95%). 

Scutellum: Unicolorous, blackish; ratio of three sides, anterior and 
two laterals, 90::70::70. 

Hemelytra: Lighter in color than remainder of dorsum, reddish- 
brown to blackish. Embolia long, narrow (length to width 80::20= 
25% ), narrow anteriorly, widest at posterior end. Hemelytra narrowly 
exposing lateral connexiva and covering abdominal tip. Fully winged, 
capable of flight, wing with one “costal”’ cell. 

Venter: Blackish anteriorly, whitish posteriorly due to heavy pelt 
of hydrofuge pile. Propleura fused medially, and free from posterior 
slope of prosternum. Connexival segments smooth-edged, I-II non- 
spinose, III-IV moderately spined at latero-posterior angles. Male gen- 
ital process prominent, medium-sized, very broadly capitate with club 
much swollen. Female subgenital plate-tip of tripartite shape, the broad 


185 


186 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


central swelling flanked by two sharp tips at the postero-lateral region 
(see illustrations). 

Legs: PROLEGS: Femur incrassate; ratio of length to greatest ven- 
tral width, 72::45 (62%); Tibia and single tarsus fused into the usual 
slender, moderately curved structure. MESOLEGS: Femur 2.5 mm. 
long, ratio of length to median ventral width 70::10 (14%); Tibia 
2mm. long, ratio of length to median ventral width 60::7 (12%), two 
transverse rows of spines across distal tip, second row from tip incom- 
plete. METALEGS: Femur 2.75 mm. long, ratio of length to greatest 
median ventral width 80::10 (12%); Tibia 3 mm. long, ratio of length 
to width 100::4 (4%), three transverse rows of spines across distal 
end, the two non-terminal rows incomplete. 


Figure 1. Ambrysus stali: (A) Terminal outline of female subgenital plate, holo- 
type. (B) Male genital process, allotype. 


Type and locality data: Holotype female, allotype male and several 
paratypes from FRENCH GUIANA~—St. Laurent, October 1937, H. E. 
Hinton. Paratypes from: DUTCH GUIANA—i4 November 1942, 
16-19 September 1943, D. C. Geiskos; TRINIDAD, British West In- 
dies, 27-29 October 1938, C. J. Drake; BRAZIL—Belem, 21 September 
1937, H. E. Hinton—Manaos, September 1937, H. E. Hinton—Rio 
Candeia, 30 November 1937, H. E. Hinton. 


New South American Ambrysus 187 


Location of types: Robert L. Usinger collection (Berkeley, Califor- 
nia) —holotype, allotype and paratypes from the type locality as well 
as from Dutch Guiana and Belem and Manaos, Brazil. Carl J. Drake 
collection (Ames, Iowa) —one paratype from Trinidad, B.W.I. La Riv- 
ers collection (Reno, Nevada) —paratypes from French and Dutch 
Guiana; from Trinidad, B.W.I.; from Manaos and Rio Candeia, Brazil. 
Snow Museum collection, University of Kansas at Lawrence—two par- 
atypes from French Guiana. U.S. National Museum collection—one 
paratype from French Guiana. British Museum (Natural History )— 
one paratype from French Guiana. 

Comparisons: the new species is closely related to Ambrysus ob- 
longulus Montandon 1897. The differences between the two species 
can be summarized as: 


OBLONGULUS STALI 


Male genital process parallel-sided, Male genital process strongly capitate. 
non-capitate. Female subgenital plate Female subgenital plate tip trisinuate 
tip quadrisinuate in outline. Known in outline. Known from the Guianas, 
only from Central America. Trinidad and neighboring Brazil. 


LITERATURE CITED 


MONTANDON, A. L. 
1897. Hemiptera cryptocerata. Fam. Naucoridae.—Sous-fam. Cryptocricinae. 
Verhandlungen zo6dlogische-botanische Gesellschaft Wein, 47:6-23. 


PROCEEDINGS OF THE ACADEMY 

The Southern California Academy of Sciences met nine times during 
the fiscal year 1961-1962. Seven of these meetings were held in the 
Jean Delacour Auditorium at the Los Angeles County Museum, the 
Botany Section was held at the Los Angeles State and County Arbo- 
retum and the Annual Meeting was at Cerritos College, Norwalk. The 
section on Earth Sciences held regular meetings at various schools and 
institutions in the area. 

The following talks were presented at the monthly meetings. The 
Section on Invertebrate Zoology met September 15, 1961, and Dr. John 
Belkin of U.C.L.A. presented a talk entitled “Entomological Work in 
the Society Islands” The Section on Experimental Biology (including 
Medical Sciences) heard the talk “Biological Clocks” by Dr. Karl 
Hamner of U.C.L.A. on October 20. On November 17, Dr. Egon T. 
Degens of California Institute of Technology presented the talk ‘“‘Proj- 
ect New Valley —Geochemical Water Studies in the Sahara Desert” 
for the Section on Physical Sciences. Dr. M. Dale Arvey of Long Beach 
State College discussed “Birds of Colombia” for the Section on Verte- 
brate Zoology on December 8. The Section on Anthropology met on 
January 19, 1962, and heard “Climatic Changes as it affected early 
human populations in the Great Basin” by Dr. Homer Aschman of 
the University of California, Riverside. On February 16, the Section 
on Earth Sciences presented Dr. Richard H. Tedford of the University 
of California, Riverside, who reported on “Cenozoic Mammals from 
Australia” The Section on Botany met at the Arboretum and heard 
the talk “Plants and People of South Africa” by Dr. William Stewart, 
the director of the Arboretum, on March 16. The Section of the Junior 
Academy met April 13 and two papers were presented by junior mem- 
bers. 

The newly elected Board of Directors and Advisory Board met on 
May 4, 1962, to hear reports and elect officers for 1962-1963. 

The Second Annual Meeting was held at Cerritos College, Norwalk 
on Saturday, May 12, 1962. More than 100 persons registered for the 
Scientific Sessions, which consisted of 34 papers presented in the six 
sections, Papers presented were: 


A species problem in the genus Branchiostoma. Alfred Egge, Long Beach State 
College. 

Factors in the ability of the eastern Pacific Green turtle, Chelonia mydas agassiz1. 
to orient toward the sea from the land. Melba C. Caldwell and David K. Cald- 
well, Los Angeles County Museum. (Presented by D. K. Caldwell) 

Carotenoid metabolism in a marine fish under heterosmotic conditions. Arthur 


S. Lockley, Long Beach State College. 
189 


190 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 3, 1962 


Intranasal chiggers (Acarima: Trombiculidae) in rodents from California and 
northwestern Mexico. Richard B. Loomis, Long Beach State College. 

Variations in the benthic fauna of Alamitos Bay marina, California. Donald J. 
Reish, Long Beach State College. 

On some effects of salinity variation in the ontogenetic development of selected 
marine invertebrate animals. DeBoyd L. Smith, Cerritos College. 

A correlation of the internal anatomy of the sand dune weevils with the geologi- 
cal history of the Pacific coast. W. Dwight Pierce, Los Angeles County Mu- 
seum. 

A new cotylosaurian reptile from the Permian of southeastern Utah, and its 
bearing on the origin of the reptiles. Peter P. Vaughn, University of California. 
Los Angeles. 

Pachyostosis in desmostylid bones. Edward D. Mitchell. Jr., Los Angeles County 
Museum. 

An approach to the study of molars in hypsodont cricetines. John A. White and 
Theodore Downs. (Presented by White) 

An analysis of heart sounds in the sidewinder. Crotalus cerastes. Robert K. 
Mullen, San Fernando Valley State College. 

Induction of melanin pigment formation in Drosophila melanogaster. A. D. Keith 
and R. A. Kroman, Long Beach State College. (Presented by Keith) 

Effect of age and stress on lipemia clearance in dogs. Harry Sobel, Veterans Ad- 
ministration Hospital, Sepulveda, and Heriberto V. Thomas, St. Thomas Hos- 
pital, Burbank. (Presented by Sobel) 

Body form and swimming performance in the scombrid fishes. Vladimir Walters. 
University of California, Los Angeles. 

Call order and social behavior in the foam-building frog, Engystomops pustu- 
losus. Bayard H. Brattstrom, Orange County State College. 

A population study of the Sideblotched lizard, Uta stansburiana, in southern 
California. Robert C. Stephens, E] Camino College. 

The avifauna of a Joshua Tree woodland community. Dennis G. Rainey and 
S. G. Van Hoose, Jr., Long Beach State College. (Presented by Rainey) 

Metabolism of Chlamydomonas in sewage lagoons. Richard W. Eppley, Northrop 
Corporation. 

A systematic study of the Capsicum baccatum L. complex. William A. Emboden, 
Jr., University of California, Los Angeles. 

A study of photoperiodic inhibition of floral induction in a single leaf system. 
Michael Stanford and Richard G. Lincoln, Long Beach State College. (Pre- 
sented by Stanford) 

Pollen of the Sarcolaenaceae, Madagascar. Sherwin Carlquist. Rancho Santa Ana 
Botanic Garden. 

Aquatic adaptations of marine and salt lake fungi. C. J. Anastasion, Rancho Santa 
Ana Botanic Garden. 

The origin of sea lions. Edward D. Mitchell. Jr., Los Angeles County Museum. 

Paleontology and stratigraphy of Miocene marine deposits on San Clemente 
Island, California. Jere H. Lipps, University of California, Los Angeles, and 
Edward D. Mitchell, Jr., Los Angeles County Museum. (Presented by Lipps) 

Oligo-Miocene marine-nonmarine relationships in southern California. Richard 
H. Tedford, University of California, Riverside. 

On some behavioral responses of intertidal mollusks in the presence of star- 
fishes. DeBoyd L. Smith, Cerritos College. 

The origin and distribution of the snout beetles of the Joshua Tree National 
Monument and the adjacent region. Elbert L. Sleeper, Long Beach State 
College. 


Proceedings of the Academy 191 


Observations of problem-solving behavior and associated sound production by the 
Pacific bottlenose dolphin, Tursiops gilli. William E. Evans, Lockheed Cor- 
poration. 

The young Pacific Grey whales in Scammon Lagoon, Mexico. Robert L. Eber- 
hart, Lockheed Corporation, and Kenneth S. Norris, University of California, 
Los Angeles. (Presented by Eberhart) 


The ecology of the late prehistoric population of the San Diego coast. Claude N. 
Warren, Venice, California. 

Colossal ground figures of the California deserts. Emma Lou Davis, Los Angeles, 
California. 

Radiocarbon dates from La Jolla laboratory. Carl L. Hubbs, Scripps Institute of 
Oceanography. 

Documentary research into polynesian ethnohistory. Richard P. Gilson, Los An- 
geles State College. 


The influence of agriculture on the aboriginal social organization of the lower 
Colorado river tribes. Frederick Hicks, University of California, Los Angeles. 


Over one hundred members and guests attended the annual dinner 
meeting which featured an outstanding lecture by Dr. Laurence M. 
Klauber on “Rattlesnakes and People?’ 

The success of the meeting was due largely to the enthusiastic co- 
operation and help of the local committee, especially Dr. Henry Childs 
and Mr. DeBoyd Smith, the co-chairmen. We appreciate the support 
of Dr. Ralph F Burnight, President of Cerritos College, who presented 
the address of welcome. 

Fellows elected to the Academy were announced at the Annual 
Dinner. They are: Dr. M. Dale Arvey, Long Beach State College; Dr. 
Loye Holmes Miller, U.C.L.A. emeritus; Dr. I. M. Newell, Univer- 
sity of California, Riverside, and Miss @rechien Sibley, Tas Angeles 
County Museum. 

The 1963 Annual Mating will be held Saturday, May 18, at the 
University of Southern California. Dr. Jay M. Savage will serve as 
the local chairman. 

Dr. Peter P. Vaughn, as chairman of the Earth Sciences Section, 
reports there were three technical meetings of the Section held during 
1961-62 academic year: In November, at the University of California, 
Los Angeles, Mr. James W. Warren spoke on “Skeletal growth zones 
in living and fossil tetrapods: In December, at the Southwest Museum, 
Dr. N. Gary Lane discussed “Faunal succession in the lower Bird 
Spring formation, Clark County, Nevada” In March, at Orange County 
State College, Dr. Bayard H. Brattstrom outlined various approaches 
in “Progress towards a paleoecology of reptiles?’ Members of the Earth 
Sciences Section presented eight technical papers at the May Annual 
Meeting of the Academy at Cerritos College. 


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Seer le yiN OF THE .. 
Southern California borane 
Academy of Sciences ae 


LOS ANGELES, CALIFORNIA 


VoL. 61 OctToBER-DECEMBER, 1962 Parr 4, 


CONTENTS 


Notes on some reptiles and amphibians from western Mexico. How- 


ard W. Campbell and Robert S. Simmons 193 
Shoulder and upper arm muscles of salamanders. William A. 
me Hilton 205 


Observations on scouting behavior and associated sound production 
by the Pacific bottlenosed porpoise ( TZursiops gilli Dall). W. E. 
Evans and J. J. Dreher 217 
Contributions from the Los Angeles Museum—Channel Islands Bio- 
: logical Survey. 34. A fossil bird, caracara, from Santa Rosa 
Island. Hildegarde Howard 297 
; A new Gulf of California Periploma. Mark E. Rogers 229 
q The occurrence of Sonoraspis californica from east-central Nevada. 
; Takeo Susuki and William W. Lumsden 233 
Notes on the occurrence, observations and public health significance 
of the Pajaroello tick—Ornithodoros coriaceus Koch, in Los 
Angeles county. William G. Waldron 
’ Errata 
A new species of columbellid gastropod from Easter Island. Leo 
| George Hertlein 
E A new species of sand-burrowing marine Amphipoda from Califor- 
i nia. J. Laurens Barnard 
nad exatoray.OlNTITeO leer wie Wiles three ei GaSe cee. wulageg ee ayia a 253 


Issued December 31, 1962 


os! 


Southern California 
Academy of Sciences 


OFFICERS 
Theodore Downs . « 2). 222-0 dice en ons 5: oe ee President 
Richard B: Loomis 2.32.2 sele 424-4 ee First Vice President 
JohniA. White: 233755322 ee eee eee Second Vice President 
Gretchen Sibley . ...\. 2... i.{. 2222000 eee ee ae Secretary 
Licyd M. Martin'Ssp)). a2. eee. oe ee Assistant to Secretary 
W. Dwight Pierce 2.0.2 2 Gon: Seine ee Treasurer 
David K. Caldwell... 22. 2 05.. e el oS. na Editor 
DIRECTORS 
Russell E. Belous Charles A. McLaughlin Ruth D. Simpson 
Theodore Downs W. Dwight Pierce Peter P. Vaughn 
Richard B. Loomis Jay M. Savage John A. White 
Lloyd M. Martin Gretchen Sibley 
ADVISORY BOARD 
M. Dale Arvey Herbert Friedmann Kenneth E. Stager 
A. Weir Bell Hildegarde Howard Richard H. Swift 
J. Stanley Brode Theodore Payne Fred S. Truxal 
David K. Caldwell James A. Peters Louis C. Wheeler 
John A. Comstock Elbert L. Sleeper Sherwin E Wood 


STANDING COMMITTEES 


Finance Library 

Russell E. Belous, Chairman Dorothy E. Martin, Chairman 
Publications Membership 

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Conservation Junior Academy 

Henry Childs, Chairman Laurel Woodley, Chairman 

Publicity 
J. A. White, Chairman 


SCIENCE SECTIONS 


Anthropology Botany 

Charles E. Rozaire, Chairman Richard G. Lincoln, Chairman 
Earth Sciences Experimental Biology 

Peter P. Vaughn, Chairman Ronald Kroman, Chairman 
Invertebrate Zoology Vertebrate Zoology 

Elbert L. Sleeper, Chairman Dennis G. Rainey, Chairman 


The Bulletin is published quarterly by the Academy. Address all communications to the 
appropriate officer at the Los Angeles County Museum, Los Angeles 7, California. 


Printed by Anderson, Ritchie & Simon, Los Angeles, California 


BULLETIN OF THE SOUTHERN CALIFORNIA 
ACADEMY OF SCIENCES 


VoL. 61 OcroBEerR-DECEMBER, 1962 Parr 4. 


NOTES ON SOME REPTILES AND AMPHIBIANS 
FROM WESTERN MEXICO 


Howarp W. CAMPBELL 
University of California! 
Los Angeles 
and 
RoBERT S. SIMMONS 
Baltimore, Maryland? 


In the course of recent field work in western Mexico a number of 
specimens were obtained that furnish additional information on the 
distribution and variation of several species of reptiles and amphibians. 
Singly these do not represent sufficient data to justify publication, 
however, it was felt that the information is of sufficient interest to 
justify a combined report. No effort has been made to list all of the 
specimens collected, only those which present new information on the 
distribution, variation, or taxonomic relationships of the species will 
be considered. 

We would like to express our appreciation to Mr. James kK, Anderson 
of Berkeley, California, and-Helen Ann Campbell for their valuable 
assistance and companionship in the field, and to Mr. Sam R. Telford 
for many valuable criticisms and assistance in identifying several of 
the specimens. Dr. Robert Stebbins of the Museum of Vertebrate 
Zoology in Berkeley, and Dr. Doris Cochran of the U. S. National 
Museum kindly allowed us to use specimens in their care for com- 
parative purposes. Dr. Kenneth S. Norris of the University of Cali- 
fornia, Los Angeles, has reviewed the manuscript and made many 
suggestions. 

The following abbreviations have been used: MVZ, Museum of 
Vertebrate Zoology, Berkeley; UCLA, Department of Zoology, Uni- 
versity of California, Los Angeles. 


1Department of Zoology. 
21305 Light Street. 


193 


— SS Oe 


194 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


ANNOTATED LIstT OF SPECIES 


Syrrhophis interorbitalis Langbartel and Shannon 


One specimen, UCLA 14,599, was collected approximately 65 miles 
N. Mazatlan, Sinaloa, on July 16, 1961. This specimen is a male, taken 
while calling at 10:30 p.m. It was found in a crack in a granite boulder 
on a densely vegetated rocky road-shoulder. 

Apparently no additional specimens of this species have been re- 
ported since the type description in 1955. The call of this specimen 
was a long, drawn-out whistle, quite human in quality. Calls similar 
to this were heard from a point well north of Culiacan, Sinaloa, to 10 
miles south of Mazatlan, lending support to Duellman’s (1958) sug- 
gestion that this species may range along the Pacific lowlands of 
Sinaloa. 

This specimen agrees with the type description in all essential details 
except the interorbital ight band, which is barely discernable and 
interrupted (see Fig. 1). It is well defined and continuous in the type 
specimen. 


Syrrhophis modestus pallidus Duellman 


Four speciments of this species were collected (UCLA 14,600-3), all 
on the night of July 19, 1961. Two were collected on the side of a steep 
roadcut, 2.8 miles N. Tepic, Nayarit, and two in a similar situation 
3.9 miles N. Tepic. The species was in full chorus following a mild 
shower. 

‘Two distinct pattern types are represented by these four frogs. ‘Two 
are very light dorsally with little dark pigmentation. In one of these the 
dark pre-ocular stripe continues posteriorly through the axilla to the 
groin as a series of spots. The other two are heavily mottled dorsally 
with dark reticulations on a light background. No structural differ- 
ences could be discerned between these two pattern types, and the calls 
are apparently identical. 

The call of this species is a short “chirp:’ and was heard from the 
northern outskirts of Tepic, Nayarit, into southern Sinaloa, a linear 
distance of 110 miles. Along Mexico Hwy. 15 from Tepic to southern 
Sinaloa these frogs are usually found calling on the steep sides of road- 
cuts. Calling individuals are usually 6”-10” above the ground in the 
vegetation, or sometimes perched on boulders. West of Mexico Hwy. 15 
on Nayarit 54, the road to San Blas, the frogs are usually found calling 
from low areas along the road shoulder, and seem to prefer calling sta- 


Mexican reptiles and amphibians 195 


Photograph by Robert S. Simmons 


Figure 1. Syrrhophis interorbitalis from Sinaloa, Mexico, x3 


tions on the ground. Too few specimens were taken to show whether 
or not these apparent ecological differences might be correlated with 
taxonomically distinct forms. 


Enulius unicolor (Fischer ) 


A speciment of this snake, UCLA 14,604, was collected 9.6 miles S. 
Santiago, Colima, on August 10, 1960. It represents the first record of 
this species from the state and extends the range north westward from 
Coalcoman, Michoacan, a linear distance of 65 miles. 

It is a female, 349 mm. total length, with a small portion of the tail 
missing. Scale counts are, ventrals: 185, subcaudals: 57, and anal 
divided. This subcaudal count is considerably less than any previously 
reported for the species, the lowest being 85 reported by Boulenger 
(1896) for a female from Lake Nicaragua, Nicaragua. As only a small 
portion of the tail is missing from this specimen it is doubtful if more 
than 10 subcaudals are lacking. This would still give a count much 
lower than the previous minimum. 

The head scales are normal for the species. Dorsal scale rows are 


196 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


17-17-17, and each has a single apical pit. The dorsal color is light 
brown and each scale is outlined with a shghtly darker brown. Ventrals 
and first scale rows are pearly white. There is no evidence of the hght 
neck collar characteristic of E.. flavitotorques. 


Gyalopion quadrangularis (Ginter ) 

Numerous specimens of this snake were collected in 1960 and 1961. 
They were found rather abundantly from Navojoa, Sonora, to 40 miles 
south of Mazatlan, Smaloa. Specimens from 47.5 and 55.2 miles south 
of Culiacan, Sinaloa, show characters intermediate between the two 
subspecies, G. g. guadrangularis and G. q. desertorum, thus confirm- 
ing Dixon and Fugler’s (1959) decision to consider them as races of 
the same species. Several specimens have been deposited in the Univer- 
sity of Florida collections and in the junior author’s personal collection. 

Specimens from the southernmost localities, 20-40 miles south of 
Mazatlan, Sinaloa, show an extreme reduction of the dorsal bands, 
these becoming small spots posteriorly and occasionally absent on the 
tail. One specimen from 40 miles south of Mazatlan, which was unfor- 
tunately not preserved, had no indication of the dorsal bands except 
the nuchal band and two small spots posterior to it. 


Lampropeltis getulus nigritus Zweifel and Norris 


Two specimens were collected, both on the night of July 22, 1961. The 
first, UCLA 14,645, an adult male, measured 1053 mm. total length, 
of which the tail was 157 mm. It was collected 5.6 miles south of the 
Sonora-Sinaloa State lme on Mexico Hwy. 15 in the state of Sinaloa. 
The second, a juvenile male measuring 419 mm. total length with a 
71 mm, tail, was taken 38.9 miles south Navojoa, Sonora. Both were 
taken on the road around midnight after a slight shower. The surround- 
ing countryside was semidesert with palo verdes (Cercidium), mes- 
quite trees (Prosopis), and vegetated roadside ponds. The type and 
paratype of this subspecies were collected under similar circumstances 
(Zweifel and Norris, 1955). 

These two specimens agree in high dorsal scale count, black colora- 
tion, and other essentials with the type description of the subspecies 
(Zweifel & Norris, supra. cit.). They are the fifth and sixth speci- 
mens of this race to be reported and extend the range of the race, as 
well as the species, southward, approximately 140 miles, mto Sinaloa. 


Pseudoficimia frontalis hiltoni Bogert and Oliver, new comb. 


One specimen of this uncommon snake was collected 11.5 miles N. 


Mexican reptiles and amphibians 197 


Espinal, Sinaloa, on July 22, 1961. Our reasons for the use of the tri- 
nominal will be covered in the following discussion. 

Our specimen, UCLA 14,644, is a male, 393 mm. total length, tail 
80 mm., 20.3% of the total length. There are 34 dark blotches on the 
body and 16 on the tail. There are 155 ventrals, 49 subcaudals. The 
dorsal scales are smooth and m 17 rows. Supralabials are 7—7, infra- 
labials 8—7, the anal is divided. 

Pseudoficimia hiltoni was named in 1945 by Bogert and Oliver on 
the basis of one specimen from Guirocoba, Sonora, Mexico. It was diag- 
nosed as being closely related to P pulcherrima but with a propor- 
tionally longer tail, wider light interspaces between the dark body 
blotches, eight instead of seven infralabials, more subcaudals, and a 
double instead of a single dark line across the frontal region of the head. 

From the data they supplied their specimen could be distinguished 
easily from pulcherrima, but falls within the range of P. frontalis in 
many characters. The tail of the type of P. hiltoni, with a small portion 
missing, comprises 18.8% of the total length; Taylor and Smith (1942) 
state the range for male frontalis to be 16.2-19.1%. The tail of the pres- 
ent specimen is 20.3% of the total length. 

The light interspaces between the dorsal blotches in the present 
specimen vary from one to three scales in width, intermediate between 
hiltoni with three or more and frontalis with one. 

Pseudoficimia frontalis and pulcherrima both possess seven infrala- 
bials on both sides, hiltoni is stated to have eight. Our specimen has 
eight on the left and seven on the right. The double band across the 
frontal region found in Ailtoni is indicated in our specimen, but the 
posterior band is incomplete. Only the anterior band is present in 
frontalis. 

Our specimen has 49 subcaudal scales, closer to Ailtoni with 50+ 
than to pulcherrima with a maximum of 43, but only one more than 
the maximum of 48 reported for frontalis (Taylor and Smith, 1942). 

Due to the intermediate nature of our specimen, and the slight dif- 
ferences between the two forms, we feel that Pseudoficimia hiltoni 
should be considered a northern subspecies of Pseudoficimia frontalis, 
ranging from northern Sinaloa into southern, and possibly central, 
_ Sonora. 


Tropidodipsas occidentalis Oliver 


Four specimens of this snake were collected in July, 1961. Two, UCLA 
14,640-1, were collected 1:05 A.M. July 20, 1.2 miles south of Acapo- 
neta, Nayarit. They were found D.O.R. within 25 feet of each other 


198 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


between a pasture and a rocky hillside. UCLA 14,642 was taken 13.7 
miles north of Mazatlan, Sinaloa at 12:00 midnight on July 20. It was 
crossing the road between two rocky hillsides. UCLA 14,643 was 
collected D.O.R., 16.2 miles north of Espinal, Sinaloa at 2:45 A.M. 
July 22 

Four species of Tropidodipsas belonging to the group characterized 
by having 15 scale rows are currently recognized from the west coast 
of Mexico. One of these, Tropidodipsas philippi, is recognized on the 
basis of pattern. It has previously been considered a “‘spottted” snake 
(Boulenger, 1896 and Shannon and Humphrey, 1959). However, a 
reconsideration of Boulenger’s description in the light of Smith’s de- 
scription of a specimen from “Colima” (1943) mdicates that it differs 
from the others only in the possession of a light-colored venter. Other 
specimens of Tropidodipsas from localities near Mazatlan, the type 
locality of 7) philippi, show a tendency for the light body annuli to 
become connected ventrally by a light stripe (Shannon and Hum- 
phrey, 1959). Should this tendency be fully developed it could produce 
the light-colored venter of philippi. The remaining dark areas between 
the annuli could then be considered large rectangular “‘spots:’ Indeed, 
this is how Boulenger (supra cit.) describes them. Additional speci- 
mens from the west coast of Mexico may show that this condition is 
a geographic variation within a wide-ranging species. 

The other three species are black with w ite or yellow dorsal annuli 
which frequently cross the venter. They are distinguished by details 
of pattern and scutellation. 

Tropidodipsas occidentalis, described from a Colima specimen by 
Oliver (1937), was previously known only from the type and one 
other specimen from Michoacan (Peters, 1954). In addition to our four 
specimens there is a specimen collected near the type locality in Colima 
by T. Papenfuss in the collection of the Museum of Vertebrate Zoology 
in Berkeley (MVZ 71367). Its characteristics are included in this 
discussion. 

The other two species, 7’ malacodryas and T. freiae, were named by 
Shannon and Humphrey (loc. cit.) from single specimens from San 
Blas, Nayarit, and central Sinaloa, respectiv ely. No additional speci- 
mens of these species have been reported since the type descriptions. 

Of the five unreported specimens at our disposal, one, MVZ 71367, 
is Clearly referable to Tropidodipsas occidentalis. The other four speci- 
mens show a varied combination of characters. 

Tripidodipsas freiae, from central Sinaloa, is more similar to 7? occi- 


Mexican reptiles and amphibians 199 


dentalis from Colima and southward, than it is to 7! malacodryas or 
any of our specimens (one of which is from within ten miles of the 
type locality of 7’ freiae). A higher ventral count and more body an- 
nuli distinguish 7: freiae from T: occidentalis (freiae: 200 ventrals, 18 
annuli; occidentalis: 175-86 ventrals, 10-13 annuli). Several of the 
other characters by which freiae was originally distinguished are vari- 
able in the specimens we have examined. T° freiae is stated to have a 
nuchal band only one scale row long. This varies from one to three and 
one-half scale rows in specimens of occidentalis. The body annuli are 
one scale row long in freiae, they vary from one to four in the others. 
T: freiae is also stated to have three preoculars in the type description, 
as opposed to two in occidentalis, however, the illustration of the type 
specimen shows only two preoculars and a loreal, which is stated to be 
absent in the type description. The three-preocular condition has also 
been reported for one side of the head in a specimen from Michoacan 
(Peters, 1954). Therefore, it seems that the length of the nuchal and 
body annuli and number of preoculars are of little value in distinguish- 
ing 7! freiae from T. occidentalis. 

Tropidodipsas malacodryas is distinguished from T: freiae and 
T: occidentalis by its lower ventral count (143) and the absence of 
preoculars. Our four specimens agree most closely with these charac- 
ters than with those of occidentalis or freiae. The ventral counts range 
from 143-154, the preoculars are absent in two, and one small pre- 
ocular is present on both sides in the other two. Our specimens do not 
agree with malacodryas in subcaudal count; the type of malacodryas 
has 96 subcaudals while our specimens have 40, 41, 43 and 44. Our 
specimens also have fewer body and tail annuli than previously re- 
ported for any of the other species. 

The presence or absence of keels on the dorsal scales has previously 
been considered of taxonomic value in defining species in this genus. 
Our specimens are highly variable in extent of keeling. The type speci- 
men of T. occidentalis is the only known specimen which completely 
lacks keels on the scales. MVZ 71367 from Colima possesses weak keels 
in the anal region. In the other specimens the keels are present but vary 
from being weakly developed in the region of the vent to clearly indi- 
cated over the posterior half, or more, of the body. No correlation of 
degree of keeling with any other character could be discerned. 

In general there appear to be two trends within these snakes from 
the west coast of Mexico. From Nayarit north to central Sinaloa there 
is a general tendency for reduction in number of body and tail annul, 


200 ~=— Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


preoculars, ventrals, and subcaudals (the type of 7’ malacodryas is 
exceptional in this last character). The type of T° freiae from central 
Sinaloa is exceptional in all these characters, beg more similar to 
specimens of 7! occidentalis from Colima. On the basis of the specimens 
we have examined we are unwilling to make any decisions on the exact 
relationships of these snakes. Our decision to assign our specimens to 
Tropidodipsas occidentalis is only tentative in the absence of any com- 
prehensive treatment of the forms. There may be one wide-ranging 
highly variable species with recognizable geographic races, or two, or 
even three, distinct species which are sympatric over part of their 
ranges (Fig. 2). 

Some comments on the status of the genus Exelencophis seem appro- 
priate here. This genus was described by Slevin (1926) as Tantilla 
nelsoni on the basis of a specimen from the Tres Marias Islands off the 
coast of Nayarit. In 1942 Smith distinguished it from Tantilla and 
erected the genus Exelencophis for the unique specimen. Since that 
time the specimen has been lost and no others have been taken in col- 
lections from these islands. 

The herpetological fauna of these islands is little different from that 
of the adjacent mainland. Most of the islands’ species may be assigned 
to the same subspecies as the mainland populations (Zweifel, 1960). 
The presence of this one endemic genus in the island group has caused 
some comment by zoologists, and Zweifel compared the known charac- 
teristics of this form with the mainland genera to which it might be 
related. In his discussion of the relationship of Exelencophis to Tropido- 
dipsas Zweifel cited the considerable difference in ventral and sub- 
caudal counts of the two forms (130, 39, respectively, for Exelen- 
cophis). The specimens of Tropidodipsas now available from adjacent 
mainland areas have reduced the distinctiveness of these characters. 
One hundred and forty-three ventrals have been recorded twice for 
Tropidodipsas (type of T: malacodryas and UCLA 14,641), and two of 
our specimens have subcaudal counts of 40 and 41, thus approaching 
the condition in Exelencophis. 

Examination of the type description of Exelencophis reveals two 
additional characters which serve to distinguish it from Tropidodipsas; 
the nasal is entire, and the first infralabials do not meet behind the 
mental scale. This combination of characters is not found in other 
forms of Tropidodipsas except in the type specimen of 7! annulifera. 
Variations in the latter character are known to occur frequently in in- 
sular populations of Tantilla in Florida (Telford, in MS), and we feel 


Mexican reptiles and amphibians 201 


that additional collecting on the Tres Marias Islands and adjacent 
mainland may justify placing Exelencophis nelsoni in the genus 


Tropidodipsas. 


DURANGO 


co Elota 


4 Acaponeta 


NAYARIT 


SJ ® Tepic 


T. OCCIDENTALIS 

T. FREIAE z 

T. MALACODRYAS Guadalajara 
NEW SPECIMENS @ 


A 
® 
| 
4 


JALISCO 


Figure 2. Localities for Tropidodipsas from the west coast of Mexico. 


202 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 
Typhlops braminus (Daudin) 


A specimen of this snake was collected under a rock at the Mazatlan, 
Sinaloa, airport on December 28, 1961, by F Howard and George 
Sphon. The specimen, of undetermined sex, measures 69 mm. in total 
length. It has been deposited in the collections of the University of 
California at Los Angeles (UCLA 14,693). 

This specimen represents the first record of this species from the 
state of Smaloa and is the most northernly record for the species in 
Mexico. Its occurrence here is not surprising in view of the species’ 
world-wide dispersal through the agency of man’s activities. 


LITERATURE CITED 


BOGERT, CHARLES M. ano JAMES A. OLIVER. 
1945. A preliminary analysis of the herptofauna of Sonora. Bull. Amer. Mus. 
Nat. Hist., 83(6): 297-426. 


BOULENGER, G. A. 
1896. Catalog of the Snakes in the British Museum (Nat. Hist.). London. vol. 2. 


DIXON, JAMES R. anv C. M. FUGLER. 
1959. Systematic Status of two Mexican Species of the Genus Gyalopion Cope. 
Herpetologica, 15(3): 163-4. 


DUELLMAN, WILLIAM E. 
1958. A review of the frogs of the genus Syrrhophus in western Mexico. Occ. Pap. 
Mus. Zool., Univ. Michigan, 594: 1-15. 


OLIVER, JAMES A. 
1937. Notes on a collection of amphibians and reptiles from the state of Colima, 
Mexico. Occ. Pap. Mus. Zool., Univ. Michigan, 360: 1-28. 


PETERS, JAMES A. 
1954. The amphibians and reptiles of the coast and coastal sierra of Michoacan, 
Mexico. Occ. Pap. Mus. Zool., Univ. Michigan, 554: 1-37. 


SHANNON, FREDERICK A. ann FRANCES L. HUMPHREY. 

1959. ‘Two new species of Tropidodipsas from the West Coast of Mexico. Herpeto- 
logica, 15 (4): 217-22. 

SLEVIN, JOSEPH R. 

1926. Expedition to the Revillagigedo Islands, Mexico, in 1925, III. Notes on a 
collection of reptiles and amphibians from the Tres Marias and Revillagigedo 


Islands, and the west coast of Mexico, with descriptions of a new species of 
Tantilla. Proc. California Acad. Sci., ser. 4, 15: 195-206. 


SMITH, HOBART M. 

1942. A resume of Mexican snakes of the genus Tantilla. Zoologica, 27: 33-42. 

1943. Summary of the collections of snakes and crocodilians made in Mexico 
under the Walter Rathbone Bacon Traveling Scholarship. Proc. U.S, Natl. 
Mus., 93(3169) :393-504. 


Mexican reptiles and amphibians 203 


TAYLOR, EDWARD H. ann HOBART M. SMITH. 

1942. Concerning the Snake Genus Pseudoficimia Bocourt. Univ. Kansas Sci. Bull., 
28 (2) :241-51. 

ZWEIFEL, RICHARD G. 

1956. A Survey of the Frogs of the augusti group, genus Eleuthrodactylus. Amer. 
Mus. Novitates, 1813:1-35. 

1960. Results of the Puritan-American Museum of Natural History Expedition to 
western Mexico. 9. Herptology of the Tres Marias Islands. Bull. Amer. Mus. 
Nat. Hist., 119(2):81-128. 


ZWEIFEL, RICHARD G. anp KENNETH S. NORRIS. 
1955. Contribution to the Herpetology of Sonora, Mexico. Amer. Midl. Nat., 
54(1):230-49. 


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SHOULDER AND UPPER ARM MUSCLES 
OF SALAMANDERS 


Wintitiam A. HILTON 


Pomona College! 


This is the fourth of a series of papers recently published on the mus- 
cles of salamanders as a group. Previous to this, a series was completed 
on the skeleton consisting of twenty or more separate articles with a 
smaller number on other features of the group. 

Certain areas or regions have been discussed separately as a matter 
of convenience, but at the end of the morphological publications it is 
planned to bring all together for a general summary as well as an ap- 
plication to other topics in connection with the world group. 

M. Cucullaris, Edgeworth ’20, Rylkoff ’24, Francis ’34; Onomas- 
toideus, Funk 1827, Meckel 1828; Levator scapulae (in part), Carus 
1828; Spini-sus-scapulare, Dugés 1834; Protractor scapulae (part), 
Owen 1866; Curcularis et Sterno-cleido-mastoideus, Riidinger 1868; 
Trapezius, Mivard 1869, Drier 1901, Osawa 1902; Capiti-dorso- 
scapularis S. Cucularis, Firbinger 1873, Hoffman 1878; Petroso- 
dorso-scapularis, Perrin 1899. 

This sheet of muscle arises from cephalic-dorsal fascia and from the 
posterior dorsal surface of the skull. It is inserted on the pro-coracoid 
and scapula. 

It is supplied by a branch of the 10th cranial nerve and a part of the 
2d spinal. 

It turns or depresses the head. When the head is firmly held it pro- 
tracts the shoulder girdle. 

M. Opercularis Gaup 98, Kingsbury and Reed 1909; Levator anguli 
scapulae, Funk 1827, Riidinger 1868, Mivart 1869; Levator scapulae, 
Carus 1828, Humphrey 1871, Osawa 1902, Rylkoff 1924; Sous-occi- 
pito-adscapulaire, Dugés 1834; Protractor scapulae (part), Owen 
1866; Basi-scapularis, Fiirbringer 1873, Hoffman 1878; Protractor du 
scapulum, Perrin 1899; Levator scapuli, Driiner 1901. 

This takes origin from the operculum of the ear capsule and is in- 
serted on the anterior border of the supra-scapula. 

It is supplied by fibers from the first spinal nerve. It acts in relation 
to audition. 


1Department of Zoology. 


206 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


M. Dorsalis scapulae, Humphrey 1872, Firbinger 1873, Hoffman 
1875, Osawa 1902, Francis °34; Sur-épmeux et Sous-épineux con- 
joints, Cuvier 1800; Scapularis, Funk 1827; —Shulterblattmuskel. 
Meckel 1828; (Subscapularis)., Carus 1828; Adscapulo-humeéral, sous- 
épineux, Duges 1834; Suprascapularis, Stannius 1854, Ryhikoff 1924; 
Subscapularis, Schmidt, Goddard, van d. Hoeven 1864; Supra- and 
infra-spinosus, Owen 1866; Deltoid, Mivart 1869; Elevateur scapu- 
laire du bras, Perrin 1899. 

This arises from the dorso-lateral surface of the suprascapularis and 
is inserted on the side of the humerus. 

It is supplied by a branch of the 3d spinal nerve. 

It elevates the arm. 

M. Dorso-humeralis, Francis 1934; Grand dorsal, Cuvier 1828: 
Latissimus dorsi, Funk 1827, Carus 1828, Stannius 1854-6, Schmidt, 
Goddard and van de Hoven 1864, Owen 1866, Riidinger 1868, Mivart 
1869, Humphrey 1872, Osawa 1902, Rylkoff 1924; Vertébrato- costo- 
humeral, grand dorsal, Dugés 1834; Dorso- humeralie Furbinger 
1873. Hoffman 1873-8; Elevator dorsal du bras, Perrin 1899. 

This muscle behind the shoulder arises from fascia, it is inserted into 
the edge of the shoulder jomt with another insertion into the anterio- 
lateral border of the humerus. 

It is supplied by the dorso-humeral nerve which comes from an anas- 
tomosis of the 3d and 4th spinal nerves. 

It retracts the humerus. 

M. Thoraci-scapularis or Serratus magnus, Fiurbinger 1873, Hoff- 
man 1873; Francis 1934; Depressor anguli scapulae inferioris, Funk 
1827; Serratus anticus magnus, Carus 1828; Grosser Riickwartszieher 
der Schulter, Meckel 1828; Costo-sous-scapulaire, ou grand denteleé, 
Duges 1834; Levator scapulaore, ou grand dentelé, Dugés 1834; Leva- 
tor scapulae et serratus anticus major, Owen 1866; Serratus anticus 
magnus, Riidinger 1868; Serratus magnus, Mivart 1869, Humphrey 
1872; Retractor et suspenseur du scapulum, Perrin 1899; Thoraci- 
superiores-serrati, Rylkoff 1924. 

This is a series of slips from the first ribs, inserted on the medial or 
mesal surface of the scapula. 

It is supplied by branches from the 2d, 3d and 4th spinal nerves. 

It retracts the scapula. 


Musc.ies EsPpEcIALLY RELATED TO THE UPPER ARM 
M. Pectoralis, Schmidt, Goddard, van d. Hoven 1864; Owen 1866. 


Salamander muscles 207 


Mivart 1869, Humphrey 1872, Furbinger 1873, Hoffman 1873-8, 
Rykoff 1924, Francis 1934; Grand pectoral, Cuvier 1800; P. inf. pec- 
toralis major, Funk 1827. Grosser Buustmuskel, Meckel 1828; Ab- 
domino-corico-humeral, portion du grand pectoral, Dugés 1834; Pec- 
toralis major, Stannius 1854-6, Riidinger 1868; Pectoralis sternalis 
with p. abdominalis, Eisler 1895; Téte sternale; Fléchisseur du bras, 
Perrin 1899; Pectoralis magnus with pars anterior, Carus 1828. 

This is a large fan-shaped flat muscle over the chest region. The 
fibers converge to a tendon inserted on the back side of the humerus. 

It is supplied by the pectoral nerve from the 4th and 5th spinal 
nerves. 

It draws the arm inwards toward the body. 

M. Supracoracoideus, Osawa 1902, Rylkoff 1924, Francis 1934; 
Portio media m. pectoralis major, Funk 1827; Portion of the big breast 
muscle, Meckel 1828; Clavi-humeéral part of the great pectoral, Duges 
1834; Pectoralis secundus, Stannius 1854-6; Pectoralis minor, 
Schmidt, Goddard and van d. Hoeven 1864; Part of pectoralis, Owen 
1866; Corico-branchialis proprius, Riidinger 1868; First part of the 
coraco-brachialis, Mivart 1869; Epicoraco-humeral, Humphry 1872. 

This arises from the cartilagenous portion of the coracoid. Its fibers 
in a flat plate are inserted by a tendon on the posterior surface of the 
humerus. Its more caudal portion is covered by fibers of the pectoralis. 
It is served by the second and third spinal nerves. 

It draws the arm towards the body. 

M. Procoraco-humeralis, Fiirbinger 1873, Hoffman 1875-8, Osawa 
1902, Rylkoff 1924; Portio superior m. pectoralis majoris, Funk 1827; 
Vorwartszieher oder Heber des Oberarms Muskel, Meckel 1828; Acro- 
mio-huméral, Deltoide, Dugés 1834; Deltoides, Stannius 1854-6, 
Riidinger 1868; Subclavious, Mivart 1869; precoraco-brachial, Hum- 
phrey 1872; Adductor inférior du bras, Perrin 1859. 

This muscle arises from the pro-coracoid and is inserted near or on 
the head of the humerus. It is supplied by the N. dorsalis scapulae on 
the lateral border and by the N. supracoracoideus on the middle border. 

When the foot is on the ground it assists in forcing the body forward. 
When the foot is free from support is tends to draw the foot forward. 

M. Coraco-radialis, Stannius 1854-6, Francis 1934; Biceps brachii, 
Ridinger 1868; part of biceps, Mivart 1869; Coraco-radialis or biceps, 
Humphrey 1872; Long fléchisseur de ’avant-bras, Perrin 1899; Co- 
raco radialis, Osawa 1902. 

This muscle arises from the ventral side of the coracoid and is not 


208 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


very distinct from the last muscle. One end is inserted near the head 
of the humerus, the other is on the middle mesial side, near the proxi- 
mal end of the radius. 

It is supplied by the Supracoracoideus nerve which comes from the 
second and third spinal trunks. 

Tt draws the arm towards the body. 

M. Procoraco-humeralis, Firbinger 1873, Hoffman 1873-8, Osawa 
1902, Rylkoff 1924, Francis 1934; Portio superior m. pectoralis ma- 
joris, Funk 1827; Vorvartszierher oder Heber des Oberarms, dreiecki- 
ger Muskel, Meckel 1828; Acromino-humeéral, deltoide, Dugés 1834; 
Deltoides, Stannius 1854-6, Riidinger 1868; Subclavious, Mivart 1869; 
Precoraco-brachial, Humphrey 1872; Adductor inférior du_ bras, 
Perrin 1899. 

It arises from the precoracid portion of the pectoral arch and is in- 
serted on the head of the humerus. 

Its lateral portion is supplied by a branch of the dorsal scapular 
nerve, its middle part is supplied by the supracoracoid nerve. 

If the arm is raised from the ground the contraction of this muscle 
draws the arm forward with something of a twist of the humerus. 

MM. Coraco-brachialis longus et brevis, Humphrey 1872, Fiirbin- 
ger 1873, Hoffman 1873-8, Osawa 1902, Rylkoff 1924, Francis 1934; 
Hakenarmmuskeln, Meckel 1828; Costo-huméral, Dugés 1834; Co- 
raco-brachialis, Stannius 1854-6, Riidinger 1868, Schmidt, Goddard 
and van d. Hoeven 1864; Second part of the coraco-brachialis, Mivart 
1869; Coraco-brachialis longus, superficialis, and brevis, Eisler 1895; 
Téte coracoidienne; flechisseur du bras, Perrin 1899; Deductor du bras, 
Perrin 1899. 

The long head arises from the postero-lateral border of the coracoid 
a little towards its dorsal side. There is a fleshy insertion along the distal 
half of the humerus on the posterior side. 

The short head arises from the proximal half of the humerus on the 
posterior side. 

The muscle is supplied by the coraco-branchial nerves from the 
ramus superficialis of the N. Branchialis. 

Both parts of the muscle draw the arm cadually; if the foot rests on 
the ground the body is propelled forward. 

M. Subscapularis, Funk 1827, Stannius 1854-6, Riidinger 1868, 
Mivart 1869, Osawa 1902, Francis 1934; Sous-scapulaire, Dugeés 
1834; Coraco-brachialis or Subscapularis, Humphrey 1872; Subcoraco- 
scapularis, Fiirbinger 1873, Hoffman 1873-8, Rylkoff 1924; Subscap- 


Salamander muscles 209 


ularis (dorsal portion) Subcoaracoscapularis, Eisler, 1895; Adductor 
supérieur du bras, Perrin 1899. 

This arises from the dorsal surface of the pro-coracoid and is inserted 
on the dorsal surface of the humerus. 

It is supplied by the subscapular nerve from the 3d spinal. 

It draws the arm backwards. If the hand(foot) is resting on the 
ground it helps advance the body. 

M. Anconaeus, Firbinger 1873, Hoffman 1873-8, Rylkoff 1924, 
Osawa 1902, Francis 1934; Anaconaeus internus et externus, Funk 
1827; Triceps brachii extensor, Carus 1828; Strecker des Vorderarms, 
Meckel 1828; Scapulo-humero-olecranien, or triceps, Dugés 1834; 
Streckmuskelmasse, Stannius 1854-6; Triceps brachii S. anconaeus, 
Riidinger 1868, Mivart 1869; Triceps and coraco-olecranalis, Hum- 
phrey 1872; Extenseur de l’avant-bras, Perrin 1899. 

This arises from the shoulder girdle and humerous by four heads 
which unite to be inserted on the olecranon process of the ulna. 

It is supplied by the extensor nerves. 

Its function is to extend the forearm. 

Head A, arises from the scapula near the posterior margin of the 
gelenoid fossa. This part was called M.a.s. medialis, by Firbinger 
1873; Hoffman 1873-8, Rylkoff 1924, Francis 1934; Anconaeus lon- 
gus, Eisler 1895; Téte antérior, Perrin 1899; Anconaeus caput scapu- 
lare, Osawa 1902. 

Head B, arises by a long tendon from the inner, bony part of the 
coracoid. It is rather small and joins the last division at about the 
middle of the humerus. It has been called, M.a. coracoideus, by 
Furbinger 1873, Hoffman 1873-8, Eisler 1895, Rykoff 1924, Francis 
1934; Coraco-olecranonalis, Humphry, Téte postérieur, Humphrey 
1872; Anconaeus caput coracoideum, Osawa 1922. 

Head C, arises from the anterio-lateral surface of the humerus, to 
later join the other parts of the muscle mass. It has been called, 
M. anconseus humeralis lateralis, by Firbringer 1873, Hoffman 
1873-8, Rylkoff 1924; Anconaeus caput humerale mediale, Osawa 
1902; With head D=Teéte profunde, Perin 1899. It arises from the 
lateral surface of the humerus. 

Head D, arises from the surface of the humerus. It has been called, 
M. Anconaeus humeralis medialis by Fiirbringer 1873, Hoffman 
1873-8, Rylkoff 1924, Francis 1934; Aconteus caput humerale lat- 
erale, Osawa 1902; With head C, Téte profunde, Perrin 1899. 

Tt arises from the surface of the humerus. 


210 + Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


M. Humero-antibrachialis, Rylkoff 1924, Francis 1934; Brachialis 
medius, Funk 1827; Flexor brachu, Carus 1828; Oberer Beuger des 
Vorderarms, Meckel 1828; Humeroradial s. biceps. Duges 1834; Hu- 
mero-radialis, Stannius 1854-6; Osawa 1902; Biceps, Schmidt, Good- 
ard, and van d. Hoeven 1864; Biceps and brachialis internus, Owen 
1866; Brachialis internus, Riidinger 1868; Part of the biceps, Mivart 
1869; Brachialis anticus, Humphrey 1872, Eisler 1895: Humero-anti- 
brachialis inferior, or brachialis inferior, Fiirbringer 1873, Hoffman 
1873-8; Court fléchisseur de l’avant bras, Perrin 1899. 

This muscle arises from the flexor side of the humerus. It is inserted 
on the proximal end of the radius. 

It is supphed by one or more branches from the superficial division 
of the brachialis nerve. 

It bends the elbow. 

CoMPARISONS 


The preceeding general account follows Francis 1934, and is largely 
based upon the muscles of Salamandra as a standard for comparison. 
In the various groups there are variations in position and in size—es- 
pecially with certain muscles. In general the pectoralis is constant in 
form and appearance and the dorso-humeralis is about the same in all 
groups. The dorsales scapulae varies in size, but the central part of the 
shoulder muscles is convenient to start from in determing the others. 
The procoracohumeralis and the supracoracoideus are seldom dis- 
tinctive. 

The greatest differences are found in the cucullaris and the so-called 
opercularis or levator scapulae. As Dunn *41 pointed out, that called 
the opercularis has really two different elements. When in Salaman- 
dridae, Hynobiidae and Amblsttomidae it is all or part of the levater 
scapulae; while in Plethodontidae it is part of the cuccularis. 


PROTEIDAE 

In Proteus, the procoraco-humeralis is especially long and slender. 
At first sight it seems continuous with the abdomino-hyoideus, being 
separed from it by a very slight line of connective tissue. The cucul- 
laris is a long and slender filament and there is a very delicate levator 
scapulae and also there are more posterior fibers which connect the 
scapula with the body wall. A very delicate omo-arcualis (discussed 
in another paper) is present. 

Necturus is similar to Proteus but the procoraco-humeralis is not 
so long in proportion to other parts. the omo-arcualis is more evident 
and like Proteus the levator scapulae is incompletely differentiated 


Salamander muscles 211 


from the dorsal trunk muscles. The cucularis is more evident and of 
two divisions. 


SIRENIDAE 


In Siren the levator scapulae is quite unusual. A single slip extends 
from the pharynx to be inserted on the cartilagenous suprascapula. I 
was not able to determine whether this was the case in Pseudobranchus 
in the material at hand. 

Siren has a well developed omo-arcualis as does Pseudobranchus as 
well. The cucullaris has two divisions in Siren, but this and other 
muscles of the region are not well developed. 


AMPHIUMIDAE 


The muscles of the shoulder girdle are greatly reduced. There is no 
levator scapulae, the cucullaris is a single rather slender band. Includ- 
ing the pectoralis there are two muscles of the region ventrally and 
three dorsally. In a larva of 50 mm, the pectoralis is evident with slight 
signs of two or three other muscles in the region. 


CrYPTOBRANCHIDAE 


The levator scapulae is incompletely separated from the trunk 
muscles. Dorsally the scapula is connected to the wall by muscle fibers. 
The cucullaris, a single band arises from the skull and dorsalfascia. 


HyYNOBIIDAE 


In Salamandrella, the levator scapulae muscle is single and forms 
the opercular muscle. The cucularis is of two slips, the upper is the 
larger. They arise from the skull and dorsal fascia. 

The general position of the muscles in this form differs from many 
other salamanders. A part of the suprascapularis muscle may be seen 
from the outer surface. 


AMBYSTOMIDAE 


The levator scapuli which forms the opercularis muscle differs in 
size in different species. It is frequently attached to a great part of the 
cephalic border of the scapula, then narrows down to a small insertion 
on the opercular plate. In some cases fibers are attached to skull parts 
near. In larval-like forms such as Siredon it is not clearly separate 
from the lateral muscles of the body. Some forms also have this con- 
dition, partly due to the fact that transformation is quite recent. In one 


212 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


specimen of Rhyocotriton which looked like an adult externally, this 
muscle of two different parts ran to the neighborhood of the ear plate, 
but all its fibers were attached to nearby bone and none to the plate 
itself, which later however, was more like a columella than an oper- 
culum. 

In this family the cucullaris was usually of two parts, sometimes 
three. 

In Ambystoma maculatum larva of 45 mm length there were dis- 
tinct opercular muscles, one seemed to be the cucullaris, but the leva- 
tor scapulae muscles were not distinct, a more or less characteristic 
larval condition. 


SALAMADRIDAE 


The levator scapulae has two slimps in some forms, one only in 
others. In Diemyctylus viridescens, 1 found two slips for this, im 
Taricha examined there was one broad one, in Pleurodellides two, in 
Tylops two and in a specimen of Triturus cristatus, one. 

The cucullaris attached to the skull and fascia in the head region 
frequently had two slips. 


PLETHODONTIDAE 


The levator scapulae arises from the base of the skull and is in- 
serted on the forward edge of the scapula. It is frequently, or is near the 
portion of the cucullaris muscle which is attached to the operculum. In 
most cases it is single, but it may have other fibers. The cucullaris is 
usually of three parts. One of the deeper or lower portions forms the 
opercularis muscle. Often parts of this slip or portions near are attached 
to the skull. In all the genera examined the conditions were much like 
those described. In Hydromantes, a powerful superficial branch of the 
cucullaris runs in the same direction as the deeper opercular muscle, 
but is very much longer and takes origin from the skull just back of 
the orbit. Typhlomolge differs from most in having the larval conditicn 
of the operculum muscle and a poorly developed levator scapulae 
which last consists of a few weak strands well separated. 

In the larval condition, the so-called operculis muscle does not end 
on the free plate of bone in the ear and the levator scapulae muscle is 
not well differentiated from lateral body muscles—a condition lke that 
of Necturus. 


Salamander muscles 213 


Plate I: 1. Proteus shoulder girdle from the front. 2. Same from the side, cephalic 
end at the right, dorsal side above. 3. Necturus, side view of shoulder muscles. 
4. Crytobranchus, same from the side. 5. Amphiuma, breast muscles from the 
front, right side. 6. Amphiuma, lateral shoulder muscles, left side. 7. Siren, side 
shoulder muscles. 8. Salamandrella, side view of shoulder muscles. 9. Same as the 
last with cucullaris removed. 10. Ambystoma maculatum adult, side shoulder 
muscles. 11. The last with superficial muscles removed. 


214 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


Plate IT: 12. Side shoulder muscles of Diamectylus viridescens. with the superficial 
muscles removed. 13. D. viridescens, shoulder muscles in place. 14. Taricha torosus, 
lateral shoulder muscles with the superficial ones removed. 15. Lateral shoulder 
muscles of Desmognathus fuscus. 16. The same with superficial muscles removed. 
17. Lateral shoulder muscles of Psepdotriton. 18. Deeper muscles of the last 
19. Lateral shoulder muscles of Hydromantes platycephalus. 20. Outer view of the 
muscles of breast and leg of Salamandra, after Francis. 21. Inner view of shoulder, 
breast and upper leg muscles of Salamandra, after Francis. 


Salamander muscles 215 
INDEX TO ILLUSTRATIONS 


A., Abdominal muscles; ACC Anconaeus coracoideus cucullaris; 
AHL, Anconaeus humeralis; AHM, Anconaeus humeralis medialis; 
C, Cucullaris; CBL, Coraco-brachialis longus; CR, Coraco-radialis; 
DH, Dorso-humeralis; DS, Dorsalis scapuli; HA, Humero-anti- 
brachialis; LS, Levator scapuli (operculus); O, Omo-arcualis; P, 
Pectoralis; PH, Procoraco-humeralis; S, Supra-coracoideus. 


SoME PERTINENT REFERENCES 

DUNN, E. R. 

1940. The “opercularis” muscle of salamanders. Jour. Morph., 69(2) :207-15. 

IWAIMOIN,, TE Jal, Wet 

1936. The myology of salamanders with particular reference to Dicamptodon en- 
satus. Jour. Morph., 19(2):31-75. 

EMERSON, E. T. 

1905. The general anatomy of Typhlomolge. Proc. Boston Nat. Hist. Soc., 32:43- 
76. 

FRANCIS, E. T. B. 

1934. The anatomy of the salamander. Oxford Press. 

UNI, J 

1938. Morphogenesis of the cranial muscles of Ambystoma punctatum. Jour. 
Morph., 63:531-587. 

SMITH, G. M. 

1927. Detailed anatomy of Triturus torosus. Trans. Roy. Soc. Canada. 3d ser. 21, 
pp. 451-484. 

WILDER, H. H. 

1891. Contribution to the anatomy of Siren lacertina. Zoologische Jahrbiich, Abte. 
Morphologie, 4:653-696. 


ADDITIONAL BIBLIOGRAPHY OF 
EarLy PuBLICATIONS DEALING WITH THE 
NoMENCLATURE OF SALAMANDER MUSCLES 
CARUS, C. G. 
1828. Tabulae Anatomium Comparativam illustrantes, Pars 1. Lipsae. 
CUVIER, G. 
1800. Lecons d’anatomie compareée. Paris. 
DRUNER, L. 
1901. Studien zur Anatomie der Zungenbein-, Keiemenbogenund Kehlpopfmuskel 
der Urodelen, I Theil. Zoologische Jahrbiich, Abteilfiir Anatomie, vol. XV. 
DUGES, A. 
1834. Recherces sur lostéologie et la mycologie des Batraciens a leurs different 
ages. Pt. II, p. 135, Paris. 
IRIS ILI, 122 
1895. Die Homologie der Extremiaten. Abhundlung der naturfulschende Ge- 
schellschact zu Halle., 18:87-348. 
FRANCIS, E. T. B. 
1934. The anatomy of the Salamander. Oxford Press. 


216 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 
FUNK, A. EF 


1827. De Salamandrae terrestris vita, evolutione, formatione. Tractatus, Berolini. 

FURBINGER, M. 

1873. Vergleichende Anatomie des Brustzhulterapparates. I Theil. Jena Zeits- 
chrift, 7:237-320. 

GAUP, E. 

1901. Zur Kenntnis des Primordialcraniums der Amphibien und Reptilien. Ver- 
handlungen anatomische Gesellschaft, 5 Versammlung in Miinchen, pp. 114- 
120. 

HOFFMAN, C. K. 

1873-8. Amphibia. In Bronn’s Klassen und Ordungen des Thierreichs. Bd. VI, Abt. 
II. Leipzig & Heidelberg. 

HUMPHREY, G. M. 

1871. The muscles and nerves of the Cryptobranchus japonicus. Jour. Anat. and 
Physiol., London, vol. VI, pp. 1-61. 

KINGSBURY, B. F, anv H. D. REED. 

1908. The columella auris in Amphibia. Anat. Rec., 2:81-91. Also Jour. Morph., 
20 (for 1909) :549-620. 

MECKEL, J. F 

1821-33. System der vergleichenden Anatomie. 6 parts Halle. French trans., 10 
vols., Paris, 1828-1838. 

MIVART, ST. G. 

1869. Notes on the Myologie of Menopoma alleghaniese. Proc. Zool. Soc. London, 
1869: 260-278. 

OSAWA, G. 

1902. Beitrage zur Anatomie des japamischen Reisensalamanders. Maitteilung 
medikament Fakultat Japan Universitat, Tokyo, vol. V, pp. 221-410. 

OWEN, R. 

1866-8. Comparative Anatomy and Physiology of vertebrates, 3 vol., London. 

PERRIN, A. 

1899. Contribution a l’étude de la myologie et l’ostéologie comparée du membre 
antérior chez un certain nombre de Batrachiens etde Sauriens. Bulletin Sct- 
ence France, Belgique, T. XXXII, pp. 220-82. 

RUDINGER, N. 

1868. Die Muskeln der vorderen Extremitaten der Reptilian und Végeln. Natur, 
Verhandlungen Hollandsche Maatschappij. d. Wetensch. te Haarlem, pp. 
1-187. 

RYLKOFF, H. 

1924, Die Entwicklung der Schultermuskeln be urodelen Amphibien. Zeitschrift 
fiir wissenschaftliche Zoologie, 122:116-171. 

SCHMIDT, F J. J.. GODDARD, Q. J.. ann VAN DER HOVEN, J. 

1864. Anateekenigen over de anatomie van Crytobranchus japonicus. Natuur- 
kunde Veh. v.d. Hollandsche Maatschappij v. Wentensch. te Harlem, 2 verz. 
19 deel, pp. 1-66. 

STANNIUS, H. 

1854-6. Handbuch der Anatomie der Wirbeltiere, 11. Aufl., Buch II. Die Am-- 


phibien. 


OBSERVATIONS ON SCOUTING BEHAVIOR AND 
ASSOCIATED SOUND PRODUCTION BY THE 
PACIFIC BOTTLENOSED PORPOISE 
(Tursiops gilli Dall)" 


W. E. Evans anp J. J. DREHER 
Lockheed California Company 


Burbank, California 


INTRODUCTION 


The sounds produced by cetacea and the possibility of their use for 
communication has long been of interest to scientists. As early as 1935, 
the sound production of cetacea in the Black Sea was observed and the 
hypothesis offered that these sounds could be communicative in nature 
(Tomulin, 1955). Since this time the literature pertaining to this hypo- 
thesis has become voluminous (see, for example, Tomilin, 1955; 
Frazer, 1947; McBride and Hebb, 1948; Wood, 1953; Lilly, 1961; 
Norris and Prescott, 1961; and Dreher, 1961). Although some of these 
contributions on the subject of cetacean sound production and com- 
munication have been based on observations of wild populations, the 
majority of the more recent work has been based on captive animals. 

The study of the communicative behavior of captive animal popula- 
tions, especially cetacea, has many advantages over field observations. 
In working with cetacea, which are normally quite mobile, the re- 
stricted environment affords better control of observational conditions 
as well as the possibility of long-term constant monitormg of sound 
production and concurrent behavior. On the other hand, one is limited 
to working with those species which readily adapt to captivity, and 
most certainly all behavior of the animals under study is modified by 
the restriction. An ideal solution to this quandry would be the capabil- 
ity to make observations of sound production and concurrent behavior 
on the same genus and/or species of cetacea both in captivity and in its 
natural habitat. The difficulties of this approach are obvious. 

With the cooperation of Marineland of the Pacific oceanarium in 
the use of its facilities and animals, and the availability of a 50-foot 
oceanographic vessel specially adapted for sound recording at sea, it 


1These studies were aided by Contract Nonr (N.R. 301-604) between the Office 
of Naval Research, Department of the Navy, and the Lockheed Aircraft Corpora- 
tion, California Division. 


218 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


has been possible to start compiling sound data on some genera of 
Delphinidae under both captive and natural conditions. At present, 
data have been collected on Tursiops truncatus (Montagu), Tursiops 
gilli (Dall), Lagenorhynchus obliquidens (Gill), Delphinus bairdi 
(Dall), and Globicephala scammoni (Cope). 

The purpose of this paper is to discuss observations of the sound 
production of the Pacific bottlenose porpoise ( Tirsiops gilli) and some 
concurrent behavior made during a cruise to Scammon’s Lagoon 
(28° N, 114° W), Baja California del Sur, Mexico. In addition, some 
comparison will be made between these data and data collected from 
T. truncatus im captivity. 

The primary purpose of the biological cruise to Scammon’s Lagoon 
from 22 January to 8 February, 1962, was to study in general the 
behavior of the California gray whale, Eschrichtius glaucus (Cope) 
and specifically to make observations of sound production. However, 
the common occurrence of 7’ gilli in the lagoon afforded excellent 
opportunity for frequent observation of the behavior of these animals 
also. 


FACILITIES AND INSTRUMENTATION 


The Lockheed California Company oceanographic research vessel 
Sea Quest is a 50-foot, diesel-powered, converted salmon trawler which 
has been especially fitted for accomplishing a wide range of oceano- 
graphic missions. From the standpoint of underwater sound recording 
one of the more important modifications has been the installation of a 
battery-powered generator which can silently provide adequate stable 
115 volts, 60 cycle, power for periods up to 4 hours of continuous 
“silent ship operation?’ 

The recording system used consisted of an AN/PQM-A1 Noise 
Measuring Set and an Ampex 351, 2 channel, Tape Recorder. Channel 
1 was used as the data channel and channel 2 as the voice channel in 
both systems. The hydrophones for the system were mounted in bo- 
tom-tripods. The hydrophone positioned alongside the ship was des- 
ignated as Hyd. #1 and the distant (200’) hydrophone as Hyd. #2. 
(Fis. te 

The frequency response of the Ampex 351 at 744 imch/sec is flat 
from approximately 30 cps to 16,000 cps and at 15 inch/sec flat (+2 
db) from 20 cps to 20,000 cps and down 11 db at 29,000 cps. The 
AN/PQM-A1 is flat from approximately 40 cps to 40,000 cps. Due to 
the limitation on the lower end of the hydrophone system (40 cps), 
and on the upper end of the tape recorder, 16 kcps-20 kcps depending 


219 


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220 ~=Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


on speed, the flat response of the complete system, +2 db can be 
assumed to be 40-20.000 cps. 


PROCEDURE 


In attempting to obtain sound production in reference to some known 
condition, a portable floating barrier was constructed which hopefully 
would divert the animals’ direction of movement and provide a good 
“sonar” target if the animals under study were using some form of 
echolocation. The barrier selected consisted of 15 air-filled aluminum 
tubes, 2 inches in diameter and 15 feet long. Each tube (spar buoy) 
was weighted with chain, and when placed in the water 12 feet were 
below the surface. During the particular phase of the cruise under 
discussion in this paper, the individual units of the barrier were placed 
at 50-foot intervals, with one end anchored and the other end moored 
to the ship. When placing the barrier an attempt was always made to 
cross a channel which was, or had been observed to be, in use by the 
animals. The hydrophones were put in place as close to the barrier 
as was possible with available cable. 


Tursiops gilli VOCALIZATIONS 


During the period of observation in the lagoon, T. gilli were sighted and 
listened to several times. The animals occurred in groups of 2 to 5 and 
approached up to 100-200 feet from the ship. During the majority of 
contacts, the only sounds heard were echolocation pulses at various 
repetition rates. These signals were very similar, if not identical, to 
those produced by 7! truncatus in captivity. Maximum range of sound 
contact was approximately 500 yards from the ship. 

On February 1, 1962, at 1600 hrs, the ship was located in Piedra 
Channel, at 27°41’8” N, 114°8’ W. Hydrophone #1 was in position, 
and the barrier was streamed between the ship and a shoal, a distance 
of approximately 250 yds. Hydrophone #1 was at a depth of 24 feet. 
At approximately 1650 hours, Wendell Tripp, the assistant master of 
the ship, sighted five Tursiops gilli moving slowly up the channel from 
the east toward the barrier at a range of about 500 yards. The sound 
gear was switched on to monitor. Very slow echolocation clicks were 
heard. At a distance of approximately 400 yards, the group, still echo- 
locating, moved over to shoal water (4-9 feet deep) and schooled into 
a tight group. After about five minutes of ver y sparse sound activity. 
a single animal left the group and headed toward the barrier. This 
animal made a “sonar” run along the barrier, turned, and headed back 


Porpoise behavior 221 


| 
Pia 
a i ies es 
pe ial pe 
ey aaa a 
I sareehsh abe sole es 


~ 
= 


> —\= 
7 
7 
a 
Se ee 
ee 


ae 


—— 
SS 
— 
~~ 
——S 


SHOAL 


ISLA PIEDRO 


Figure 2. Schematic diagram of the location of research vessel, barriers and ani- 
mals during vocalization incident discussed in text. 


222 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


to the group. At this time, whistle activity started. This represented the 
first time on the cruise that prolonged whistling activity was observed. 
The single animal runs on the net, followed by whistling, were re- 
peated 2 to 3 times then the group left the shoal and proceeded cau- 
tiously up the channel. After passing the barrier, sound activity ceased 
and the animals went west and through the barrier, whereupon visual 
contact with the animals was lost. A schematic diagram of the location 
of ship, barrier and animals during this incident is presented in 
Figure 2. 

It is interesting to note that on the several other occasions of Tursiops 
sighting, only intermittent echolocation signals were recorded. How- 
ever, the placement of a sound reflecting barrier across a normally 
non-restricted channel did elicit some form of cautious behavior ac- 
companied by what appeared to be scouting. Also of interest is the 
sequence of echolocation scanning behavior-whistling-scanning be- 
havior, in this order. Although it is possible that similar events occur in 
captivity with 7) truncatus, the small confines of the usual holding 
facility make specific comments on groupings and orientation of 
animals difficult if not impossible. 


T. gilli-T. truncatus: TONE ConTOUR COMPARISON 


Having obtained a relatively good sample of 7: gilli whistles, a com- 
parison was made between these signals and similar vocalizations pro- 
duced by 7! truncatus in captivity. To aid in this comparison, a simple 
notation system was developed to symbolize the various whistle con- 
tours (Fig. 3). In each of these symbols the abscissa represents time 
and the ordinate represents frequency or pitch. The contours used in 
the following comparison, it should be poimted out, were transcribed 
by ear from sound tapes slowed down four times. This is a relatively 
easy and accurate process after several preliminary auditions of the 
tape. While linguistics recognizes some eight different forms of tone- 
modulated language (Pike 1948), involving pitch change, register 
change, durations, relative beginning and end registers, and various 
combinations, these analyses were assumed to follow the pattern of a 
Class I language wherein lexical significance is denoted by changes 
in pitch shift only. This does not preclude sandhi (linking) accom- 
modation, or semantic contour profiles, but does relegate them to 
second or higher order determinants. 

Earlier analyses of porpoise whistles (Dreher, 1961) dealt with 


Porpoise behavior 


bo 
to 
Ov 


FREQUENCY 


TIME 


Figure 3. Typical example of the notation system used in this study to symbolize 
cetacean whistle contours. 


some statistical aspects of Tursiops truncatus signals as produced in a 
relatively restricted tank with nine interacting animals over several 
situations of feeding, play, and rest. The question naturally arises: 
Do other species (or genera) use the same whistle contours, and if not, 
what differences are noted? A partial answer is possibly present im 
analyses of the vocalization of 7? gilli in the process of responding to 
the sound-reflecting barrier. The previously described caution and 
approach behavior serves as a context for the observed whistles. In all, 
from the first detection of the barrier to the successful passing, some 
66 whistles were noted. These comprised 16 different contours, 8 of 
which were similar to those used by the captive 7! truncatus in the 
12 most frequent calls. 

One salient difference between 7) truncatus whistles and T gilli 
whistles appears. The wild 7. gilli whistles, while sometimes similar 
in contour, were usually longer in duration than those produced by 
captive animals. This is an acoustically predictable happening, based 
on the sound production behavior of other animals, notably man. That 
is, as the reverberation of the environments is decreased and anecho- 
icity is approached, any given utterance becomes longer in time. This is 
equivalent to saying that humans in a small room with sound reflecting 
walls will produce words shorter in duration than if they are some 


224 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 
TABLE | 


OCCURRENCE OF WHISTLE CONTOURS FOR 
TURSIOPS TRUNCATUS AND TURSIOPS GILLI 


ANIMAL 

__ CONTOUR T. TRUNCATUS RANK OT. GILLI CRANK 
ut, USED | USED 13.5 
LeeN USED 2 —— — 
VEX USED 3 USED 1.0 
lav) USED 4 USED 45 
S\N USED 5 USED 13.5 
W\ USED 6 USED 45 
LIVI* USED 7 USED 135 
\ USED 8 USED 2.5 
NG USED 9 — ——- 
WIN USED 10 a — 
—— USED i USED 13.5 
VU USED 12 USED 13.5 
Su USED 13 — — 
NSW USED 14 — — 
I\SISS\SN\ _ ——USED 15 — — 
iol, — — USED 13,5 
—_-\ —— — USED 9.0 
Wt — — USED 6.5 
JNREIN — = USED 9.0 
ING —. — USED 9.0 
TIN — — USED 6.5 
=o — — USED 25 


distance apart in an open field. Table 1 summarizes the whistle con- 
tours noted in interchange between captive 7) truncatus and those 
observed in 7. gilli during the barrier problem, with due allowance for 
the lengthening phenomena. 

It will be noted that several identical rank occurrences have been 
entered for 7’ gilli, a reflection of the small sample observed. The 
rankings tell only how often a signal was produced, not when. Several 
of the contours were produced in isolation, with several short sequences 


Porpoise behavior DDN 


also occurring. It is not known at this juncture just what constraints 
exist in their syntax, although the increasingly greater library of their 
sounds will eventually make such a determination possible. 


DiscussIon 


Direct conclusions as to the communicative value of whistles produced 
by either species of Tursiops considered are of course not possible from 
the aforementioned observations. However, these observations do add 
some interesting support to the porpoise whistle communication hy po- 
thesis. The occurrence of whistling in a precise relationship to specific 
behavior, 7.e. response to a new situation, does indicate the possibility 
of purposeful sound production. Of further interest is the use of some 
identical or near identical contours by different species. In fact, it 
would seem almost mandatory for mutually understandable calls to 
exist in light of observation of cooperative behavior as pointed out by 
Brown and Norris (1956) and Norris and Prescott (1961). Enough 
importance has been placed on the significance of the interspecific 
whistle similarities observed by authors in this present study that 
further analyses into possible intergeneric similarities are in progress. 


ACKNOWLEDGMENTS 


We wish to thank Dr. K. S, Norris of the Department of Zoology of the 
University of California, Los Angeles, and R. L. Eberhardt of the 
Lockheed-California Company, Burbank, for their assistance in field 
observations. Also we wish to thank Miss Ruth M. Haugen, of Lock- 
heed and Dr. David kK. Caldwell, Los Angeles County Museum for 
their constructive comments on the manuscript. In addition we ap- 
preciate the cooperation of Messers. D. H. Brown and J. H. Prescott of 
the Marineland of the Pacific Oceanarium in the collecting of data on 
T. truncatus. 


LITERATURE CITED 
BROWN, D.N., ann kh. S. NORRIS. 
1956. Observations of captive and wild cetacea. Jour. Mamm., 37 (3) :120-125. 
DREHER, JOHN J. 
1961. Linguistic considerations of porpoise sounds. Jour. Acoustical Soc. Amer., 
33(12) 1799-1800. 
FRAZER, E C. 
1947. Sounds emitted by dolphins. Nature, London, 160, (4074). 
LILLY, J. G. 
1961. Man and dolphin. Garden City, N. Y.: Doubleday and Co., Inc., 312 pp. 


226 ~=Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


McBRIDE, A. F, anv D. O. HEBB. 

1948. Behavior of captive bottle-nosed dolphin, Tursiops truncatus. Jour. Comp. 
and Physiol. Psych., 41:111-123. 

NORRIS, K. S., anp J. H. PRESCOTT. 

1961. Observations on Pacific Cetaceans of Californian and Mexican Waters. 
Univ. California Publ. in Zool., 63(4)291-402. 

PIRES Kas 

1948. Tone Languages. Ann Arbor: Univ. of Mich. Press. 

TOMLIN, A. G. 

1955. O povedenii i Zvukovoi signalizatsii Gitvobraznykk. (On the Behavior and 
Sound Communication of Cetacean.) Trudy Instituta Okeanologit ANSSSR. 
18, 28-47. 

WOOD, E G., 

1953. Underwater sound production and concurrent behavior of captive porpoises 
Tursiops truncatus and Stenella plagiodon. Bull. Marine Sci. Gulf and Carib- 
bean, 3:120-133. 


CONTRIBUTIONS FROM THE LOS ANGELES MUSEUM 
—CHANNEL ISLANDS BIOLOGICAL SURVEY 
34. A FOSSIL BIRD, CARACARA, FROM 
SANTA ROSA ISLAND 


HiILpEGARDE Howarp 
Los Angeles County Museum 


The Los Angeles Museum Channel Islands Biological Survey was 
curtailed precipitately in December, 1941, with the bombing of Pearl 
Harbor, Expedition No. 13 was at that time on Santa Rosa Island. 
Comstock (1946: 105-107) gave a brief report of this last expedition. 
Regarding the paleontological work, there is the notation that King 
A. Richey, John C. Stock and Harry Fletcher prospected in the Tecolote 
Canyon area, November 11-14, 1941, and “reported a number of 
exposures suitable for working” Two of the paleontologists returned to 
the mainland on November 15, presumably with the intent of report- 
ing their investigations to the late Chester Stock, then Senior Curator 
of Earth Sciences at the Los Angeles Museum, and also of the geo- 
logical faculty of the California Institute of Technology. The Institute, 
under Dr. Stock, had some years before conducted paleontological 
work in Santa Rosa Island. All plans for continuing excavations by 
either institution were, of course, set aside with the advent of the war. 
The few Dwarf Elephant fossils collected on the prospecting trip of 
1941 (in field bags labelled “S.R.I.-1” to “S.R.I.-4) were stored for 
future study. No one noted that the three fragments in the bag marked 
“S.R.1.-2” were avian. 

As part of a project concerning fossil birds of the southwest (sup- 
ported by a John Simon Guggenheim Foundation Fellowship) I was 
assisted in the summer of 1962, by Wm. D. Arvey, Long Beach State 
College. A search was made through the entire vertebrate fossil collec- 
tion of the Los Angeles County Museum for unrecorded avian ma- 
terial. In this manner we came upon the fragments from Santa Rosa 
Island. 

In addition to the “S.R.I.—2” notation on the field bag, the inscrip- 
tion with the bird bones reads, “Santa Rosa Island, near mouth of Cor- 
ral Canyon in sea cliff. Richey, 11.12.41” Phil C. Orr, who in recent 
years has led the Santa Barbara Museum of Natural History’s paleon- 
tological investigations on the Channel Islands, describes in detail the 
wavecut platforms of Santa Rosa Island and the deposits that cover 


227 


228 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


them (Orr, 1960) and refers the upper, or Tecolote member of the 
Santa Rosa Island Formation to Wisconsin age, based on radiocarbon 
datings of from 10,400 to beyond 30,000 years B.P. In correspondence 
he states that the deposits in Corral (better known as Arlington) Can- 
yon are all of the Tecolote member. 

' The specimens collected by Richey consist of a proximal and distal 
end of tarsometatarsus and a distal end of tibiotarsus, probably all of a 
single individual. The bird represented was a caracara, which I assign 
to the species Caracara prelutosus (Howard) described from the Pleis- 
tocene of Rancho La Brea. The tibiotarsus shows no diagnostic charac- 
ters, but both fragments of tarsometatarsus agree with this element of 
C. prelutosus (Howard, 1938; 235-236) as follows: distalmost fora- 
men on posterior side very slightly above intertrochlear space and 3.5 
mm. below upper foramen; near proximal end, internal side of shaft 
with deep central depression, and greatest depth of internal face of 
shaft at approximately 7 mm. below distal extremity of calcaneum. 
Other measurements are as follows: tarsometatarsus, breadth of proxi- 
mal end 13.6 mm., breadth distal end 14.5 mm., breadth middle 
trochlea 5.0 mm.; tibiotarsus, breadth distal end 13.4 mm. 

Other recorded species of birds from Santa Rosa Island (Howard, 
1944) were collected by the California Institute of Technology in the 
late 1920’s and early 1930's, and were referred to recent species of 
goose and ducks (Branta canadensis, Anas carolinensis, and Anas, sp.). 
The Santa Barbara Museum of Natural History’s expeditions to Santa 
Rosa Island during the last fifteen years, have yielded a few more avian 
fossils. A report on this material will be forthcoming. 


LITERATURE CITED 


COMSTOCh, JOHN A. 

1946. Contributions from the Los Angeles Museum—Channel Islands Biological 
Survey. 33. Brief notes on the expeditions conducted between March 16, 1940 
and December 14, 1941. Bull. So. Calif. Acad. Sci., 45(2):94-107. 

HOWARD, HILDEGARDE 

1938. The Rancho La Brea Caracara: A new species. Carnegie Inst. Washington, 
Publ. No. 487, pp. 217-240. 

1944. Miscellaneous avian fossil records from California. Bull. So. Calif. Acad. 
Sci., 43(2):74-77. 

ORR: PHilEEG: 

1960. Late Pleistocene marine terraces on Santa Rosa Island, California. Bull. Ge- 
ological Soc. Amer., 71:1113-1120, 8 figs., 1 pl. 


A NEW GULF OF CALIFORNIA PERIPLOMA 


Mark E. RocGers 


Loma Linda, California 


INTRODUCTION 


Among the many interesting and unusual mollusks taken by the 
“Ariel” Expedition, August 28 through September 2, 1960, was a new 
member of the genus Periploma. This new species is of especial interest 
as it represents the second member of the subgenus Halistrepta, Dall 
1904, and the first of that taxon to be recorded from the Gulf of Cali- 
fornia. It may be known as: 


Periploma (Halistrepta) myrae, NEW SPECIES 


Figure 1. Periploma (Halistrepeta) myrae, new species. Exterior view of right 
valve of holotype, x2. 


Figure 2. Periploma (Halistrepta) myrae new species. Hinge area of right valve 
of holotype, x3. 


Description: Shell rotund, with a concave dorsal margin both anterior 
and posterior to the beaks, so thin as to be fragile, white, the left valve 
flatter than the right. The right valve is ornamented by somewhat dis- 
contimuous undulant sculpture. Left valve less coarsely sculptured, the 
undulations seemingly more continuous, though discontinuity does 
exist. Surface of shell smooth. Under high magnification the texture of 
the shell appears granular. The anterior and ventral margins are 


229 


230 ~=3©>— Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


broadly rounded; three minute lines extend anteriorly from the beaks 
but are visible only with difficulty (shghtly overemphasized in Fig. 1). 
These lines are apparently very shallow furrows, and are shown as 
such in Figure 1, but this could not be determined with certainty. 
Posterior margin abruptly truncated, producing a broad, short ros- 
trum. This rostrum is set off by a low rib, which extends diagonally 
from the beak, formed by the folding of the concentric undulations. 
This rib forms the boundary between the posterior and ventral mar- 
gins. The rostrum is ornamented by faint continuations of the con- 
centric undulations and by fine growth lines. Interior subnacreous. 
Posterior muscle-scar quite large, wedge-shaped, very faint. Anterior 
muscle-scar and pallial line obscure. The pallial smus broadly wedge- 
shaped, with the anterior end rounded. Resilifer small, narrowly 
spoon-shaped, nearly vertical. A dull area bordered by a more polished 
narrow margin indicates the resilium attachment. A short, narrow 
clavicular prop supports the resilifers. The props extend diagonally 
and posteriorly, fading out along the anterior-dorsal edge of the pos- 
terior muscle-scar, It is possible that this prop serves more as a muscle 
attachment than as a support for the resilifer. 

Holotype: To be deposited in the Stanford Univ. Paleo. Type Coll., 
number 9499. Length, 20.2 mm., altitude 16.4: mm. The holotype was 
broken subsequent to collecting and is represented by the repaired 
right valve and fragments of the left valve. 

Ty pe locality: Trawled in 15-25 fms., just off Loreto, in the channel 
between Loreto and Carmen Id., Baja California, Mexico. 29 August 
1960. “Ariel” Expedition. 

Commentary: In 1904 Dall described the very rare Periploma sul- 
cata, from San Pedro, California, and established the sectional name 
Halistrepta for it. In both 1908 and 1915 he used Halistrepta in a sub- 
generic sense, which rank it is accorded here. 

The discontinuous, undulant sculpture and prominent rostrum place 
Periploma myrae in this subgenus. It is very unfortunate that the 
lithodesma was not present for comparison. 

While Periploma myrae is related to P. sulcata it may be distin- 
guished at a glance by the truncated posterior end, by the doubly con- 
cave dorsal margin, prominent rostrum, and three faint furrows rather 
than a single furrow extending toward the anterior end. The clavicu- 
lar prop is also less heavily developed and the resilifer is vertical rather 
than extending forward. 

Periploma myrae resembles P. discus Stearns, 1890 in general shape, 


New Periploma 231 


but that form lacks the undulant sculpture in the adult form. The 
prominently truncated posterior and rostrum are also not present on 
P. discus. Young specimens of P. discus occasionally have undulant 
interior sculpture (as shown by a specimen in the Burch coll. from off 
Redondo Beach, California) which disappears as the shell grows older. 

This beautiful new form is dedicated to Dr. A. Myra Keen of Stan- 
ford University in grateful appreciation of her many contributions to 
malacology and paleontology. 


ACKNOWLEDGEMENTS 


I wish to express my thanks for much assistance to Mr. and Mrs. E. P. 
Chase of the San Diego Museum of Natural History, to Dr. A. Myra 
Keen, to Mr. and Mrs. J. Q. Burch and to Dr. S. Stillman Berry for his 


help and guidance. 
The excellent hand drawings are by Dr. Bruce Campbell. 


REFERENCES CONSULTED 


DALL, WILLIAM HEALEY 

1904. A new species of Periploma from California. The Nautilus, 17 (11) :122-123. 

1908. Reports on the dredging operations off the west coast of Central America to 
the Galapagos, to the west coast of Mexico, and in the Gulf of California. XIV. 
The Mollusca and Brachiopoda. Bull. Mus. Comp. Zool., Harvard, 43 (6) :205- 
487, pls. 1-22. 

1915. A review of some bivalve shells of the group Anatinacea from the West 
Coast of America. Proc, U.S. Natl. Mus., 49 (2116) :441-456. 

1921. Summary of marine shellbearing mollusks of the northwest coast of Amer- 
ica. Bull. U.S. Natl. Mus., no. 112,217 pp., 22 pls. 


GRANT, U.S., IV, ann H. R. GALE 

1931. Catalogue of the marine Pliocene and Pleistocene Mollusca of California 
and adjacent regions. Mem. San Diego Soc. Nat. Hist., 1:1-1036, 15 figs., pls. 
1-32. 

KEEN, A. MYRA 

1937. An abridged check list and bibliography of west North American marine 
Mollusca, Stanford University, Calif. 84 pp. 

1958. Sea shells of tropical west America. Stanford University, Calif. 624 pp., 
illus. 

OLDROYD, IDA S. 

1924. The marine shells of the west coast of North America. Stanford Univ. Pubs., 
Univ. ser., Geol. Sct., (1) :1-247, 57 pls. 

OLSSON, AXEL A. 

1961. Mollusks of the tropical eastern Pacific. Panamic-Pacific pelecypoda. 
Ithaca, N.Y.. Paleo. Research Institution, 574 pp., 86 pls. 


THE OCCURRENCE OF SONORASPIS CALIFORNICA 
FROM EAST-CENTRAL NEVADA 


TAKEO SUSUKI 
University of California, Los Angeles 
AND 
WitiiamM W. LuMspDEN 
Long Beach State College 


In several publications the genus Sonoraspis Stoyanow (1952:50-53) 
was either regarded as a paleontological puzzle (Lochman, 1952: 137) 
or overlooked in its stratigraphic significance (Palmer, 1954:67), or 
even misrepresented (Moore, et al., 1959:0224), but invariably con- 
fused with Glossopleura Poulsen (1927:268), from which it sharply 
differs in its paleontological and easily observable nature, its strati- 
graphic position, and its paleogeographical distribution. This genus 
possesses clearly outlined characteristics: a thorax always with eight 
segments, and certain mixed features of Anoria Walcott (1924:54) 
and Glossopleura but never all of them. This provides a basis for spe- 
cific separation and an easy identification. It should be stated that thus 
far Sonoraspis has never been located in the same strata with Glosso- 
pleura. The purpose of this article is to present additional data acquired 
in the recent research of the writers. This paper is the first in a series 
of proposed reports dealing with the distribution of Sonoraspis and its 
relation to Glossopleura. 

A significant clue as to the presence of an eight-segmented form in 
southern California was briefly mentioned by Clark (1921:6) while 
discussing Bathyuriscus howelli var. lodensis. Based on Clark’s discus- 
sion, Stoyanow and Susuki (1955:467-470, pl. 1, figs. 1, 2) conducted 
an extensive research in the Marble Mountains, San Bernardino 
County, California (Bristol Mountains of Clark), and located there 
specimens of Sonoraspis about fifty feet below the base of the Bonanza 
King formation (Hazzard and Mason, 1936:234-238). This find by 
Stoyanow and Susuki revealed that a seaway containing Sonoraspis 
fauna existed between the Sonora Basin, Mexico and southern Califor- 
nia. Before Sonoraspis was discovered in southern California, McKee 
(1947:288) postulated an early Middle Cambrian seaway continuous 
from western Sonora north through Arizona and into the Grand Can- 
yon area. McKee (1947:288) based this seaway on the presence of 
Glossopleura in the Harquahala Mountains to the south of the Grand 


233 


234 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


Canyon. Stoyanow (1948:323-324) was not in agreement with 
Mchee’s interpretation, pointing out the lithological similarity be- 
tween the Cambrian of Harquahala Mountains and the Grand Canyon, 
and that it seemed more probable that the Harquahala trilobites be- 
longed in the Cambrian basm of the Grand Canyon, masmuch as 
southwestern Arizona was occupied by a pre-Cambrian Massive of 
Mazatzal Land (Stoyanow, 1942:1264; 1936:462). A westward 
Paleozoic portal im the Sonora Basin was first suggested by Schuchert 
(1910: pl. 52) and Schuchert’s idea was corroborated by Stoyanow 
(1942: 1263-1264). 

The presence of Sonoraspis thirty feet below the stratum containing 
Albertella was established in the Inyo Mountains, California (Stoya- 
now, 1958:347), thus extending northwestward the geographic dis- 
tribution of Sonoraspis. The occurrence of Sonoraspis in the Inyo 
Mountains may well suggest a connection with the Sonora Basin 
through the Marble Mountains during early Middle Cambrian time. 
This discovery is not only significant paleogeographically, but ex- 
tremely important stratigraphically in that from this relationship, the 
stratigraphic position of Sonoraspis was determined. It should be noted 
that in the Canadian Rockies (Rasetti, 1951:109), Glossopleura zone 
is well above the Albertella zone and a similar stratigraphic relation 
exists in the Grand Canyon section (McKee, 1945:30, 33). Although 
Sonoraspis and Albertella had been found in the Sonora section, their 
stratigraphic interrelation there was not clearly understood, smce 
Lochman (1952:180) did not recognize Sonoraspis as a valid genus 
but equated it both paleontologically and _ stratigraphically with 
Glossopleura. In discussing Glossopleura species, Lochman (1952: 137) 
was unable to explain why Glossopleura mckee Resser from the Bright 
Angel shale of the Grand Canyon section showed consistently seven 
segments in the thorax, yet all the species from Sonora, even though 
the cephalon and pygidium appeared identical to G. mckee, showed 
eight thoracic segments. 

The find of Sonoraspis by the present writers in Currant Creek Gap 
section, east-central Nevada (Fig. 1) indicates the first recorded extent 
of the Sonoraspis fauna considerably northeastward within the main 
Cordilleran trough. 

This specimen from Nevada is closer to S. californica Stoyanow and 
Susuki (Holotype, UCLA Invert. Paleo. Cat. no. 23460) than to 
S. gomezi Stoyanow (1952:54-55) by tuberculation on all thoracic 
segments which increase in size (from circular to elongated) toward 


Sonoraspis californica from Nevada 235 


\ 
Buvelse WHITE PINE 


U.C.L.A. Loc. 4444 


Currant 


LINCOLN CO. 


Pioche 


Figure 1. Map showing east-central Nevada where Sonoraspis californica was 
found. 


236 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


the segmented pygidium, whereas S. gormezi has only six posterior 
thoracic segments bearing tubercles and an unsegmented pygidium. 
Ty pe: Hypotype, UCLA Invert. Paleo. Cat. no. 34974, Figure 2. 
Locality: UCLA Invert. Paleo. Loc. no. 4444. 
Occurrence: The described type has been collected at a roadcut on 


Figure 2. Sonoraspis californica Stoyanow and Susuki, Hypotype, ucta Invert. 
Paleo. Cat. no. 34974, specimen enlarged x2 to show posteriorward development 
from circular to elongated tubercles on the thoracic segments. 


Sonoraspis californica from Nevada 237 


the northside of U.S. Highway 6, in thin-bedded, platy, gray limestone, 
300 feet north and 400 feet west of the southeast corner of Sec. 24, 
T. 11 N., R. 58 E., Currant Mountain quadrangle (U.S.G.S., 1957 
ed.), Currant Creek, Nye County, Nevada (Figs. 3 and 4). 


Figure 3. Currant Creek Gap, Nevada. Westward view. Lighter colored exposure 
at the base, on the right side of photograph is where Sonorapsis californica was 
found. 


Pe ES 


Figure 4. Close-up of the limestone outcrop. 


238 ~=— Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 
ACKNOWLEDGMENTS 


The writers wish to acknowledge the support given by the Dept. of 
Geology, University of California, Los Angeles for the field work and 
to Mrs. Opal L. Kurtz for the drafting. 


LITERATURE CITED 


CLARK, CLIFTON W. 

1921. Lower and Middle Cambrian formations of the Mohave Desert. Univ. Calif. 
Pub. Geol. Sci. Bull., 13:1-7. 

HAZZARD, JOHN C., sno MASON, JOHN FE 

1936. Middle Cambrian formations of the Providence and Marble Mountains, 
California. Geol. Soc. America Bull., 47:229-240, 1 fig. 

LOCHMAN, CHRISTINA 

1952. Trilobites, pp. 60-161 pls. 15-31, in Cooper, G. A., et al., Cambrian strati- 
graphy and paleontology near Caborca, northwestern Sonora, Mexico. 
Smithson. Misc. Coll., 119: (1):184, 31 pls. 

McREE, EDWIN D. 

1945. Stratigraphy and ecology of the Grand Canyon Cambrian, pp. 1-168, pls. 
1-15, in Mckee, E. D. and Resser, C. E. Cambrian history of the Grand 
Canyon region. Carnegie Inst. Washington Pub. 563, 232 pp., 27 pls. 

1947. Paleozoic seaways in western Arizona. Amer. Assoc. Petroleum Geol. Bull., 
31:282-292. 

MOORE, RAYMOND C., et al. 

1959. Treatise on invertebrate paleontology, Part O, Arthropoda. Geol. Soc. 
Amer., 560 pp., 415 figs. 

PALMER, ALLISON R. 

1954. An appraisal of the Great Basin Middle Cambrian trilobites described be- 
fore 1900. Geol. Surv. Prof. Paper 264-D, pp. 55-85, pls. 13-17. 


POULSEN, CHR. 

1927. The Cambrian, Ozarkian and Canadian faunas of northwest Greenland. 
Meddelelser om Gronland, 70:233-348, pls. 14-21. 

RASETTI, FRANCO 

1951. Middle Cambrian stratigraphy and faunas of the Canadian Rocky Moun- 
tains, Smithson, Misc. Coll., 116:(5):1-277, 34 pls. 

SCHUCHERT, CHARLES 

1910. Paleogeography of North America. Geol. Soc. Amer. Bull., 20:427-606, pls. 
46-101. 

STOYANOW, ALEXANDER 

1936. Correlation of Arizona Paleozoic formations. Geol. Soc. Amer. Bull., 47:459- 
540, 1 pl., 5 figs. 

1942. Paleozoic paleogeography of Arizona. Geol. Soc. Amer. Bull., 53:1255- 
1282, 5 pls., 3 figs. 

1948. Some problems of Mississippian stratigraphy in southwestern United States. 
Jour. Geology, 56:313-326. 


Sonoraspis californica from Nevada 239 


1952. The original collection of Cambrian trilobites from Sonora, pp. 49-59, pl. 
14, in Cooper, G. A., et al., Cambrian stratigraphy and paleontology near 
Caborca, northwestern Mexico. Smithson. Misc. Coll, 119: (1):7-184, 31 pls. 

1958. Sonoraspis and Albertella in the Inyo Mountains, California. Geol. Soc. 
Amer. Bull. 69:347-352, 1 pl. 

STOYANOW, ALEXANDER, anp.SUSUKI, TAKEO 

1955. Discovery of Sonoraspis in southern California. Geol. Soc. Amer. Bull., 
66:467-470, 2 pls. 


WALCOTT, CHARLES D. 
1924. Cambrian geology and paleontology. Cambrian and lower Ozarkian trilo- 
bites. Smithson Misc. Coll., 75 (2) :53-60, pls. 9-14. 


\ 


“ 


val 


NOTES ON THE OCCURRENCE, OBSERVATIONS AND PUBLIC 
HEALTH SIGNIFICANCE OF THE PAJAROELLO TICK — 
ORNITHODOROS CORIACEUS KOCH, 

IN LOS ANGELES COUNTY 


WiLuiAM G. WaLpron! 
Los Angeles County Health Department 


Recently, several reports in non-scientific publications have disclosed 
that little is known locally about the Pajaroello tick—Ornithodoros 
coriaceus Koch. Because of these reports, studies were conducted by 
the author in the Los Angeles National Forest, the suggested habitat 
of this tick, to gather data on the prevalence of O. coriaceus and infor- 
mation on the severity and virulence of its bite. 

Nutall (1908) first described the Pajaroello in Mexico around the 
Gulf of Tehuantepec. He also found this tick in 1908 in Santa Barbara 
County. Hermes (1960) and Kohls (1944) have collected numerous 
specimens in the Mount Hamilton area where it “flourishes in the deer 
beds among the low scrub oaks; Quercus dumosa. Hermes likewise 
acknowledges that this species occurs commonly in the more moun- 
tainous coastal counties in California. Hoffman, quoted in Cooley & 
Kohls (1944), states “O. coriaceus is a native of the hot and temperate 
regions along the Pacific extending all the way from California to 
Chiapas (Mexico)? 

The first step on this Pajaroello study involved checking records of 
collections made locally. Surprisingly few were available. A total of 
only 4 collections were locatéd which had been made in the San Gabriel 
and Santa Monica Mountains. 


O. coriaceus collections: 


A. UCLA, Dr. J. N. Belkin 
1. Los Angeles County, Pasadena, ex: soil, [V-7-53 
22 nymphs, coll. student (UCLA) 
B. Los Angeles County Museum, Dr. Fred Truxal 
1. Los Angeles County, Township T2N-R11W, Sec. 29 
SW 14, ex: dirt, 5800’, 6/49, coll. 
Dr. R. O. Gilbert, 1 nymph, 1 ad. 
2. Los Angeles County, Devil’s Punch Bowl, Ex: man, 
8/21/52, 1 nymph, 1 larvae 


1Entomologist. 


242 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


C. Long Beach State College, Dr. Elbert Sleeper 
1. Los Angeles County, San Dimas Exp. Forest., 
VII-II-59, #2, 1-ad. Coll. M. Knox and E. Sleeper. 


Forest rangers, patrolmen and fire guards of the U.S. Forest Service 
were quite familiar with the Pajaroello. Many advised the author that 
they first became acquainted with the tick and its bite while assigned 
to the Los Padres National Forest in the Santa Barbara area. 

A great deal of interesting and pertinent information was gathered 
from these men, who not only were aware of the Pajaroello, but were 


Figure 1. Ventral (upper) and dorsal (lower) view of Ornithodoros coriaceus 
Koch. Photos courtesy of Dr. Fred Truxal, Los Angeles County Museum. 


Pajaroello tick 243 


likewise highly trained and capable observers. In practically every 
instance the bites, which allegedly implicated the Pajaroello, occurred 
while the Forest Service personnel were working on a forest fire. This, 
of course, implies that the men were away from their stations and 
offices and were living in a more or less primitive fashion for the period 
of the fire. Also, the men were working in the brush where they had 
ample opportunity for exposure to arthropod bites. 

According to Merrell (1948), the U.S. Forest Service in the Angeles 
National Forest reported in 1940 that several members of a CCC main- 
tenance crew allegedly were bitten by arthropods thought to be Paja- 
roello ticks. This is the first recorded reference to O, coriaceus in Los 
Angeles County. 

According to Herms (1950) the main host of O. coriaceus is the 
mule deer, Odocoileus species. It is therefore reasonable to assume that 
ectoparasites may be spread into new areas as the deer herds migrate 
from one section to the other due to the pressures of food, fire, water 
and hunters. The first collections above 6,000 ft. known to the author 
were made at Table Mountain in the Big Pines area which is about 
7200 ft. altitude. The other 5 collections made by the author were 
primarily between 4500 and 7000 ft. It was observed that the highest 
density of Pajaroellos should occur in the spring of the year. 


O. coriaceus specimens in the author’s collection: 


1. Los Angeles County, Camp 37, Calif. State Highway Camp, 
Angeles Forest Highway, Ang. Forest, 6000’, 9/58, 
coll. W. G. Waldron, 3 ad., ex: brush. 


2. Los Angeles County, 300’ S of Table Mt. Reservoir, 
So. Slope, 7200 ft., Big Pines, Ang. Forest, 
4/2/59, 1 ad., coll. W. G. Waldron, Ex: brush, 162-59. 
3. Los Angeles County, Table Mt. Campgrounds at West Bend, 
Big Pines, Ang. Forest, coll: W. G. Waldron, 1 ad, 
1 nymph, ex: brush, 7500 ft. 
4. Los Angeles County, Swartout, Big Pines, 
Ang. Forest, 6/30/58, 
1 ad., 2 nymphs, coll. W. G. Waldron, ex: man, 6800 ft. 
5. Los Angeles County, Jackson Lake, Big Pines, Ang. Forest, 
4/1/59, coll. E.S. Cross (USFS), 2 ad, 5 nymphs, 
ex: brush, 6500 ft. 


Although most Forest Service personnel are not trained entomolo- 
gists and do not usually have equipment necessary for identification 


244. Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


of the arthropods, they accurately described and even furnished the 
proper tick upon request (collection #5 above). Apparently other 
Ornithodoros species such as O. hermsi, O. turicata and O. parkeri 
which could be mistaken for O. coriaceus are not readily found where 
the Pajaroellos have been collected. 

Nutall (1908) graphically described the effect of the bite and feed- 
ing habits of O. coriaceus. Hermes (1950) also describes an account of 
two bites which a former student suffered some years ago. The descrip- 
tions by Nutall and Hermes are very similar. Hesper N. MacMillen 
and E. C. Loomis, in personal communications to the author, likewise 
give the same general description of the Pajaroello tick bite from their 
own personal experiences. 

The author has gathered a great deal of data on the Pajaroello bite, 
particularly from Forest Service personnel, and they all follow more 
or less the same pattern and in most instances no pain is felt at the 
time the tick attaches its mouth parts to the skin of the human host. 
This is the converse of descriptions cited above from Nutall and others. 
Possibly, most of the tick bites investigated by the author were inflicted 
while the person was at rest and therefore no noticeable pain was evi- 
denced. It is also possible that the tick actually causes pain as a result 
of feverishly attaching its mouth parts to the host while the body of 
the host is in motion, the tick thus protecting itself from being dislodged 
from the point of attachment. This is an assumption upon the part of 
the author and does not reflect any valid data. The author also has 
noted that the reaction to the bite appears to be a direct allergic re- 
sponse of the host to the saliva of the tick. 

Despite comments to the contrary, the Pajaroello is not as dangerous 
as a rattlesnake. The tick injects no toxin per se, and very few bite 
patients have been hospitalized. Actually only two cases of this severity 
have been recorded locally. The bite may be more painful and annoy- 
ing to those people who react strongly to the material in the saliva. 
Likewise, as MacMillen and Loomis graphically indicated to me, a 
person may become increasingly sensitized to the bite to the poimt 
where additional exposures could cause acute discomfort to the human 
host. 

Records are available where the “incise and suck” treatment has 
been employed for Pajaroello as well as for rattlesnake bites. The 
method suggested by the Health Officer of the Los Angeles County 
Health Department and the local office of the American Red Cross 
suggests suction might be of aid if applied scon enough after the bite, 


Pajaroello tick 245 


but neither recommend the need for incision. One of the most impor- 
tant steps is the application of a good salve, preferably with an anal- 
gesic, and extreme care against secondary infection. No records are 
available which in any way implicate the Pajarcello as being a vector 
of disease. This is not to say that this Argasid is not a potential, but 
rather that it has never been implicated, circumstantially or otherwise, 
as a transmitter of disease to men or animals. 


SUMMARY 


It appears that the Pajaroello, contrary to some reports, is not as dan- 
gerous as a rattlesnake although its bite can be painful and uncom- 
fortable. Also, O. coriaceus is not new in California, although there is 
a definite dearth of collecting information for the area between Santa 
Barbara and the Mexican border. The Pajaroello is part of the fauna 
in the Angeles Forest and care should be taken to prevent unnecessary 
exposure to its bite; however, it does not appear to be a ““menace”’ to 
public health at this time. 


LITERATURE CITED 


COOLEY, R. A., anp G. M. KOHLS. 

1944. The Argasidae of North America, Central America and Cuba. Amer. Midl. 
Nat., Mono. No. 1, Univ. Press, Notre Dame, Ind. 

HERMS, W. B. 

1950. Medical Entomology. New York: The Macmillan Co., 643 pp. 

LOOMIS, E. C. 

1950. Ticks in California. Vector Views, 2:1. 

MERRELL, MURIEL L. 

1948. Pajaroello. U.S. Forest Serv. Rept., Angeles National Forest. 

NUTALL, G. H., ez al. 

1908. Ticks, a monograph of the Ixodoidea, Part I., Argasidae, pp. 1-104, Part II, 
pp. 105-116. 

ROBINSON, C-. S. 

1942. Some observations on the Pajaroello tick (Ornithodoros coriaceus). Jour. 
Forestry, 40:659-60. 


246 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


ERRATA 


The following errors regrettably appeared in the article “Blood para- 
sites of mammals of the Californian Sierra Nevada foothills, with spe- 
cial reference to Trypanosoma cruzi Chagas and Hepatozoon lepto- 
soma Sp. N? by Sherwin EF Wood which appeared in volume 61, part 
3, of the Bulletin: 


1. In Table 1, page 163, the numbers ‘‘2,1” in the TZ. cruzi column 
should appear opposite Peromyscus truei gilberti, and not opposite 
Reithrodontomys megalotis longicaudus. 


2. The last two full sentences on page 168 should read: “Where the 
central aperture has enlarged and the circular, band-form nucleus 
has become more labile, the parasite may extend into the aperture 
pushing a band of nuclear material before it. Most of these immature 
neutrophils have not yet developed their distinctive granulation” 


3. On page 170, line 2, the correct statement should be “2 yw thick” and 
not “2 u thick?’ 


A NEW SPECIES OF COLUMBELLID GASTROPOD 
FROM EASTER ISLAND 


Leo GrorGE HERTLEIN 
California Academy of Sciences! 


San Francisco 


Two lots of shells from Easter Island were presented to the California 
Academy of Sciences, one by Ray Summers and the other by Mrs. 
Paul Steele. These specimens were collected by Frey Sebastian 
Englert. 

Among these shells is an interesting little columbellid which appears 
to be quite distinct from any described species. Accordingly, it is here 
described as new. 

The writer here expresses his gratitude to several persons who 
compared specimens of this species with specimens in the collections 
_ in their respective institutions as follows: Dr. A. Myra Keen, Stanford 
University; Dr. Alison Kay, University of Hawaii and Bernice P. 
Bishop Museum; Dr. Harald A. Rehder, United States National 
Museum. 

The photographs, retouched by Mrs. Margaret Hanna, were pre- 
pared by Maurice Giles. Measurements were made by Dr. G Dallas 
Hanna. 


The new species may be known as: 


Zafra pascua Hertlein, NEW SPECIES 


l 2 3 


Figures 1-3. Zafra pascua Hertlein, new species. Fig. 1. Holotype. Height, 6.15 
mm., maximum diameter, 2.83 mm. Fig. 2. Paratype. Height, 5.91 mm., maxi- 
mum diameter, 2.80 mm. Fig. 3. Paratype. Height, 6.05 mm., maximum diameter, 


3.06 mm. (Photographs by Maurice Giles, retouched by Margaret Hanna). 


1Department of Geology. 


248 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


Shell small, elongately ovate, fairly thick; 41% whorls, the sutures 
impressed; protoconch dome-shaped, smooth; whorls sloping with a 
slight concavity on the posterior third; axial sculpture consists of 
rounded, axial ribs, about 15 on the penultimate whorl, extending 
from suture to suture but fading out on the anterior half of the body 
whorl; interspaces narrower than the ribs and crossed by very fine in- 
cised spiral striae; canal short, sculptured with 5 or 6 fine, well-defined, 
obliquely sloping spiral grooves which above and below give way to 
finer striae; aperture rather narrow, columella slightly curved, interior 
of inner and outer lips smooth; shell white except for a narrow black 
band beginning near the middle of the edge of the columellar lip and 
extending obliquely downward around the base of the canal, visible 
through the inside of the shell. Length, 6.15 mm.; maximum diameter, 
2.83 mm. 

Holotype: no. 12408 and paratypes, nos. 12409, 12410 (California 
Academy of Sciences Department of Geology Type Collection), from 
Loc. 34203 (CAS), Easter Island; Frey Sebastian Englert, collector. 

On some specimens of the paratypes of this species the axial ribs 
are nearly obsolete on the body whorl. The fine lines of growth are 
crossed by concentric striae lending a decussate appearance under 
magnification. 

The general shape and sculpture of the present species bear a 
general resemblance to that of Anachis. However, the narrow aper- 
ture and smooth inner and outer lips are quite different from mem- 
bers of that group. The characters of the aperture suggest the reason 
for placing it in the genus Zafra A. Adams. Dr. Rehder agreed in this 
general placement of this species (written communication May 18, 
1962) in or near the Zafra group and it is here provisionally so placed. 

The general shape and external sculpture of Zafra pascua, new 
species, resemble that of the species described as Columbella pumila 
Dunker? from Japan which later was placed in the genus Zafra 
by ‘Taki and Oyama’. The shell of the new species differs from that 
of Dunker’s species in the smooth interior of the outer lip and in 
the dark band of color around the canal, whereas the corresponding 
lip of Z. pumila was described as plicated or subdenticulated and the 
color of the exterior uniformly fuscous. 


2Columbella pumila Dunker, Malakozool. Blatter, Bd. 6, p. 224, January, 1860. 
Japan.—Dunker, Moll. Jap. Descript. et Tab. Trib. Icon., p. 6, pl. 1, fig. 4, 1861. 


‘Zafra pumila (Dunker), Taki and Oyama, Palaeo. Soc. Japan, Special Paper 
No. 2, pl. 43, fig. 10, 1954. 


A NEW SPECIES OF SAND-BURROWING 
MARINE AMPHIPODA FROM CALIFORNIA 


J. Laurens BARNARD 


Beaudette Foundation! 


Another species of the magnificent crustacean genus Kohaustorius 
J. L. Barnard (1957) is described. These small but complexly orna- 
mented animals apparently represent adaptations in the Amphipoda 
that are known for various sand-crabs in the Decapoda, namely greatly 
expanded, processiferous appendages for digging in sands and strongly 
setose antennae for filtering particles. The weakly armored mandibles 
apparently are related to the particulate kind of food obtained by 
many haustoriids. 


FaMILy HAUSTORIIDAE 


Genus Eohaustorius J. L. Barnard 
Eohaustorius J. L. Barnard 1957: 81; Gurjanova 1962: 400. 


Key To EOHAUSTORIUS 


1. Peraeopod 5 with erect cusp on dorsal posterior edge 

OH COCCI ters Taek ese er care En washingtonianus 
Peraeopod 5 lacking cusp on article 2 of peraeopod5 ........ Y 
Article 7 of gnathopod 1 as long as article 4 ..... sencillus, n.sp. 
Article 7 of gnathopod 1 half as long as article4............. 3 
Hind lobe of article 5 on peraeopod 2 large and long, reaching 
down to end of article 6,/article 6 rectangular ........ cheliferus 
Hind lobe of article 5 on peraeopod 2 smaller, not reaching to end 
Olmanicleon article6 bulbous = .....04505. 2408s 4 5. eous 


Www > 


es 


Eohaustorius sencillus, NEW SPECIES 
Figures 1 and 2 


Diagnosis: Article 6 of gnathopod 1 long, bulbous, bearing a short 
apical spine; peraeopod 5 lacking a cusp on dorsal posterior edge of 
article 2; hind lobe of article 5 on peraeopod 2 small, not reaching 
downward strongly (see figure). 

Holotype: Allan Hancock Foundation (AHF) No. 5736, female, 


4.5 mm. 


1Santa Ynez, California. 


249 


250 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


Sey 


> ms ~ b Me 
R . NP Y 
C= ZW 


RX 


fy) 
ted 
hh 
\V' 4" 
x \ 


Figure 1, Eohaustorius sencillus, n.sp. Female, 4.5 mm, station 4812. Lateral view 
of animal. 


New marine amphipod 251 


| 
| 


Ss) 
a 


C2 
1 oa a 
VE 


SSOH 


Le 


= 
Jip 


Figure 2. Eohaustorius sencillus, n.sp. Female, 4.5 mm, station 4812. A, B, man- 
dibles; C, D, gnathopod 1; E, gnathopod 2; F, peraeopod 2; G, coxa 3; H, uropod 
3; I, telson. ; 


Type locality: Station 4812, near Pt. Conception, California, 
34°34’05” N, 120°39’05” W, in 14 meters depth, January 16, 1957, 
collected by R. V. Velero IV of University of Southern California. 

Material: 42 specimens from 3 stations. 

Relationship: Unlike other species of the genus, this bears a large, 
inflated sixth article of gnathopod 1, armed with a small spine. In other 
species this sixth article is short and bears a spine either as long as the 
article or as much as twice as long. In southern California, the species 
is quickly distinguished from E. washingtonianus (Thorsteinson ) 
(see J. L. Barnard 1957) by the lack of cusp on article 2 of peraeo- 
pod 5. 

Ecology: In southern California this species has been found in only 
two samples at Pt. Conception in depths less than 10 fms and thus, 


252 Bulletin So. Calif. Academy Sciences / Vol. 61, Pt. 4, 1962 


scarcely penetrates into the southern area and its frequency in south- 
ern California is negligible. The species is better represented in Monte- 
rey Bay (station 6444, 18 specimens). 


LITERATURE CITED 


BARNARD, J. L. 

1957. A new genus of haustoriid amphipod from the northeastern Pacific Ocean 
and the southern distribution of Urothoe vavarini Gurjanova. Bull. So. Calif. 
Acad. Sci., 56 (2) :81-84, pl. 16. 

GURJANOVA, E. E 

1962. Bokoplavy severnoi chasti Tixogo Okeana (Amphipoda—Gammaridea), Ch. 
1. Akademiia Nauk SSSR, Opredeliteli po Faune SSSR 74:1-440, 143 figs. 


SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 


VOLUME 61, 1962 


INDEX OF SUBJECTS 


algodonensis, Macrobaenetes ... .1(2 
Amblystira angella, n.sp......... 137 
PAmalbyiStOMMIC ACM rass ae 5 os cls. 211 
Ambrysus stali,n.sp............. 185 
LNSTTHO) CHWUTOCES «00580080 ome eon nae 211 


Analysis of the habitat, web design, 
cocoon and egg sacs of the tube 
weaving spider Diguetia canities 
(McCook) (Aranea, Diguetidae) 65 

A new Ambrysus from South 
America (Hemiptera, Naucori- 


GINS) 6 5.6:6°5'5 5 Cia eee neem aon eae 185 
A new Gulf of California Peri- 
IOI 5 0.0 do See ee eee 229 


A new Megahippus from the Bar- 
stow formation San Bernardino 
CountyaCaliformia =. .)........ 113 

A new Pasiphaea (Crustacea, Deca- 
poda, Natantia) from southern 
California waters .............. 15 

A new species of chigger, genus 
Euschoengastia (Acarina, Trom- 
biculidae), with notes on other 
species of chiggers from the Santa 


Ana Mountains, California ..... 177 
A new species of columbellid gastro- 
pod from Easter Island ........ 247 


A new species of sand-burrowing 
marine amphipoda from Califor- 


TATE 4 ‘5’ arnt bog see biG ay ene eee ee 249 
angella, Amblystira ............. 137 
Argenna fossilis Petrunkevitch .. .143 


Arizona elegans eburnata Klauber. 31 


Blood parasites of mammals of the 
Californian Sierra Nevada foot- 
hills, with special reference to 
Trypanosoma cruzi Chagas and 
Hepatozoon leptosoma Sp. N. 


We 161, 246 
Caracara prelutosus (Howard) ...228 
chacei, Pasiphaea ............... 18 
CGlvalicidoideal 2.2.53 4..040506 aves 150 


Chionactis occipitalis occipitalis 
(Hallowell) 


Chthonuidaewiaoe meee: 145 
Comstock, John Adams (Resolution 
TO) ctsteh altars mintin eer ites eenale 1 
Contributions from the Los Angeles 
Museum—Channel Islands _bio- 
logical survey. 34. A fossil bird, 
Caracara, from Santa Rosa Is- 
lane aro eee ee OT 
Crotalus cerastes Hallowell ...... 34 
Crotalus mitchelli pyrrhus ....... 35 
Crotalus viridis helleri ........... 36 
Cryptobranchidaeymer-see one seek 211 
Dependence on temperature of Ca/ 
Mg ratio of skeletal structures of 
organisms and direct chemical 
precipitates out of sea water .... 45 
Dictyonota maroccana Ribaut ... .141 
Dictyonota tricornis (Schrank) ...141 
Diguetia canities (McCook) ...... 65 
elitha, Leptodictya .............. 136 
Engynoma isolata, n. nam. .......134 
Eintonovny Garner et 146 
Enulius unicolor (Fischer) ....... 195 
Eohaustorius sencillus, n. sp. ..... 249 


Epiplatymetra grotearia Packard . 3 
Euschoengastia californica 


GEwanee a Sie ener eas eas toc 182 
Euschoengastia criceticola 

Brernmancy ciate ask at 183 
Euschoengastia frondifera Gould ..179 
Euschoengastia otophila, n. sp. .... 179 
Euschoengastia radfordi 

Brennan and Jones ............ 183 
Euschoengastia terrestris Gould. . 182 
Fossil arthropods of California. 24. 

Some unusual fossil arthropods 

from the Calico Mountains nod- 

ul Sheen aera rs ts remiaa eta 143 
Galeatus maculatus (Herrich- 

Schae tier ero cine ne ae 135 
Galeatus spinifrons (Fallén) ..... 35 
Gambusia affinis patruelis Baird 

aingl (Girl Gooouorooctoe ccs 37 


Gyalopion quadrangularis 


(Giinther, aeons tae 196 
Haedus ojos, n. sp. ............-. 139 
Hamingia arctica Danielssen and 

INOGeM oii sae ena enn ator nee: 123 
Hepatozoon citellicola (Wellman 

BLIGH VV Orgy, ivevkspess Steichen eae 168 
Hepatozoon leptosoma n. sp. ..... 170 
Hepatozoon muris (Balfour) ..... 169 
Elymobuidae. v.55 st lees aston 211 
Hypsiglena torquata deserticola 

WaMWer \65. ces pecvolio sansa tes Beas 31 
Ichneumonoideats .42.).. .¢- sss an 150 
isolata, Engynoma .............. 134 
juliae, Palaeosminthurus ......... 147 
kelsoensis, Macrobaenetes ....... 93 


Lampropeltis getulus californiae 


(Blammville) eee ee iieciebttesa 32 
Lampropeltis getulus nigritus 

Zweifel and Norris ............ 196 
eChy tia? rgcncetrck 2 Pape tec 145 
Repidopteray 22)... earn cesines 150 
Leptodictya elitha, n.sp.......... 136 
leptosoma, Hepatozoon ......... 170 
Lichanura roseofusca gracia 

Kilauiberins us cscnese ieee ieee ioe 30 
Life history notes on Epiplatymetra 

grotearia Packard <7. 3...:2..+- 3 
Macrobaenetes, n. gen. .......... 91 
Macrobaenetes algodonensis, n. sp. 102 
Macrobaenetes kelsoenis, n. sp. .. 93 


Macrobaenetes sierrapintae, n. sp..105 
Macrobaenetes valgum 


(Strohecker) ian acento ee 98 
Masticophis flagellum piceus 

(Cope) scy2.see ae eer 32 
Masticophis lateralis lateralis 

@elalllowellli) Seer 32 
mckennai, Megahippus .......... 113 
Megahippus mckennai, n. sp. ..... 113 
Miyatrombicula scottae 

(Brennan) ye sarencracre oe 179 


Monarchistic dominance in small 
groups of captive male mosquito- 
fish, Gambusia affinis patruelis . 37 


myra, Periploma ............... 229 
Naitingis, n. gen. ............... 133 
2 


Or 


Neotrombicula californica 


(Ewing) ........@5pepReeerer 178 
Neotrombicula dinehartae (Bren- 
nan and Wharton) ........... 178 


Nonmarine molluscs from recent 
sediments near Vernon, Apache 


county, Arizona’; pene 25 
Notes on some reptiles and amphib- 
ians from western Mexico ..... 193 


Notes on the occurrence, observa- 
tions and public health signifi- 
cance of the Pajaroello tick—Or- 
nithodoros coriaceus Koch, in Los 
Angeles, county, Sn eeppee err 241 


Observations on scouting behavior 
and associated sound production 
by the Pacific bottlenosed por- 


poise (Tursiops gilli Dall) ..... D7 
Odontacarus linsdalei (Brennan 

and Jones) . ; 2 4-fescoeee nee 178 
oios, Haedus ................... 139 


On the recognition of a second spe- 
cies of the genus Pelagophychus .153 


Ornithodorus coriaceus Koch ..... 241 
otophila, Euschoengastia ........ 179 
Palaeosminthurus, n. gen. ....... 146 
Palaeosminthurus juliae, n. sp. ...147 
Palaeosminthuridae, n. fam. ..... 146 
pascua, Zafra .................. 247 
Pasiphaea chacei, n.sp........... 18 
Pelagophycus giganteus 
(Areschoug) =. 3; - eee 156 
Pelagophycus porra (Leman) ... .153 
Periploma myrae, n. sp. ........ .229 
Phyllorhynchus decurtatus perkinsi 
Klauber) ....css2) ee Seeeeeooe 32 
Pituophis melanoleucus deserticola 
Steyneger: ... sc: sce eee 33 
Plethodontidae ..2..25500enbnooe 912 
Proceedings of the Academy ..... 189 
Proteidae ... .. ..2 sage 210 
Pseudoficimia frontalis hiltoni Bo- 
gert and Oliver =o eee 196 
Pseudoschoengastia occidentalis 
Brennan’ . +... ieee 183 


Records of snakes from Joshua Tree 
National Monument, California. 29 
Report of a scyphozoan Stephano- 
scy phus simplex Kirkpatrick from 
the Arctic Ocean ............. 64 


Report of an echiuroid worm Ha- 
mingia arctica Danielssen and 


Koren from the Beaufort Sea .. .123 
Rhinocheilus lecontei lecontei Baird 

eincl Gamal . 53 0¢esedeoue sees 33 
Salamadridae .................. 912 
Salvadora hexalepis (Cope) ...... 34 
sencillus, Eohaustorius ........... 249 
Shoulder and upper arm muscles of 

salamanders ................. 205 
sierrapintae, Macrobaenetes ..... 105 
Sumer acura s is cares sl hae eas 211 
Sonoraspis californica Stoyanow 

AICS USUI ees keys ae a ko 233 
stali, Ambrysus ................. 185 
Stephanoscyphus simplex Kirkpat- 

TAUGIS, | din 0/06. SEEN a ee 64 


Studies on Neararctic desert sand 
dune Orthoptera. Part VI. A new 
genus and three new species of 
large sand-treader camel crickets 
from the Colorado Desert with 
KeySHAM COLES) Gara te. ss ae 89 

Suspected melanophore movement 


in fishes beyond the larval stage . 129 
Syrrhophis interorbitalis Langbar- 


tel and Shannon .............. 194. 
Syrrhophis modestus pallidus 
Dwellers eee ss ao els eee 8s 194. 


Taxonomic changes and descriptions 
of new Tingidae (Hemiptera) . .133 
The occurrence of Sonoraspis cali- 
fornica from east-central Ne- 
Via Calpe isiee mene raricta amine eee 233 
The significance of the petroliferous 
nodules of our desert mountains. 7 


Tingis stachydis (Fieber) ........ 135 
dimeusestmctulan (utom) sea 134: 
Trimorphodon vandenburghi 

ilauberc tree ecw ee ena 34 
GEOL Ae masse 8, Oe eat 148 
Tropidodipsas occidentalis Oliver . .197 
Trypanosoma cruzi Chagas ....... 161 
Tirsiopspevliia (Dall) Sense eae ene Di 


Typhlops braminus (Daudin) ....202 


Zafra pascua, n.sp.............. QA7 


INDEX OF AUTHORS 


Nlbi,, IRebyroaoraGl WS oo ees one sa deen: 113 
Barnard, J. Laurens ............. 249 
BralmeCarolym 2 s-....--.-- - 64, 123 
nae NMlanalivad Sse... |. oe 177 
Caldwell, David K............ 37,129 
Caldwell, Melba C. .......... 37, 129 
Campbell, Howard W............ 193 
(GarzerMeNV IVA Ais om 2 eee RS 65 
Clauilinewe, Geos We oeoee con neue « 45 
Comstock, John Adams .......... 3 
IDENweoIL, 1h, VENI soecseoecneosbee 153 
Drakes Garlic. sac ade vik 133 
Drake, Robert J. .............. 25, 44 
Dre lve ree ne yan a eae 217 
ATT SMAV VME eis cnc cioors ehh Ae 917 
GCrilorroval, ||@ aval Sih gies om ome codes 6 143 
Elertlemlkeo|George 7 .).5 54). 4. 446 247 


Yaldwyn, John C. . 


255 


Hilton, William A. ............. 205 
Flovwandssrnidesardemne see 297 
ILal IRinvenes, IFA os co caw ococoe oon es 185 
Loomis, Richard B............ 29,177 
Lumsden, William W............ 933 
IMiolare, Olin I, -oeds0cescc00ce 64, 123 
Mortenserm MEPAVE are eee 65 
Riexcew Va Dwacihityaeer ser ne 7, 143 
Inkoyeteras, IMMeme Ih, oo 6 on cb orn 008 5 ow) 
Ruhoff, Florence A. ............. 133 
Simmons, Robert S.............. 193 
Stephens. Roberti aes ar: 29 
Susulksilakecomnenr ase eee aL oO 
Medford RichardeHees 55 oso ase 113 
Tinkham, Ernest R.............. 89 
Waldron, William G............. 244 
Wood, Sherwin E........... 161, 246 
ROM are 5 gts 15 


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oa AF 4 
BULLETIN OF THE 
Southern California sso 


BOTANICAL 


Academy of Sciences EN 


1963 


‘ LOS ANGELES, CALIFORNIA 


January-Marcu, 1963 Part 1 


CONTENTS 


Orchestoidea gracilis, a new beach hopper (Amphipoda: Talitridae) 
from Lower California, Mexico, with remarks on its lumines- 
peucewe. f Bousfeld and Wak, Klawe\). 0... 62. oie. 1 


The late Pleistocene 150 foot fresh water beach line of the Salton 
NearancanRoOvert GMWHOTILOSH ee eigen ae 


CC CC TN OC TT TT 


Notes on the larva and pupa of Euphydryas eurytion (Lepidoptera, 
INyauplalidae)is@ omas C."Ermel 0... eo ges 


In Memoriam to Bess Reed Peacock. Bonnie C. Templeton ........ 


Further studies on the benthic fauna in a recently constructed boat 
harbor in southern California. Donald J. Reish .............. 


Notes on the life histories of two southwestern phalaenid moths. 
Wimlere id airs (GorestOChjaer teh. CAEL. oles 3 he sia lslaye vps ate, «ye 33 


The early stages of Pero macdunnoughi Cassino and Swett (Lepi- 
doptera, Geometridae). John Adams Comstock .............. 


Issued April 1, 1963 


Southern California 
Academy of Sciences 


OFFICERS 
Theodore Downs 2.00/25 < lsiecieove «1s © eceleleieiel sticks = ote 11) ee President 
Richard.B. Loomis))..2. 2). ce:02 nes osteysckin se eee First Vice President 
John As White aio e275 Saree | Ue ete cn cian me Second Vice President 
Gretchen Sibley. .2).'¢.. 0. cinjb's sierete te oie « ayelshecorelentia </o lcie: «cee ene Secretary 
Tloyd Mi. Martin fers. sicjerarie stereo oir teers cnetternonelene Assistant to Secretary 
Wi Dwight Bierce ej.) </cjcteje ce qalefelce aie cet la.e)-/aye -ls)el icici Treasurer 
David K. Caldwell .2:..5 057%. cu 5. cons notes «fe 6 0 ie ee Editor 
DIRECTORS 
Russell E. Belous Charles A. McLaughlin Ruth D. Simpson 
Theodore Downs W. Dwight Pierce Peter P. Vaughn 
Richard B. Loomis Jay M. Savage John A. White 
Lloyd M. Martin Gretchen Sibley 
ADVISORY BOARD 
M. Dale Arvey Herbert Friedmann Kenneth E. Stager 
A. Weir Bell Hildegarde Howard Richard H. Swift 
J. Stanley Brode Theodore Payne Fred S. Truxal 
David K. Caldwell James A. Peters Louis C. Wheeler 
John A. Comstock Elbert L. Sleeper Sherwin E Wood 


STANDING COMMITTEES 


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SCIENCE SECTIONS 


Anthropology Botany 

Charles E. Rozaire, Chairman Richard G. Lincoln, Chairman 
Earth Sciences Experimental Biology 

Peter P Vaughn, Chairman Ronald Kroman, Chairman 
Invertebrate Zoology Vertebrate Zoology 

Elbert L. Sleeper, Chairman Dennis G. Rainey, Chairman 


The Bulletin is published quarterly by the Academy. Address all communications to the 
appropriate officer at the Los Angeles County Museum, Los Angeles 7, California. 


Printed by Anderson, Ritchie & Simon, Los Angeles, California 


BULLETIN OF THE SOUTHERN CALIFORNIA 
ACADEMY OF SCIENCES 


Vou. 62 January-Marcu, 1963 Part 1 


ORCHESTOIDEA GRACILIS, A NEW BEACH HOPPER 
(AMPHIPODA: TALITRIDAE) FROM LOWER CALIFORNIA, 
MEXICO, WITH REMARKS ON ITS LUMINESCENCE 


E. L. BousFiELp 
National Museum of Canadat 
and 


W. L. Kiawe 


Inter-American Tropical Tuna Commission? 


During a recent research cruise “La Blota” of the Inter-American 
Tropical Tuna Commission in Mexican coastal waters, Mr. James 
Joseph, Mr. Enrique Diaz, and the junior author made a small night 
collection of beach hoppers on a sandy beach at Cabo San Lucas, Baja 
California (22° 53’ N, 109° 53’ W). The animals were easily located 
through their luminescence as they hopped about on the wet sand near 
the water’s edge. They were collected individually with the aid of a 
flashlight. Upon close examination the amphipods were found to be 
distinct from any of the eight species of beach hoppers previously 
known from the Pacific coast of North and Central America (Bousfield 
1957, 1960) and from other species of the genus Orchestoidea and are 
herewith described as Orchestoidea gracilis n. sp. 

The present species is closely related to Orchestoidea meridionalis 
Schuster from E] Salvador and O. biolleyi Stebbing from the Pacific 
coast of Costa Rica. The three species may constitute a natural sub- 
group within the genus Orchestoidea as presently defined. This tro- 
pical and warm-temperate subgroup differs from the cool-temperate 
typical forms (e.g. O. twberculata, O. pugettensis, O. californiana) in 
having relatively long and slender body and limbs, shallow head, 
pronounced inferior antennal sinus, large maxilliped palp, complex 
and specialized peraeopod dactyls (especially in peraeopod 3), elon- 


1Ottawa, Canada. 


2ZLa Jolla, California. 


2 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


gate peraeopod 5, slender pleopods, non-telescopmg urosome, well-de- 
veloped prepeduncle in uropod 1, elongate uropod 3, and spade-shaped 
bilobed telson. Moreover. the group exhibits unusual features of sexual 
dimorphism, not only in the expected differences between antennae 2. 
gnathopods. and peraeopods, but also in the differential development 
of the rami of uropods 2 and 3. Most of these features are shared also 
by “Talorchestia” brito Stebbmg from western Europe and by Or- 
chestoidea brasiliensis Dana from the Atlantic coast of South America. 
a fact indicating that presently accepted generic criteria have outlived 
their usefulness. However, formal recognition of these relationships 
must await a revision of the entire beach-hopper complex within the 
terrestrial amphipod family Talitridae. 


Key to TroprcAL AMERICAN-PACIFIC SPECIES OF Orchestoidea 
1. Dactyl of peraeopods 4 and 5 with 4-5 stiff bristles along mner 
margin; pleopods very weak, rami with only 3-5 plumose seg- 
ments; uropod 3 of mature male longer than uropod 1 ......... 
ER a ae Peet ee os Orchestoidea meridionalis Schuster 


Dactyl of peraeopods 4 and 5 with single marginal bristle; pleopod 


rami with 7-14 plumose segments; uropod 3 of male short .... 2 

2. Outer ramus of uropod 1 smooth, inner ramus with terminal spine 
about half its length; pleopod rami with 7-8 plumose segments: 
uropod 2 of mature male, inner ramus much longer than outer 
sr Guacbtesya teen er enti SE Orchestoidea biolleyi Stebbing 
Outer ramus of uropod 1 armed with small outer marginal spines. 
inner ramus with terminal spine about one-quarter its length: 
pleopod rami with 11-15 plumose segments; uropod 2 of mature 
males ranm subequal’ ee sige Orchestoidea gracilis n. sp. 

ABBREVIATIONS FOR FIGs. 

A1 Antenna 1 Mxpd Maxilliped 

A2 Antenna 2 Gnt Gnathopod 1 

Hd Head Gn2 Gnathopod 2 

Es E ee HP P1-5 Peraeopods 1-5 

shu sower 1p : A : l : ] tes 11-3 

Rt.Md. — Right Mandible ae ae devs: 

Lft.Md. Left Mandible see teopedsite 

Mx1 Maxilla 1 U1-3 Uropods 1-3 


Mx2 Maxilla 2 ar Telson 


New Beach Hopper From Mexico 3 


Family Taurrriar Bulycheva 1957 
Genus Orchestoidea Nicolet 1849 


Orchestoidea gracilis, new species 


Description: Male (17.5 mm.). Head about as deep as long, convex 
above; inferior antennal sinus well incised. Eye very large, black, 
sub-trapezoidal, more than half length of head. Antenna 1 short, 
peduncular segments subequal, flagellum of 6-7 short segments. An- 
tenna 2 very long, about equal to body length; flagellum much longer 


Figure 1. Orchestoidea gracilis new species from Cabo San Lucas, Baja Cali- 
fornia, Mexico. 1. Male, 17.5 mm, 2. Female, 14.5 mm. 


4 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


than peduncle. of about 45 short untoothed segments, peduncular seg- 
ments strong but not powerfully expanded, surfaces spinulose. 

Upper lip slightly wider than deep, apically pilose. Lower lip with 
prominent lateral lobes, densely pilose along inner margin. Mandible. 
cutting edge with 4-6 teeth; left lacinia is 4-cuspate, right lacinia is bifid; 
molar process finely striate (about 30 striations). Maxilla 1, apical 
spine-teeth of outer lobe relatively long and slender, all pectinate, a 
row of fine bristles at base of inner spine teeth; palp minute, appearing 


Figure 2. Orchestoidea gracilis new species from Cabo San Lucas, Baja California, 
Mexico. 1. Male, 17.5 mm. 2. Female, 14.5 mm. 


New Beach Hopper From Mexico B 


one-segmented; inner plate with rather short plumose apical setae. 
Maxilla 2, inner lobe with strong proximal plumose seta; outer lobe 
larger, apically rounded. Maxilliped, mner lobe apically sub-truncate, 
with 3 subequal conical spine-teeth and about 8 marginal plumose 
setae; outer lobe apically rounded, extending well beyond inner lobe, 
outer apical setae are plumose, inner setae are minutely pectinate; 
palp very large, 3-segmented, outer marginal spines with compound 
ups. 

Coxal plates 2-4 successively broader, each with distinct posterior 
lobe or knob. Coxa 5 much longer than deep, anterior lobe a little 
larger than posterior lobe. Gnathopod 1, coxal shelf prominent, forming 
a complete inner lower margin armed with long stiff spines; segment 
5 with large and prominent pellucid process below; segment 6 rather 
broad, lower margin distally tumid and armed with heavy compound 
spines; dacty] strong, inner margin pectinate. Gnathopod 2 very large 
and robust; propod (segment 6) sub-ovate, spinose palm very oblique, 
slightly convex, with spinulose knob near base of dactyl, posterior 
angle defined by twin prominences each armed with 4-5 stout spines; 
dactyl strong, inner margin lined with minute spinules and a spinulose 
knob near hinge. 

Peraeopods generally slender, anterior and posterior margins well 
armed with compound spines. Dactyls are relatively short, nails very 
small, outer margin convex, well chitinized, and minutely pilose dis- 
tally, inner margin with single stiff bristle. Dactyl of peraeopod 2 with 
prominent spur on inner margin. Dactyl of peraeopod 3 very short, 
inflated and strongly convex behind, nail vestigial. Anterior marginal 
spines of segment 5 of peraeopod 3 not exceptionally long and strong. 
Peraeopod 5 distinctly longer than peraeopod 4, especially in segment 
6. Coxal gills small, sac-like, somewhat elongate on peraeopod 4 and 
enathopod 2. 

Pleosome side plates 1-3 smoothly rounded below, posterior margins 
lined with 4-6 minute spinules. Pleopods 1 and 3 subequal, 2 largest; 
peduncles slender throughout, not laterally expanded, outer margin of 
2 and both margins of 3 sparsely spinose; two small coupling spines; 
rami subequal, shorter than peduncles, each with 11-15 plumose 
segments. 

Urosome segment 2 distinct, not telescoping dorsally with segment 1. 
Uropod 1 with strong prepeduncle; rami slender, subequal, nearly 
equal to peduncle, outer ramus with outer marginal spines only, ter- 
minal spines not spade-tipped nor exceptionally long. Uropod 2, rami 
and peduncle subequal, inner ramus with both margins spose, outer 


6 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


ramus with inner margin bare. Uropod 3 somewhat elongate, ramus 
slightly longer than peduncle, not laterally compressed, posterior 
margin minutely spmose, apex with tuft of short slender spines. Telson 
spade-shaped. longer than wide, dorsally and apically spinose, consist- 
ing of two medially fused lobes. 

Female (14.5 mm.). Generally smaller and more compact than 
male. Antenna 1 much shorter, flagellum of 29-30 segments, slightly 
longer than peduncle. 

Gnathopod 1 strong, blister lackmmg on lower margin of segments 
5 and 6. Gnathopod 2. segment 2 moderately expanded in front, an- 
terior margin sparingly spinose, distally smuous or indented; segment 
4 with short posterior lobe; segment 5 rounded beneath, upper margin 
minutely spinose; segment 6, “mitten” lobe extending well beyond 
dactyl. minute lateral spines rather sparse. Brood plates on segments 
2-4 rather slender throughout, somewhat linear, with about 12 simple 
marginal setae; that of segment 5 shorter but broader, with fewer 
apical setae. 

Uropod 2, rami subequal. Uropod 3, ramus and peduncle subequal. 
Telson short, shghtly longer than wide. 

Material Examined: Cabo San Lucas, Territorio Sur de Baja Cali- 
fornia, Mexico, on wet sand at water’s edge. LW level. May 31, 1962. 
W.L. Klawe, J. Joseph. and E. Diaz coll. 

Male (HOLOTYPE), Female (ALLOTYPE), NMC No. 6634; 
7 female, 1 mmm. male PARATYPES, NMC No. 6635. 


HABITAT 


The beach hoppers were collected on the Cabo San Lucas beach be- 
tween one-half and one mile east of the Cabo San Lucas Pier. The ani- 
mals were hopping on the wet. medium-to-coarse. arkosic sand. 


LUMINESCENCE 


Some individuals of Orchestoidea gracilis were luminescent, which 
aided in their location and capture. The entire body of the animals 
appeared to emit light. 

There are several reports on luminous gammarids (sensu lato). 
According to Harvey (1952) the oldest is perhaps that by Thulis and 
Bernard, published in 1786, of a fresh water species from southern 
France. The earliest report on luminous marine gammarids is most 
hkely that of Viviani (1805). Tilesius (1819) presents a figure of a 


New Beach Hopper From Mexico 7 


luminous marine amphipod and Van Vollenhoven (1860) reports on 
luminous Orchestia littorea. 

Giard (1889, 1890) and Giard and Billet (1889) are to be credited 
with demonstrating the bacterial origin of luminescence in beach fleas. 
They moculated non-luminous individuals of Talitrus, Orchestia lit- 
torea, Hyale nilssoni, and even some other crustacea, with light-pro- 
ducing bacteria from Talitrus. According to the above authors, the in- 
fection leads ultimately to death. 

In 1927 Inman reported the bacteria-induced luminescence of Talor- 
chestia longicornis and Orchestia platensis. He examined about 20,000 
individuals, mostly of 7! longicornis. The contents of the digestive 
tracts of some of the non-luminous individuals revealed the presence 
of a bacterium, causing the luminescence, which he referred to as 
Bacterium giardi. Although present in small numbers, it was an 
ubiquitous component of the intestinal flora of the animals he in- 
vestigated. He postulated that under certain conditions these bacteria 
invade the muscles of the animal and increase in number so rapidly 
that the animal becomes luminous and finally dies. He pointed out the 
possibility that the bacteria may sometimes cause the death of a beach 
hopper without its becoming luminous. 

Similarly, in the case of Orchestoidea gracilis collected in the area of 
Cabo San Lucas, we are dealing not with a self-induced luminescence 
but with one caused by luminous bacteria. Three facts lead to this 
conclusion: (1) The origin of light was not restricted to any particular 
part of the body, but the entire animal was glowing, (2) there is no 
morphological evidence that the animal possesses special light-produc- 
ing organs, and (3) not all of the hoppers observed and collected were 
luminous. 

Of pertinent interest is our observation that, at the water’s edge 
where O. gracilis was collected, wave-cast fish bones emitted a faint 
glow. As beach hoppers in general are scavengers, the animals pos- 
sibly become infected through ingestion of foodstuff on which the 
luminous bacteria are growing. 

For a more detailed discussion, and also for a summary of reports 
and investigations on luminescence of amphipods caused by bacteria, 
we refer the reader to the classical treatise on bioluminescence by 
Harvey (1952). 


8 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 
LITERATURE CITED 


BOUSFIELD. E. L. 
1957. Notes on the amphipod genus Orchestoida on the coast of North America. 


Bull. So. Calif. Acad. Sci.,58(3): 119-129. 
1960. New records of beach hoppers (Crustacea: Amphipoda) from the coast of 
California. Bull. Natl. Mus. Canada, no. 172, pp. 1-12. 


GIARD, A. 

1889. Sur V’infection phosphorescente de talitres et autres crustacés. Comptes 
Rendus Hebdomadaires des Séances et Mémoires de la Société de Biologie, 
Paris, 109: 503-506. (Translation: On the phosphorescent infection of the 
talitri and other crustaceans. Ann. Mag. Nat. Hist., 6(4): 476-478.) 

1890. Nouvelles recherches sur les bactéries lumineuses pathogenes. Comptes 
Rendus Hebdomadaires des Séances et Mémoires de la Société de Biologie. 
Paris, 2(ser. 9): 188-191. 

GIARD, A., anv A. BILLET 

1889. Observations sur la maladie phosphorescente des talitres et autres crus- 
tacés. Comptes Rendus Hebdomadaires de Séances et Mémoires de la Société 
de Biologie, Paris, 1(ser. 9): 593-597. 


HARVEY, E. N. 
1952. Bioluminescence. New York: Academic Press, xit+649 pp- 


INMAN, O. L. 
1927. A pathogenic luminescent bacterium. Biol. Bull... 53: 197-200. 


SCHUSTER, O. von 
1954. Zwei neue Crustaceen von der Pazifischen hKiiste Mittel-Amerikas (Am- 
phipoda und Isopoda). Senckenbergiana, section Biologica 35(%): 103-105. 


STEBBING, T. R. R. 
1908. A new amphipod crustacean, Orchestoidea biolleyi, from Costa Rica. Proc. 
U.S. Natl. Mus., 34: 241-244. 


TILESIUS von TILENAU, W. G. 

1819. Leuchten des Meeres (arranged by L. W. Gilbert). Annalen der Physik. 
61: 36-44, 142-176, 317-330. 

VAN VOLLENHOVEN, SNELLEN 

1860. De Dieren van Nederland. Natuurlijke Historie van Nederland. vol. 1. 
Haarlem: A. C. Kruseman, 296 pp. 

VIVIANI, DOMINICO 


1805. Phosphorescencia maris. Quattuoredecim lucescentium animaculorum novis 
speciebus illustrata. Genoa: J. Giossi, 17 pp. 


THE LATE PLEISTOCENE 150 FOOT FRESH WATER BEACH 
LINE OF THE SALTON SEA AREA 


Ropert G. THomas'! 


Los Angeles, California 


INTRODUCTION 


The physical situation in the Salton Sea region in recent years will be 
briefly discussed in order to provide some ideas which are important 
in the interpretation of a 150 foot fresh water beach line which is re- 
ported here for the first time. Earlier lake levels will then be discussed, 
followed by a brief description of sea level changes, and finally by a 
tentative outline of Pleistocene events. 

Through the early years of recorded history of this region, what is 
now called the Salton Sea was a dry lake bed. In 1905, however, the 
Colorado River inundated the area, and the sea which it formed re- 
mains today, even though the major river flow was turned back toward 
the Gulf of California in 1907. 

Since the forming of the Salton Sea, it has not been connected with 
the Gulf of California, even though the historic high water level in 
1907 was still far below sea level and the waters of the sea have pres- 
ently the approximate composition of sea water. It is separated from 
the Gulf of California by the southwestward trending ridge of the 
Colorado River Delta (Fig. 1.). The lowest natural elevation on the 
delta is about 30 feet and is reported to be located at Volcano Lake 
southeasterly of Mexicali near the north end of the Rio Hardy shown 
on Figure 1. The Colorado River in pre-historic times has flowed south 
to the Gulf and at times to the north. It is likely that it has flowed in 
both directions at the same time, breaking into two or more rivers. The 
depositional processes have built a delta with the north slope consider- 
ably steeper than the south slope although the difference is apparent 
only on a profile with greatly exaggerated scale. 

The prominent shoreline of an ancient lake at about 40 feet above 
sea level in the Salton Sea area has been discussed by many authors. 
This lake, variously called Lake Coahuila, Lake Cahuilla, Lake Le 
Conte and Blakes Sea, extended from the vicinity of Volcano Lake 
some 105 miles north to the vicinity of Indio, and was 35 miles wide 
and about 300 feet deep. The shoreline features are well preserved and 


'Thomas M. Stetson, Civil and Consulting Engineers. 
215 W. 7th Street, Los Angeles 14. 


10 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


SELECTED FEATURES OF THE 
SALTON SEA REGION 


4 PALO VERDE 
VALLEY 


QO 
° 
» “}) 
Te. 
J 
°° 


Brawley @ a ee 


PERIAL VALLEY; 3} 
. -\.? 


DELTA 


| 
-( 
\ 


\\ 


Yuma esa [ 


2p e11aISg 


‘ 


Apaeyy ory 
gure uno eto 


zaie 


rs 
wh 
Re 
SP Az) 
Zen 
a = 


Desierto 


—— 


Figure 1. Selected features of the Salton Sea region. 


Pleistocene beach line 11 


little weathered. Fresh water fossil molluscs are abundant around its 
higher shorelines. Radiocarbon dating indicates that the forty foot 
beach line was last occupied only about 300 years ago, but that it 
existed at least 1600 years ago (Hubbs et al., 1960). Hubbs and Miller 
(1948) discuss many of the problems concerning this ancient, though 
relatively recent, lake. 

Much work remains to be done on this phase of the geologic history 
of the region, but the purpose of this note is to discuss the significance 
of even higher shorelines near the present Salton Sea. 

Figure 1 shows many of the geographic and physiographic features 
in the region of the Colorado River Delta. Figure 2 is a more detailed 
sketch of the location of the preserved 40, 100 and 150 foot beach lines. 


150 aANp 100 Foor SHORELINES 


Brown (1923) suggested that there are higher shorelines than the ob- 
vious and extensive 40? foot shoreline around the Salton Sea. Since the 
author had never observed any obvious higher shorelines in the region, 
a search was started for them on new U.S. Geological Survey 1/24,000 
scale topographic maps. Features resembling beach bars were noted 
on the maps at an elevation of about 150 feet above sea level. They are 
located west of El Centro and extend about 15 miles from the Mexican 
border to north of Highway 80. These were found to be fresh water 
beach bars. 

Figure 3 shows the gravel pits which contain fresh water fossils at 
the 150 foot beach line west of El Centro near Highway 80. These 
fossils, identified by the author, are the same kind as found near the 
40 foot shoreline, but here they are pitted and poorly preserved. Sub- 
surface caliche deposits shown on Figure 4 are very well developed, 
but no surface tufas or lime deposits were noted at those areas visited. 
The surface soil is weathered to a reddish color and the pebbles above 
and below the beach line are thickly coated with desert varnish. Most 
of the beach bars are discontinuous and have been partly dissected by 
stream erosion. These conditions suggest considerably greater age than 
the well-preserved 40 foot beach line. A poorly preserved remnant of 
a beach bar was also noted at about 100 feet above sea level. Except 
for the presence of fresh water fossils, these older beach bars closely 
resemble older bars found in closed desert basins in the Great Basin 
region. 


240, 100 and 150 hereafter refer to approximate elevations above sea level 


12 


Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


40 FOOT BEACH LINE 
35 1 
Sit CSs aaa 


24. 


| - VN ' 


OT BEACH 


150 FO 


So 
345 


- Pa eral 
$2 
? 


100 FOOT BEACH LINE >)” 


Shy es 


by 
0 
4 
fo) 
i= 
< 
18) 
eo) 
4 
f) 
ial 
< 
z 
ad 
(2) 
4 
Aa, 
a 
< 


BEACH LINES WEST OF 


SCALE IN MILES 


EL CENTRO 


A CALIFORNIA 


—_— 
Wad 


Figure 2. Approximate location of beach lines west of E] Centro, California. 


Pleistocene beach line 13 


Figure 3. Gravel pit north of Highway 80 about 1.4 miles west of Plaster City. 
Hammer leans against gravel containing pitted and poorly preserved fresh water 
molluses. 


SEA LEVEL CHANGES 


Studies in coastal southern California (Thomas, 1961) and in other 
parts of the world (Russel, 1957), mdicate that effects of world-wide 
changes of sea level on stream systems has been considerable. Erosion 
of considerable lengths of alluvial filled valleys near the coast occurred 
during the period of Wisconsin low sea level. These eroded valleys 
have refilled as the post-Wisconsin sea level rise occurred (Shepard 
and Suess, 1956). The sea level changes must have also affected the 
Colorado River. 


TENTATIVE OUTLINE OF QUATERNARY GroLocic History 


The Pliocene and possibly Miocene marine Imperial formation was 
deposited from the Gulf of California to the north of the present Salton 
Sea while alluvial fans were bemg deposited from the nearby moun- 
tains. These mountains were probably considerably lower than they 
are today. Formations deposited in late Pliocene and in early and 
middle Pleistocene times include parts of the Canebrake conglomerate, 


14. Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


Figure 4. Gravel pit south of Highway 80 (in distance) about 1.4 miles west of 
Plaster City. Clay and silt layers are shown near the bottom of the cut. The upper 
portion of the cut is gravel with thin horizontal caliche layers and thick vertical 
caliche columns. 


the Palm Spring, Brawley and Borego formations and possibly part 
of the sediments mapped as the Ocotillo conglomerate (Dibblee, 1954). 

In the Vallecitos Creek area south of Borego Valley, Downs (per- 
sonal communication) and Downs and Woodward (1961) have re- 
ported a vertebrate fauna of Irvingtonian age (early Pleistocene of 
most California geologists at the time of this writing, but middle Pleis- 
tocene of vertebrate paleontologists) throughout the upper 2500 feet 
thickness of the Palm Spring formation. The vertebrate fauna indi- 
cates that the area was a prairie grassland with wooded areas near 
stream margins (White and Downs, 1961:32). These conditions sug- 
gest that more rain fell than at present in that now desert area. A sim- 
ple explanation would be that the Peninsular ranges were lower, allow- 
ing a greater moisture flow over them, as has been suggested for the 
Sierra Nevada (Axelrod, 1958; Putnam, 1960). Downs and Woodard 
(1961) also report at least one thin marine shell layer in these sedi- 
ments which suggests a lagunal embayment. This is confirmed by 
Arnal (1961) from microfossils in the Borego formation. The possi- 


Pleistocene beach line 15 


bility that these marine organisms may have lived in ancient, closed 
saline lakes, as some species do now in the Salton Sea, has not been 
adequately explored. 

The available data, then, suggest that the Salton Sea area was, from 
time to time, mundated in limited areas by the Gulf, and that at cther 
times a fresh water lake or lakes existed, bordered by extensive alluvial 
fans. The considerable thickness of these early and middle Pleistocene 
sediments exposed around and buried beneath the Imperial and Coa- 
chella Valleys and the presence of the many large fault systems, includ- 
ing the San Andreas, suggest that tectonic activity was continually in 
progress during deposition. 

As a result of erosion due to uplift, large pediments or eroded sur- 
faces with little or no alluvial cover were formed on the earlier Pleisto- 
cene folded sediments by late Pleistocene time. Some large alluvial 
fans related to pediment forming were deposited near the mountains. 
Remnants of both pediments and fans are still abundant along the 
western mountains from the Salton Sea to the Mexican border and 
along the eastern mountains in the vicmity of the Salton Sea. 

The Colorado River had no doubt entered the scene prior to this 
period of folding and erosion. The first clear evidence of its appearance 
is recorded by the 130 foot terrace on Palo Verde Mesa near Blythe, a 
part of the East Mesa near Yuma, and part of the Gran Desierto. These 
are probably remnants of a more widéspread alluvial deposit which 
extended unbroken to the Gulf of California. The deposit probably 
formed a delta some 100 feet higher than the present delta surface. 
The reason for the higher: surface may well have been the higher 
eustatic sea level durmg Sangamon interglacial time. The Colorado 
River must have flowed both directly to the Gulf and to the Salton Sea 
region forming a southwesterly trending nose or ridge. 

When the ancient delta had built up to about 150 feet above present 
sea level, one of many fresh water lakes was formed. Remnants of the 
eastern shore line may be buried under the Algodones sand hills east 
of El Centro, but the features west of FE] Centro described above are 
still preserved. 

With the onset of Wisconsin glaciation, the Colorado River probably 
had greater runoff than at present. World-wide sea level fell, causing 
erosion of the ancient delta. Much of the material of the Algodones 
sand dunes may have originated as the lake level fell and exposed the 
beaches. The 75-foot terraces at both Palo Verde Mesa and Yuma Mesa 
may be erosion features formed during a relatively stable period. It is 


16 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


possible that the very poorly preserved 100 foot beach line (Figure 2) 
may represent a lake level corresponding to this stable period. 

Sea level eventually fell to about 300 or 400 feet below present sea 
level during the Wisconsin Glacial period and the Colorado River 
eroded great quantities of the old delta away as it adjusted to the lower 
sea level. It is possible that the delta was completely removed in the 
present Imperial Valley, allowing the Whitewater River to flow to the 
Gulf. Such a condition may help explain the lack of recent sediments 
on both the east and west sides of the Salton Sea, where the folded and 
eroded Borego and Brawley formations are exposed or covered by only 
a few feet of younger material over wide areas. During this period of 
downcutting, most of the alluvial fans previously deposited and sedi- 
ments previously formed were deeply eroded and partly removed. 
Remnants of alluvial fans of similar age to that on which the 150 foot 
beach is found may be seen along the mountains on both sides of the 
Salton Sea, but only at considerably higher elevations. Some prelimi- 
nary work with air photos and in the field clearly shows that the fans 
are of various ages. The various aged fans can be recognized and rea- 
sonably correlated over much of the region by their colors in photos 
and by geomorphic relationships. 

As the Wisconsin glacial period ended, sea level once again rose. 
The Colorado River, adjusting to this rise, deposited materials once 
again. Basal gravels were deposited in Palo Verde Valley and in Yuma 
Valley and then finer materials covered them, finally forming the 
present delta surface. 

Ancient lakes probably existed from time to time throughout the 
various Pleistocene phases described above, their salinity varying with 
rate of inflow (depending upon the vagaries of the Colorado River), 
outflow and evaporation. The most recent pre-historic lake, with the 
40 foot shoreline, was probably formed as the delta accumulated dur- 
ing post-Wisconsin sea level rise. It existed up to the 40 foot beach line 
only a few hundred years ago and dried up when the Colorado River 
flowed directly south to the Gulf of California. 

The 150 foot shoreline is essentially at the same elevation in the 
few mules of its known exposure. It appears that tectonic activity since 
its formation has been relatively small compared with that which 
resulted in the major middle or late middle Pleistocene unconformity. 
The meager regional physiographic evidence available does not sug- 
gest large regional upwarping of the 150 foot shoreline, although it 
remains a distinct possibility. 


Pleistocene beach line 17 


It is hoped that this brief note will stimulate mterest in the Pleisto- 
cene problems of the region and encourage the collecting of evidence, 
not only in the Salton Sea region, but also in the Colorado River drain- 
age further upstream, and in the northern part of the Gulf of Califor- 
nia. The events described here should have left much evidence in these 
other areas. 


LITERATURE CITED 

ARNAL, ROBERT E. 

1961. Limnology, Sedimentation and Microorganisms of the Salton Sea, Cali- 
fornia. Bull. of Geol. Soc. Amer., 72:427-478. 

AXELROD, D. I. 

1958. Pliocene Verde Flora of Western Nevada. U. of Calif., Pub. in Geol. Sci., 
34(Q2). 

BROWN, 'S. J. 

1923. The Salton Sea Region, California. U.S.G.S. Watter Sup. Paper 497. 

DIBBLEE, THOMAS W, JR. 

1954. Geology of the Imperial Valley Region, California. Bull. Calif. Div. Mines, 
170, Chap. IT, Cont. 2. 

DOWNS, THEODORE anv G. DAVIDSON WOODARD 

1961. Middle Pleistocene Extension of the Gulf of California into Imperial Valley. 
Paper presented at a meeting of the Geol. Soc. Amer. Meeting at San Diego. 
Calif. 

HUBBS, CARL L. anp ROBERT R. MILLER 

1948. Correlations between Fish Distribution and Hydrographic History in the 
Desert Basins of Western United States in the Great Basin. Bull. U. of Utah, 
38(20). 

HUBBS, CARL L., GEORGE S. BIEN ann HANS E. SUESS 

1960. La Jolla Natural Radiocarbon Measurements. Amer. Journ. Sci., Radio- 
carbon Supp., 4:197-223. 

NORRIS, ROBERT M. anp KENNETH S. NORRIS 

1961. Algodones Dunes of Southeastern California. Bull. Geol. Soc, Amer., 
72:605-620. 

PUTNAM, WILLIAM C. 

1960. Faulting and Pleistocene Glaciation in the East-Central Sierra Nevada of 
California, U.S.A. Report 21st Sess., Intl. Geol. Cong., Part XX1:270-274. 

RUSSELL, R. J. 

1957. Instability of Sea Level. Amer. Scientist, 45 (5). 

SHEPARD, FEF P. ann H. E. SUESS 

1956. Rate of Postglacial Rise of Sea Level. Science, 123 (3207) : 1082-1083. 

THOMAS, R. J., J. J. LANDRY ann R. J. TURNEY 

1961. Ground Water Geology of the Coastal Plain of Los Angeles County. Bull. 
Calif. Dept. of Water Res., (104) App. A. 

WHITE, JOHN A. ann THEODORE DOWNS 

1961. A New Geomys from the Vallecito Creek Pleistocene of California. Los An- 
geles County Mus., Cont. in Sct., 42. 


18 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


ERRATUM 
The following correction should be made in the title of the article by Hildegarde 
Howard, “A fossil bird, Caracara, from Santa Rosa Island:’ which appeared in 


volume 61, part 4, of the Bulletin: 
The paper is number 36 in the “Contributions from the Los Angeles Museum 


Channel Islands Biological Survey?’ not number 34. 


NOTES ON THE LARVA AND PUPA OF 
EUPHYDRYAS EURYTION (LEPIDOPTERA, NYMPHALIDAE) 


THomas C. EMMEL 


Reed College, Portland, Oregon 


The checkerspot butterfly Euphydryas eurytion Mead and its forms 
range through the mountains and foothills of northern New Mexico 
to southern Wyoming. Although E. eurytion and its presumed parent 
species anicia are very widespread and fairly abundant in favored 
localities of the West, Dr. J. A. Comstock (personal communication ) 
states that nothing is known of the early stages of eurytion, and that 
the only information on record for the metamorphosis of a@nicia is an 
early paper on the larval habits (no description and no illustration ) by 
W. H. Edwards (1892). The following notes describe the present 
author’s observations on the last instar larva and pupa of E. eurytion. 

In Colorado, this member of the anicia complex is found from the 
southwestern mountains to the higher areas east of the Continental 
Divide (Brown, Eff, & Rotger, 1957). The E. eurytion larva to be 
described was collected in a dry meadow habitat (elevation 8,700 
feet above sea level) about one mile southeast of the ranch house at 
Big Spring Ranch, Florissant (Teller County), Colorado, on July 9, 
1962. When found, it was not on a plant but was crawling along the 
ground, Food-plant genera reported for closely-related EKuphydryas 
species were in the larva’s immediate vicinity, but a search of these 
plants (Penstemon and Castilleja) did not produce more larvae. 

Description of Mature Larva. Length, 26 mm. Ground color, ivory 
white, mottled irregularly with black. 

Black branching spines are present in the usual dorso-lateral rows. 
Unfortunately, more precise notes were not taken before the larva 
pupated on July 15. Before pupation, the larva shrunk to a length of 
about 17 mm. The imago emerged August 2, 1962. 

Description of Pupa. Length, 16 mm. Maximum width, 7.4 mm. 
Ground color, lustrous silvery-white, marked with black blotches and 
striations as shown in the illustration, Figure 1. 

Wing cases, silvery-white divided obliquely by a prominent broad 
band of black. Slightly tinged with orange-brown along the shoulders, 
around the black blotches, and along the wing tips. 

Antennal sheaths, heavily checked with black squares which are 
separated by very narrow white segmental lines. 


19 


20 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


= 


Co * cathe & js - wot at 
Figure 1. Lateral view of the pupa of Euphydryas eurytion, showing the dis- 


tinctive wing case markings. 


| 


ance 


RO A sn 


= 


; Figure 2. Dorsal view of the pupa of Euphydryas eurytion. 


Larva and pupa of Euphydryas eurytion 21 


Dorso-lateral nodules (representing vestiges of the larval spies) 
are orange with black blotches surrounding the nodules on all but the 
posterior aspects. 

Cremaster, dark brown at the end. 

From comparison with published figures of pupae of other Euphy- 
dryas species, it appears that the wing-case markings on this E. eury- 
tion pupa are the most distinguishing features, particularly with 
regard to the broad, continuous, obliquely-crossing band of black. 


LITERATURE CITED 


BROWN, E M., D. EFF, anv B. ROTGER 
1957. Colorado butterflies. Denver: Denver Museum of Natural History, 368 pp. 


EDWARDS, W. H. 
1892. Miscellaneous notes on butterflies, larvae, etc. Canadian Entomologist, 
24: 49-56. 


DP Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


IN MEMORIAM 
BESS REED PEACOCK 
1887-1962 


Bess Reed Peacock, a member of the Southern California Academy of 
Sciences since 1951, passed away on November 24, 1962, at her family 
home in Los Angeles. 

Miss Peacock was born in Independence, Missouri, on March 2, 1887, 
the only child of Judge J. Mack and Bettie Reed Peacock. Shortly there- 
after, the family moved to Roswell, New Mexico, where her father served 
as Circuit Judge for many years. 

Miss Peacock received her elementary and secondary education in 
Roswell, and there demonstrated her aptitude for instruction by teaching 
classes before she had finished high school. She attended the University 
of Missouri in 1906, and later went to the University of Chicago where 
she received her AB degree in education in 1912, with Phi Beta Kappa 
honors. 

In connection with her studies at the University of Chicago, she did 
welfare work in the settlement houses in Chicago, This led to her doing 
physiotherapy work in army hospitals during “World War I. 

She attended San Diego Teachers College in 1921, but returned to the 
University of Chicago to complete her MA degree in botany in 1923. 

When her father retired from his judgeship 1 in 1920, the family moved 
to Los Angeles. In 1923, after receiving her graduate degree, she taught 
fora dia time in Orange County hetore eee ing the lee Angeles City 
school system. In 1924, she taught at Roosevelt High School, rail then 
at Mt. Vernon Junior High Schools in 1926, where she taught science in 
the eighth and ninth grades until she retired in 1952. 

Bess Peacock had a lively interest in all things in science. This kept 
her constantly in the pursuit of knowledge w hich she passed on to her 
many students duri ing the approximately 35 years she served as a 
teacher. She earned her teaching credential in science at the University 
of Southern California in 1930, and became a member of Delta Kappa 
Gamma. She felt her greatest ability was in teaching boys and often 
preferred the supervision of “boys home room?’ 

She was active in the Southern California Academy of Sciences, the 
Southern California Botanical Society, the Nature Conservancy and 
nature study groups. Her other activities included Women’s University 
Club and the Los Angeles County Museum Association. 

Her chief interests were in nature everywhere and the cultural wel- 
fare of all her students. Her many financial gifts, most of them anony- 
mous, went toward conservation of nature, gave culturally deprived 
youngsters camping experiences, and promoted the advanced education 
of a number of w orthy students. Each student that she Sponsored and 
suided to higher education became her adopted “niece” or “nephew” 
during the period she assumed the obligation of his education. 

Her close friends and teaching colleagues cherish her for her dedica- 
tion to learning, integrity, and devotion to the truth as she saw it. 

Bonnie C. TEMPLETON 


FURTHER STUDIES ON THE BENTHIC FAUNA IN A 
RECENTLY CONSTRUCTED BOAT HARBOR 
IN SOUTHERN CALIFORNIA? 


Donatp J. REIsH 
Long Beach State College? 


INTRODUCTION 


In a previous study of a newly constructed harbor the author (Reish, 
1961) found no evidence that succession occurred in the subtidal ben- 
thic environment. The principal species were dominant during the 
entire 2.5 years of study. However, a decrease in the number of species 
and specimens occurred about 1.5 years after the study was initiated 
which was believed to be related to the effects of limited water circula- 
tion in Basin 1 (Fig. 1). A drop in the dissolved oxygen content of the 
water and subsequent appearance of sulfide odors to the sediments 
was cited as supporting evidence. In the course of dredging Basins 2, 
3, and 4 (Fig. 1), a channel was formed connecting one part of the bay 
(Fig. 1, Station I) with the Marine Stadium (compare Fig. 1, Reish, 
1961 with Fig. 1, herein). This provided an opportunity not only 
to verify whether or not benthic succession occurs, but also to determine 
whether or not a similar reduction occurred in an area with unim- 
paired water movements. The purpose of this paper therefore is to 
describe the benthic fauna in a newly established marine area possess- 
ing adequate water circulation. 

The subject of colonization of animals in the subtidal benthos has 
been reviewed by the author in the previous study (1961); included 
were the papers by Brandt (1897), Shelford, et al (1935), and the 
author (1956, 1957). 

The author wishes to thank Miss Emile Bender, Mrs. Bettye Byrne, 
Miss Ruth Zakem, Mr. Harold Pope, and Mr. Alfred Stone for their 
assistance in the collecting and sorting of the material. 


DESCRIPTION OF ALAMITOS Bay MARINA 


Alamitos Bay (Fig. 1) is a small body of water used primarily for 
recreational purposes, which is located within the City of Long Beach. 


1This study was supported by research grant NSFG-8914 from the National 
Science Foundation. 


“Department of Biological Sciences. 


23 


€ ulseg 


i 


| 


Basin 2 


MARINA 


ALAMITOS BAY 


ALAMITOS’ BAY and 
SAN GABRIEL RIVER 


ALAMITOS BAY 
MARINA 
440 


we 


é 


@y, Vj (gs 


of, 


Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


33°46 
118° 08 


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J 7 | 


ALAMITOS 


24 


California. The boat basins and station locations 


amitos Bay, 


Al 


imse 


Map of 


Figure 7. 


il 


are shown in the 


Benthic Faunal Studies 26 


Basin 1 was dredged in 1955, while the dredging of Basins 2, 3, 4, and 
the connection to Marine Stadium commenced August 1959. The 
marina was dredged to a depth of —12 feet and the main channel (Fig. 
1, Stations J, K, and R) to a depth of —15 feet. The sides of the marina 
consist of cement bulkheads and rock rick-rack. See Reish and Winter 
(1954) for the early history and development of the other parts of 
Alamitos Bay. 


MATERIALS AND METHODS 


Quantitative samples were taken with a size one Hayward orange 
peel bucket (Reish, 1959b). The samples were preserved m formalin, 
then washed later through a screen having 20 openings to the inch 
(0.85 mm.). The animals were separated and weighed. The data 
given in Figure 2 are wet weights and include structures formed by 
the animal such as shells of molluscans and tubes of polychaetes. Col- 
lections were made August 19, September 30, and November 17, 
1959, January 17, May 10 and August 31, 1960, January 6, and June 
29, 1961, and April 16, 1962. Currents were measured during July 
and August 1962, with a pygmy current meter (G. M. Manufacturing 
Co., New York). 


RESULTS 


The data have been summarized in Table 1, and Figures 2 and 3. 
Table 1 shows the principal species and the number of specimens taken 
each sampling period; the species of lesser incidence are given below. 
Figure 2 summarizes the occurrence and abundance of the principal 
bottom-dwelling species. Figure 3 represents graphically the number 
of species and specimens and the biomass for each survey. 

The first animals were not encountered until September 30, 1959, 
seven weeks after dredging began (Table 1, Fig. 2). No animals were 
collected during an earlier sampling period in August 19, 1959. Ad- 
ditional species appeared rapidly with a peak being reached about 10 
months later (Fig. 2). Fluctuations, possibly seasonal, were noted in 
the total number of species present following the initial rise. Addi- 
tional species not listed in Table 1 include the polychaetes: unidentified 
polynoids (5 specimens), Eumida sanguinea (Oersted) (3), un- 
identified phyllodocids (9), Exogone sp. (1), Platynereis bicanalicu- 
lata (Baird) (3), Boccardia uncata Berkeley (1), Polydora pauci- 
branchiata Okuda (2), unidentified spionids (10), Sprochaetopterus 
sp. (3), Polyophthalmus pictus (Dujardin) (1), Capitella capitata 


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28 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


(Fabricius) (3), Asychis disparidentata (Moore) (2), Aziothella 
rubrocincta (Johnson) (2), Owenia fusiformis collaris Hartman (1). 
unidentified ampharetids (3), unidentified terebellids (3), and 
Fabricia limnicola Hartman (1); crustaceans: Corophium acherusi- 
cum Costa (2), Caprella equilibra Say (1), unidentified amphipods 
(12), unidentified isopod (1), cumacean (1), shrimp (1), and crab 
(1); molluscans: Chione undatellum (Sowerby) (1). Lysonia califor- 
nica Conrad (1), Protothaca staminea Conrad (2), Tellina carpenteri 
Dall (1), unidentified juvenile clams (19), Bullaria gouldiana (Pils- 


bry) (3); the holothurian Leptosynpta albicans (Selenka) (3), un- 


OCCURRENCE AND ABUNDANCE OF DOMINANT SPECIES 


Eteone dilatae 


Nephtys caecoides 
Goniada littorea 


Lumbrineris) minima 
Dorvillea articulata 


Haploscoloplos elongatus 
Prionospio cirritera 


Chaetozone corona 
Tharyx parvus 


Cossura candida 
Armandia bioculata 
Amaeana occidentalis 


Nemerteans 


Tagelus californianus e 


Scale (| = 60 Specimens 


Macoma nasuta 


Capitita ambiseta 


Euchone limnicola 


Scale q = 150 Specimens 


Figure 2. Diagramatic representation of the occurrence and abundance of the 
dominant benthic species collected from the marina channel stations J, K, and R. 


Benthic Faunal Studies 29 


identified sea anemone (1), sipunculid (1), phoronids (11), and an 
ophiuroid (1). A total of 78 species were collected during 2.7 years of 
study of which 49 (63%) were polychaetes, 13 (17%) were mollus- 
cans, 9 (12%) were crustaceans, and the remaining 7 (8%) were 
sea anemones, turbellarians, nemerteans, sipunculids, phoronids, and 
echinoderms. 

The curve for the number of specimens collected each sampling 
period lagged the species curve during the first year (Fig. 2). A sharp 
decrease occurred between August 1960, and January 1961. This 
decrease was primarily the result of near disappearance of the poly- 
chaete Euchone limnicola Reish. The number of specimens taken 
the last two sampling periods were nearly identical. 

The growth of the animals proceeded slowly as indicated by the 
curve for biomass (Fig. 2). The drop in biomass following the peak in 
August 1960, is again the result of the decrease of Euchone limnicola. 


NUMBER OF SPECIES SPECIMENS & BIOMASS FROM THE CHANNEL OF ALAMITOS BAY MARINA 


Oy, pom species 


— Specimens 


8/59 9/59 11/59 1/60 5/60 8/60 vel 6/61 4/62 


Biomass in grams |18 


No. Specimens 
No. Species 


Figure 3. Graphic representation of the number of species, the number of speci- 
mens, and the biomass in grams collected each survey from the marina channel 
stations J, KR, and RB. 


30 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


DiscussIoNn 


As pr eviously found in Basin 1 of Alamitos Bay Marina, there was 
no indication of a succession of animal populations in the subtidal 
benthic bay environment. Since the collections that were made shortly 
after dredging was completed contained no animals, it is reasonable to 
assume that the animals taken in September 1959, were the first in- 
habitants of the area. One of these, Capitita ambiseta Hartman, con- 
tinued to be one of the principal species throughout the 2.7 years of 
study (Table 1, Fig. 3). 

All but two of the 44 more frequently occuring animals, Pinnixa 
franciscana Rathbun and Olivella biplicata Sowerby, were collected by 
August 1960. (Table 1). Since P. franciscana is a commensal crab 
that lives in the burrow of the ghost shrimp Callianassa californiensis 
Dana, it is evident that the appearance of P. franciscana must await the 
development and growth of a population of C. californiensis. It is 
possible that O. biplicata may be only an occasional inhabitant since 
it has not been reported in the previous studies of Alamitos Bay (Reish 
and Winter. 1954; Reish, 1959a. 1961). 

Several early inhabitants appeared, then later dropped out. These 
were the spionid polychaetes Polydora ligni Webster, Prionispio hetero- 
branchia newportensis Reish, P- pinnata Ehlers. and Nerinides pig- 
mentata. (Reish). None of these species was a dominant member at 
any time. The two former species are abundant in other parts of 
Alamitos Bay and the two latter are present offshore. 

The occurrence and abundance of the principal species have been 
summarized in Figure 3. These species may be grouped according to 
the characteristic feature of their curve, namely, species which reached 
a peak then decreased, species which show fluctuations (possibly sea- 
sonal), species which reached a plateau, and species which are ap- 
parently still increasing in numbers. 

Euchone limnicola, known only from Alamitos Bay, demonstrates 
mar kedly a build up in the population followed by sharp decrease in 
numbers. The number of individuals (1309) taken during August 
1960, strongly influenced the values for the number of specimens and 
the weight of biomass summarized in Figure 2, as discussed above. 
Four additional species may be included here, Eteone dilatae Hartman. 
Chaetozone corona Berkeley, Tharyx parvus Berkeley, and Macoma 
nasuta (Conrad). While not as many specimens of these species were 
collected in comparison to E. limnicola, the results are nearly identical 
to what was found previously in Basin 1. Perhaps these species can not 


Benthic Faunal Studies 31 


successfully compete with the other species or are unable to repopulate 
themselves in an established population. Many specimens of E. limini- 
cola were observed with eggs in their coelom, but it is not known 
whether or not spawning, fertilization, and development occurs in the 
marina. 

The polychaetes Nephtys caecoides Hartman, Dorvillea articulata 
(Hartman), and Haploscoloplos elongatus (Johnson ) show at least two 
population peaks followed by a drop in the number of specimens. With 
the latter two species the peaks apparently are seasonal with the highs 
occurring during the warmer months of the year. No seasonal con- 
clusions can be made with respect to V. caecoides. 

The population of possibly two species, Goniada littorea Hartman 
and Capitita ambiseta, is still increasing. The greatest number of spe- 
cimens for each species were collected at the time of the last sampling. 
Definite conclusions regarding these species must await further 
samples. 

Following the imitial rise, a more or less stable population has been 
reached by Lumbrineris minima Hartman, Prionospio cirrifera Wiren, 
Cossura candida Hartman, and the unidentified nemerteans. While 
fluctuations in numbers occurred with these species, it may be the 
result of either patchiness in distribution or collecting techniques. The 
conclusions regarding the unidentified nemeteans may not be valid 
since it is possible that more than one species is involved. 

Tagelus californica Conrad did not fit into any of the above four 
categories because one-half of the total number of specimens was col- 
lected during one sampling period. 

The findings reported herein differ from the previous study in that 
no marked drop off in the animal population was observed in the 
channel as was the case in Basin 1 and more recently (Reish 1962) in 
the innermost stations of Basins 2, 3, and 4 (Fig. 1, L, N, O, Q, S). 
This supports the earlier view that the decrease in the population is the 
result of limited water circulation. The dissolved oxygen content of 
the water drops and the substrate, which origially was gray im color 
and odorless, became black and possessed a strong sulfide odor. Ap- 
parently it takes about one year for the effect of limited water move- 
ment to alter the benthic environment of a newly established marine 
area. In the channel where the water movement is unobstructed, 
there is no marked change in the animal population. The decrease in 
the number of specimens between August 1960 and January 1961 
was largely the result of the near extinction of one dominant species 
rather than the decrease of specimens in all species of the population 


32 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


(Fig. 2 and 3). Currents have been determined to move at the rate of 
300 to 1000 feet per hour in the channel but no movements have been 
measured in the basins. 


SUMMARY 


1. A periodic, quantitative study was made in a newly dredged boat 
channel to determine whether or not succession occurs in the 
subtidal benthic environment. 


2. There was no indication that the settlement of an earlier assem- 
blage of animals was necessary for the settlement of a subsequent 
one. 


we) 


Variations in the population were noted. The principal species 
were grouped according to: (1) those that reached a peak in 
number of specimens then decreased, (2) those that showed 
fluctuations, possibly seasonal, (3) those that reached a plateau, 
and (4) those that apparently are still increasing in numbers of 
specimens present. 


4. The relationship between the amount of water circulation and its 
effect upon benthic animals in boat basins and channels was 
discussed. 


LITERATURE CITED 

BRANDT, k. 

1897. Das Vordringen Marine Thiere indem hkaiser-Wilhelm Canal. Zoologische 
Jahrbiicher Abteilung Systematik Geographie und Biologie, 9:387-408. 

REISE D: 

1959a. A discussion of the importance of the screen size in washing quantitative 
marine bottom samples. Ecology, 40:307-309. 

1959b. Modification of the Hayward orange-peel bucket for bottom sampling. 
Ecology, 40:502-503. 

1961. A study of benthic fauna in a recently constructed boat harbor in southern 
California. Ecology, 42:84-91. 

1962. A study of succession in recently constructed marine harbors in southern 
California. Proc, First National Coastal and Shallow Water Research Con- 
ference, Baltimore, Los Angeles, and ‘Tallahassee, pp. 570-573. 


SHELFORD, V. E., et al. 


1935. Some marine biotic communities of the Pacific coast of North America. 
Ecol. Monogr., 5: 249-354. 


NOTES ON THE LIFE HISTORIES OF 
TWO SOUTHWESTERN PHALAENID MOTHS 


JoHn ApAmMs CoMsTOCK 


Del Mar, California 


LACINIPOLIA QUADRILINEATA (GROTE) 


This small and contrastingly marked moth was described by Grote 
in 1873, as Mamestra 4-lineata, from ““California”’ 

Very little has been published concerning its life history, and I have 
no record of adequate illustrations. 

Dyar published a brief description of the larva in 1898, from ma- 
terial supplied by Koebele, “Los Angeles County, food plant not 
stated?’ 

Crumb, m 1956, p. 135, gave an excellent description of the mature 
larva, compiled from material obtained in Arizona, and recorded its 
range as California, Arizona, New Mexico, Nevada and Oregon. His 
food plant records were given as “taken among dead leaves and other 
plant debris about clover, Haplopappus heterophyllus, and other 
herbs”’ This suggests a sweeping operation rather than beating specific 
plants. 

During June of this year (1962) I did some extensive beating of 
Adenostoma fasciculatum H. & A., (Chamise) in Del Mar, California. 
Among other captures I took two larvae that were new to me. These 
were drawn, measured, and carefully noted. Later they pupated, and 
when they finally emerged, proved to be Lacinipolia quadrilineata. 
As this moth was coming in to black light at the time, I confined several 
females, and secured numerous eggs. This made possible the following 
notes. 


Ecc 


Hemispherical, the base flattened, and top rounded, with a clearly 
defined micropyle. Height, 0.5 mm. Diameter at base, 0.75 mm. The 
walls are covered with from 25 to 30 ridges which arise at the base 
and run towards the margin of the micropylar depression. Some of 
these ridges end considerably short of this margin. Others fuse with 
their nearest neighbor. The majority reach the edge of the micropylar 
depression. 


33 


34 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


eh MOLI OO AE OMe 


<a 


rere 


, 


os (0 apf we 


Mee Pes: 


On wet rb bongs 


Figure 1. The egg. mature larva and pupa of Lacinipolia quadrilineata. | A. egg. 
lateral aspect, tipped forward slightly to show micropyle. Enlarged < 40; B. egg. 
superior aspect, enlarged < 40; C. mature larva, lateral aspect, enlarged X ap- 


prox. 5: D. pupa, ventral aspect, enlarged X 7.] Reproduced from colored drawing 
by the author. 


Life histories of phalaenid moths 35 


The micropyle proper is a small crater-like structure in the center 
of this depression. Radiating from it are numerous fine ridges. These 
join the raised margin and help to form an elevated circlet along 
with the large ridges that converge on the outer wall. All of these 
ridges are white, and are topped with pearly nodules. The troughs 
between them are comparatively deep, and their floors are crossed 
transversely by low lines or ridges. 

The color of the egg when first laid is a pale yellow-green. Later 
the ground color changes to cream, with a large irregular patch of 
salmon-red on the floor of the micropylar depression, and a wide circlet 
of the same color covering most of the upper half of the egg. Some 
examples show a wider distribution of this salmon-red band. 

The egg is illustrated as Figure 1, A and B. 


Mature Larva 


Regularly cylindrical throughout most of the body, with tapering 
anterior segments and cauda. Length, 25 mm. 

Head; width, 2.75 mm. Color, dark gray, with brown and blackish 
mottling. It is retracted into the first segment when the larva is at rest, 
resulting im only a small portion showing, as recorded in the illustra- 
tion, Figure C. 

Body; dull mottled gray throughout. There is a suggestion of a 
longitudinal middorsal line, topped throughout its length by minute 
white spicules. A dorso-lateral line of the same character is also present, 
the latter being paired. 

The spiracles are very small, but are black, and therefore con- 
spicuous. 

The entire body is covered by minute spiculiferous points that are 
lighter in color than the ground on which they rest, and hence give the 
larva a frosted appearance. 

Near the dorso-lateral paired line there is a row of white setae, (two 
to each segment). A similar row of shorter setae occurs supraspira- 
cularly. The remaining setae are still shorter, and are black. 

The legs are gray with a tinge of yellow, and are heavily covered 
with short setae. The prolegs are concolorous with the body, and the 
crochets are brown. 

The larva is sluggish in its movements at all times. 

The two larvae pupated June 27 and 28 in a fragile cocoon on the 
floor of the rearing jar. 


36 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 
Pupa 


Length, 11.5 mm. Width through center, 4 mm. Color, red-brown, 
with a darker brown on the segmental junctures of the abdomen and 
the caudal tip. 

The head is evenly rounded, and the eyes large and protruding. 
The maxillae extend to the wing margins, and the antennae terminate 
about .5 mm. short thereof. 

The surface texture is smooth and glistening, except for the slightly 
granular surface of the eyes and the punctate surfaces of the caudal 
margins of the movable segmental junctures on the abdomen. The 
spiracles are concolorous with the body and relatively inconspicuous. 
The cremasteric tip bears two long thin spines without recurved tips. 

The pupa is illustrated as Figure D. 


AUTOGRAPHA BILOBA (STEPHENS ) 


This striking moth, which is illustrated in Holland’s Moth Book on 
Plate XXVIII, figure 24, and which has been shifted about in various 
genera, such as Plusia, Phytometra and Autographa, has received scant 
attention in the literature, so far as its life history is concerned. 

It ranges throughout the Americas from Canada to Argentina, and 
its larvae feed on several plants of economic importance, yet I have 
failed to find any worthy illustrations of its early stages in the litera- 
ture. 

The larva is a rather general feeder, having been reported on let- 
tuce, cabbage, tobacco, alfalfa, parsnip, clover, thistle and Collinsza. 
Apparently it is seldom abundant, in contrast to others of our pest 
insects. I have only taken a single example at Del Mar during many 
years of collecting. Fortunately this lone specimen was a female who 
rewarded me with a fine cluster of eggs. 

The most complete account of the species’ metamorphosis was pub- 
lished in 1882 by French. Hampson, in 1913, briefly mentioned 
French’s paper. Rosewall, in 1922, listed one of its food plants, and 
i 1956 Crumb published pertinent facts concerning its larva, seasonal 
occurrence and range. My notes supplement and amplify these pre- 
viously published records, and add illustrations. 

Eggs from the single female collected at Del Mar, April 6, were 
laid April 8, and hatched April 19 to 23, 1962. The young larvae made 
egress at the side of the egg, leaving the shell intact. They immediately 
took to lettuce. 


Life histories of phalaenid moths 37 


Figure 2. Early stages of Autographa biloba (Stephens). [A. egg, lateral view, 
slightly tipped to show micropyle, enlarged x 46; B. first instar larva, lateral 
aspect, greatly enlarged; C. mature larva, dorsal aspect, enlarged « 4; D. pupa, 
ventral aspect, (black phase) enlarged < 3; E. pupa, lateral aspect (olive green 
phase), enlarged X 3.] Reproduced from colored drawing by the author. 


EGG 


Hemispherical, the base flat, the top well rounded and the small 
micropyle deeply depressed. Color, glistening white. The surface is 
covered with vertical ridges, approximately 40 to 42 in number, each 
topped with a row of minute pearly nodules. Many of the ridges 
coalesce in the upper portion of the egg. There are apparently no 
horizontal striae between the ridges. 

The basal diameter of the egg is 0.60 mm. and the height 0.35 mm. 
The micropyle is relatively small and depressed. 

The egg is pictured as Figure 2 A. 


38 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 
First INstar LArvA 


Body cylindrical, constricted on segmental junctures, translucent. 
with a slight tinge of yellow. The length of larva was not noted. 

The head is black, larger than first segment, and measures 0.4: mm. 
in width. A prominent black scutellum occurs on the prothoracic 
segment. 

The body is thickly covered with relatively long black setae, each 
arising from a black papillus. These papillae occur in transverse rows 
across each segment, arranged in zig-zag lines. The two pairs of pro- 
legs and the anal pair are concolorous with the body. The true legs are 
dark. Ecdysis on April 19 to 23, 1962. Figure 2 B illustrates this 


instar. 
SECOND INSTAR 


Length of body, 3 mm. Head width approximately 0.5 mm. Ground 
color, translucent light green. The dark setae arisimg from black papil- 
lae are still present. The head is translucent light green; ocelli, black. 
and mandibles tipped with brown. 

From this point on, the rate of growth and time of ecdysis varied so 
widely that specific instars were not noted. One intermediate stage is 
recorded. 


Larva oF 11 Mm. LENGTH 


Body width, 2.1 mm. Head width, 2.3 mm, Ground color of head 
and body, liek green. A narrow darker green longitudinal middorsal 
stripe is present, bordered laterally by three narrow pale white stripes 
on the first three or four segments, becoming obsolete posteriorly. A 
similar stigmatal line runs the length of the body. Suprastigmatally on 
the 4th to 8th segments a black pa alee bearing a white seta is a notice- 
able feature. There are numerous white setae scattered over the body. 
each arising from a white papillus. The legs are black. and the prolegs 
c colores with the body. 


MATURE LARVA 


Length, ice extended, 22 mm. Body width through center, 4.5 mm. 
Head width, 2.3 mm. Our illustration of the larva (Fig. 2 C) appears 
foreshortened, due to the resting habit with the central segments 
arched upward. 

The ground color of the head and body is ight green. The head is 
glistening green except for a conspicuous black margin around the 


Life histories of phalaenid moths 39 


cheeks and across the mandibles. The ocelli are black on this black 
margin. The bases of the antennae are green and the terminal seg- 
ments black. The body is cylindrical and plump. A dark green longi- 
tudinal middorsal band extends the length of the body. In some exam- 
ples this is less clearly defined. Lateral to this dark green band is a 
wide whitish band, somewhat mottled in a few examples. This fuses 
laterally with the green ground color. A faint longitudinal stripe runs 
immediately superior to the spiracles. The latter are white centered, 
with narrow brown rims. Superior to this there is a line of black 
nodules, one to a segment placed above each spiracle. These are not 
always present, particularly in the caudal and thoracic areas. 

The legs are black, and the prolegs concolorous with the body. The 
crochets are dark brown. Numerous white setae, arisimg from minute 
white papillae occur sparingly over the body. There is also a heavy 
vestiture of minute white hairs, discernible only under high magni- 
fication. 

The larvae began spinning loosely woven white transparent cocoons 


on May 5, 1962. By May 16 all had pupated. 


PuPA 


Length, 16 mm. Greatest width through the patagia, 5 mm. When 
first formed the color is dark olive-green. This gradually darkens to a 
uniform shiny black. 

The eyes are prominent and bulging. The wing tips and antennae 
form part of an overhanging bulge extending caudalward over the 
fifth abdominal segment. In the green stage the spiracles are clearly 
distinguishable as brown ovals, with narrow dark brown rims. In the 
black phase they show as brown pits, surrounded by a brown shading. 

The cremaster is a stubby prominence, rounded at the tip. It bears 
two relatively long straight spines with recurved tips, and at the base 
of these there is a cluster of very small red-brown recurved hooklets. 
These features are shown in the two drawings of the pupa as Figure 2 D 
and E. 

The first imago appeared May 22, and a pair emerged May 23, 
1962. Progressive hatchings followed thereafter. 


BIBLIOGRAPHY 
CRUMB, S. E. 
1956. Larvae of the Phalaenidae. Tech. Bull. No. 1135, U.S.D.A., pp. 135, and 
257-258. 


40 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


DYAR, HARRISON G. 

1898. Descriptions of the larvae of fifty North American Noctuidae. Proc. Ent. 
Soc. Wash., 4: 319. 

FRENCH, G. H. 

1882. The preparatory stages of Plusia biloba. Papilio, 2(7): 113-115. 

GROTE, AUG. R. 

1873. Descriptions of Noctuidae principally from California. Bull. Buffalo Soc. 
Nat. Sci., 1: 140. 

HAMPSON, GEORGE F 

1913. Catalog. Lep. Phalaen. Brit. Mus. XIII, pp. 512-513. 

HOLLAND, W. J. 

1908. Moth Book. New York: Doubleday Page Co., p. 238. 

ROSEWALL, O. W. 

1922. Insects of the Yellow Thistle. Ent. News, 33(6): 179. 


THE EARLY STAGES OF 
PERO MACDUNNOUGHI CASSINO AND SWETT 
(LEPIDOPTERA, GEOMETRIDAE) 


JoHN ApAmMs Comstock 
Del Mar, California 


The species of Pero commonly occurring around Los Angeles and 
in the area southward to the Mexican border was long mislabeled Pero 
peplarioides in local collections. This was prior to the issuance of the 
McDunnough paper of 1949, and that of Rindge in 1955. These two 
papers clarified the mix-ups in our southern California Peros in 
proving that our so-called P peplarioides was actually Pero macdun- 
noughi C. & S., and that the real peplarioides, properly called P. occi- 
dentalis peplarioides (Hulst) was a southern Rocky Mountain species. 

A correction will therefore have to be made as to my paper of 1930, 
containing notes on the (presumed) life history of Pero peplarioides, 
as this should refer to P- macdunnoughi. 

This paper was somewhat sketchy, in addition to its incorrect de- 
signation of the species. Recent rearings of macdunnoughi in San 
Diego County (type locality, San Diego) would seem to justify a more 
thorough account of its metamorphosis. 

The moth has come to light continuously from April to late Sep- 
tember of this year (1962), and larvae have been beaten from Adenos- 
toma, Lonicera and Rhamnus crocea Nutt. It has been reared on 
Privet and Broom, and reported on “Pyrocanthus:’ Probably it is a 
general feeder on many vines and bushes. 

Females lay readily in captivity, depositing their eggs indiscrimi- 
nately. Frequently these are laid in rows on the edges of leaves or on 
the netting of the jar cover. They are usually laid on their sides. 


EGG 


Suboblongate, slightly flattened at one end and rounded at the other. 
Length, 1 mm. Width, 0.6 mm. The color is olive-green at the base, 
shading to straw at the rounded end. The surface is covered with 
minute hexagonal cells with very low side walls, discernible with high 
magnification but appearing to be granular under a low powered lens. 
The egg is pictured in lateral aspect on Fig. 1 A. 


41 


42 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


a Te 


D 


Figure 1. Early Stages of Pero macdunnoughi Cassino and Swett. | A. egg, lateral 
aspect, enlarged X 30; B. first instar larva, dorsal aspect, enlarged x 17; C. head 
of mature larva, enlarged < 9; D. larva, penultimate instar, lateral aspect, en- 
larged X 4; E. pupa, ventral aspect, enlarged x 414; EK and G. cauda and 
cremaster, ventral and lateral aspects, enlarged <X 20.] Reproduced from water 
color painting by the author. 


First Instrar LARVA 


Length, 4 mm. Head width, 0.45 mm. Width of body through 5th 
abdominal segment, 0.3 mm. 

The head is markedly bilobed. Its color is orange-yellow. The ocelh 
are black. See Fig. 1 B. 

The first two thoracic segments taper outwardly toward the head 
and the caudal segments widen to 0.4 mm. The body is translucent, 
and soiled yellow in color. There is a faint suggestion of a middorsal 
longitudinal paired gray stripe, shaded a lighter gray internally. The 
caudal area is orange-yellow. The setae are relatively long and dark. 


Early stages of Pero macdunnoughi 43 
PENULTIMATE INSTAR 


Length, 25 mm. Width through 5th abdominal segment, 1.8 mm. 

Head; width, 1.75 mm. Subquadrate, tapering toward mouth parts. 
The crown is accented by triangulate points supero-laterally. The 
outer margin is black, the face and front speckled brown, and the labia 
white. The mandibles are dark brown, streaked with black. See Fig. 
iC, 

The body is mottled dark and light gray, speckled with black. There 
are red-brown patches on the lateral surface near each segmental 
juncture. On the 5th segment there is a large black warty tubercle, 
placed substigmatally, (one on each side). This protrudes ventrally. 
There are numerous small black papillae scattered over the body, each 
giving rise to short black setae. The larva is a perfect camouflage, 
mimicking a gray twig. It is illustrated on Figure 1 D. 


FINAL INSTAR 


Length, 45 mm. Head width, 2.5 mm. The appearance is similar to 
penultimate instar except that the triangulate patches in front of each 
segmental juncture on the lateral surface are black, without a trace of 
the former red-brown coloring. From the upper angle of each triangle 
a black ridge extends upward and over the dorsum, and a similar ridge 
extends downward across the venter. There is also a ridge running 
longitudinally, in line with the spiracles. This 1s topped by a narrow 
black line. The spiracles are gray, with black margins, and narrow 
black central slits. The legs and prolegs are mottled black and gray. 
Most of the setae are black, and arise from black nodules. Those on the 
last caudal segment are gray. There is considerable variation in the 
stippling and spotting of individuals. 


Pupa 


Length, 17 mm. Width through center, 5.5 mm. Color, dark brown, 
shading to black on eyes, segmental junctures and cauda. Texture, 
smooth, except for the posterior margins of the movable segmental 
junctures, which are pitted. 

The maxillae and antennae reach to the wing margins. The spiracles 
are slightly darker than the ground color, and are relatively incon- 
spicuous. The anterior portion of the pupa is evenly rounded. 

The cremaster is the most distinctive feature. It is composed of a 
lobulated pyramidal element from which arise two stout shafts that 


AA, Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 1, 1963 


taper to recurved tips. These cross over each other. There are also 
two short recurved hooklets on each side. The shape of the last caudal 
elements is complicated, and difficult to describe. It is best defined by 
the accompanying illustrations on Figure 1 E, F and G. 

Three aspects of the pupa were shown in my paper of 1930, Plate 13. 


LITERATURE CITED 

COMSTOCK, JOHN A. 

1930. Studies in Pacific coast Lepidoptera. Bull. So. Calif. Acad. Sci., 29(1): 
29-31, pls. 13-15. 

McDUNNOUGH, JAMES H. 

1949. Critical notes on certain Pero species (Lepidoptera, Geometridae). Amer. 
Mus. Novitates, 1393: 1-11. 

RINDGE, FREDERICK H. 

1955. A revision of some species of Pero from the western United States (Lepi- 
doptera, Geometridae). Amer. Mus. Novitates, 1750: 1-33. 


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VoL. 62 APRIL-JUNE, 1963 Part 2 


CONTENTS 


eee 


Fe 


Functional Morphology of the External Appendages of Emerita 
Analoga. Cameron Knox and Richard A. Boolootian 


Fossil Arthropods of California. No. 25. Silicified Leafhoppers from 
California Mountains Nodules. W. Dwight Pierce 


A Contribution to the Biology of the Gray Garden Slug. R. O. Arias 
and H. H. Crowell 


Surf-Riding by the California Gray Whale. David K. Caldwell and 
Melba C. Caldwell 


A Se ee OE ae eee 


Molluscs from Pacific Northwest Archaeological Sites, 2. Washing- 
ton: 45-CA-30, A Coastal Shellmidden in the Ozette Area. 
Robert J. Drake 


Issued June 24, 1963 


Southern California 
Academy of Sciences 


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PUL IN OF THE SOUTHERN CALIFORNIA 
ACADEMY OF SCIENCES 
VoL. 62 APRIL-JUNE, 1963 Part 2 


FUNCTIONAL MORPHOLOGY OF THE EXTERNAL 
APPENDAGES OF EMERITA ANALOGA' 


CAMERON KNox AND RicHARD A. BOOLOOTIAN 
University of California? 


Los Angeles 


INTRODUCTION 


The sand crab, Emerita analoga (Stimpson), is the only species of the 
family Hippidae recognized from the western coast of the United 
States. Its distribution extends from Washington (Banner and Mc- 
Kernan, 1943) to Chile (Schmitt, 1935). The closely related species, 
Emerita emerita (Linn.), ranges from Baja California southward. In 
the family Albuneidae there are two species. The larger is Blephari- 
poda occidentalis Randall and the smaller Lepidopa myops Stimpson. 

Emerita analoga is by far the most numerous. It inhabits the sandy 
beaches of southern California, and may be found there in great num- 
bers. Anyone who has visited the seashore in this vicmity may have 
observed the many small “v” shaped channels which the water 
follows in the outwash of the waves. In the daytime under ordinary 
conditions this is the only visible indication of the presence of these 
animals other than an occasional discarded skeleton. The tiny ripples 
are created as the water rushes over the extended flagella of the 
antennae. As “soft-shelled” crabs they are dug up or dredged by fisher- 
men who recognize them as good bait for surf fish. Because individuals 
in the hard- and soft-shelled conditions are found living together, there 
has arisen the erroneous popular idea that they are different species. 

Common as these creatures are, not a great deal has been written 
concerning their anatomy or morphology. Most of the investigations 
conducted upon Emerita analoga have dealt with their habits, life 
history, and ecology. An article by MacGinitie (1938) gives an ex- 
cellent account of their movements and mating habits. Weymouth 
and Richardson (1912) too were concerned with the ecology and 
habits of the crabs. Mead’s work (1917) dealt chiefly with move- 


1Supported in part by a grant from the National Science Foundation (G 9561). 
*Department of Zoology. 


45 


| 
\ 


40 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


ments. Many of his experiments were conducted with animals not in 
their natural environment, and therefore the conclusions he reached 
are subject to critical examination. 


MeEtTHOops 


Data were collected on the sand crab, Emerita analoga, both by 
direct observation and photographic technique. The animals used im 
these studies were kept in small, individual glass tanks provided with 
six inches of sand covered with water. 

Normal behavior was studied, including swimming, digging, and 
feeding. Observations were then made where appendages were re- 
moved singly, in pairs, and in various combinations. 

The photographic equipment used was a sixteen millimeter Arriflex 
with 25, 50, and 150 millimeter lenses. The animals were photo- 
graphed ventrally, laterally, and dorsally at 24 frames per second. 
High speed photography, 50 to 250 frames per second, was used to 
better understand the use of external appendages. ‘Time lapse photog- 
raphy, 1 to 10 frames per second was used for studying the action 
of the antennae during feeding. 


OBSERVATIONS 


Between the high- and low-water marks Emerita analoga is most 
abundant. Some individuals have been found several feet beyond low 
water but they are not numerous and probably do not remain there 
for any great length of time. Occasionally these animals have been 
found above the reach of the waves, but in such cases they were deeply 
buried and not very active. They had undoubtedly advanced with a 
previous high tide and were then trapped by a rapid ebb. Upon ob- 
servation after the next high tide, it was found that practically all of 
the stranded individuals had made their way back to lower levels. 

The area that is covered by the wash of the waves is the favored 
locality for the sand crabs. Since this area is constantly being shifted 
by the variation of the tides, the animals are found to advance and 
retreat periodically. This movement has certain characteristics which 
are of interest and are quite readily observed. 

During the period of high tide, the population of the zone farthest 
inshore is dominated by males. Seaward, the males decrease in relative 
number so that at the outermost limit of the waves the sexes are prac- 
tically equal or slightly in favor of the females. Still farther seaward 
beyond the wash, there are more females than males. At low tide these 
conditions are reversed; the females are most abundant at the shore- 


Functional morphology of Emerita analoga A7 


ward limits of the waves and the males at the seaward limits. A single 
male, however, does not migrate the entire distance; the group move- 
ment gives rise to the observed conditions. Experiments using marked 
animals have indicated that many of the males which occupy the zone 
at the lower limits of the wash of the waves during low tide move in- 
shore at high tide so that they are still at the lower limits of the waves. 
Likewise the males at the shoreward limits durmg low tide make up 
most of the population at the highest reach of the water at high tide. 

In the course of the studies upon population movements, several 
different habitats have been observed. At Venice, California, a large 
area of gently sloping beach is shielded from the sun by piers, the 
pilmgs of which aid materially in reducing the force and errosive 
effect of the waves. In this relatively quiet shaded environment the 
sand crabs were found to be more numerous than on any of the un- 
covered beaches. A different type of habitat is found on the open, 
exposed beaches which make up most of the shoreline. This type may 
further be subdivided according to whether the seaward slope is steep 
or gentle. When steep, the errosive effect of the waves is relatively 
more pronounced. There is much shifting in the surface sands, thus 
creating an extremely unstable substratum. As the slope of the shore 
increases, the wave-washed zone becomes more narrow, thereby re- 
ducing the area which the sand crabs inhabit. On the gently sloping 
shores, the wave-washed zone is much broader and the errosive effect 
is relatively weak. Since Emerita analoga live under all of these vari- 
able conditions, many of their habits must be interpreted with con- 
sideration for the special environment they inhabit. Table 1 illustrates 
the distribution of sexes with varying slope and tide conditions. 


TABLE 1 
Distribution of the sexes of Emerita analoga at high and low water. 
Zones at high tide Zones at low tide 
In- Off- In- Off- 
Station shore Middle shore shore Middle — shore 
1 Males 39 42 18 27 22 Dy 
Females 11 37 25 40 29 LY 
9 Males 29 31 ila 20 18 35 
“~~ Females 10 21 33 32 27 24. 
3 Males 13 15 11 i 10 15 
Females 3 17 17 19 19 12 
Males 24: il 15 4 13 19 
seaetemales! = 07 20 26 12 17 9 
Males 15 29 23 10 17 22 
5 Females 2 31 28 11 28 13 


48 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


At Stations 1 and 2 the beach was very gently sloped and entirely 
shaded by a wide pier. Because of the resistance of the pilings, the 
force of the waves was greatly diminished. At Station 3 the beach was 
steep, uncovered, and exposed. Stations 4 and 5 were similar to Station 
3 except that the slope was gentle. Each entry in the above table 
represents the average of eight counts made as nearly simultaneous as 
possible. A screen twenty-four by twenty inches was used to collect 
the sand crabs. 

The normal habitat of Emerita analoga presents factors which are 
both advantageous and disadvantageous to the animals. In their parti- 
cular zone they are almost without competition except for the oc- 
casional appearance of one of the Albuneidae mentioned before. Their 
food consists of microplankton which is brought to them by the waves. 
The constant wash of the water makes it unnecessary for them to 
perform their own pumping as many other plankton feeders are 
obliged to do, Their task is simply to sift the organisms from the water. 

Among the disadvantages of the environment are the difficulties 
which the animals meet in maintaining their position on an unstable 
substratum and escaping the shock of the waves. They protect them- 
selves by burrowing into the sand, and by this means also hide from 
predators such as birds and fish. At the same time, the sand crab must 
be able to get sufficient food and oxygen from the water. 

As might be expected, this unusual mode of life has resulted m 
striking modifications of the typical decapod anatomy. 


ANATOMY 


Carapace. The somites of the head and thorax of Emerita analoga 
are fused with one another in the usual way. The whole cephalothorax 
is covered by a heavily calcified carapace. It is found that the mem- 
branous dorsal body wall is not united with the carapace but only 
closely applied to it. This is especially evident in the posterior portion 
where by careful dissection one part may be separated from the other 
without much difficulty. 

In the course of collecting the sand crabs, a series was taken from La 
Jolla to Monterey, California, with most of the work bemg done on 
specimens gathered near Santa Monica. 

The carapace (Fig. 2) is ovate and measures 15-30 mm. in length 
in the adult female and between 10-22 mm. in the adult male. The 
width in the female ranges from 9-23 mm. and in the male from 


Functional morphology of Emerita analoga 49 


@) 


=a 6). 
ow ee ena — - @® 
eo & sb = 
c WW) 0 ore 
ro) 

oOo ®O -: {uo} 
> oOo c 
oOo — ne} 
> 
ag re 3 SS 
@ 
=} fo) . 
~ oO — Ee 
c iS fe) 
wo : (6) oO 
3 i oO c : 
a) ener) a wo re) 
c a wo i ‘e 
Chan) = ro) a >) 


{o) oO 
Q ® 
wo = 

QO 


Fig. 1, ventral view of Emerita analoga, with the appendages removed and the 
abdomen extended. x 2; fig. 2, dorsal view of Emerita analoga. x 2; fig. 3, trans- 
verse section between the second and third walking legs. x 2. 


50 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


8-17 mm. In a series of individuals of either sex it is found that the 
relation of the width to the length of the carapace is quite variable. 
‘Two specimens measuring the same in length may vary as much as 
4 mm. in width. Measurements of the carapace of an average female 
would be about 16 mm. in width and 22 mm. in length; the male would 
be about 10 mm. in width and 14 mm. in length. 

The carapace is strongly convex, with its lower margin overlapping 
the sterna and covering the basal joints of the walking legs (Fig. 3). 
The branchiostegites (Figs. 1 and 3) of Emerita analoga have the 
outer layer of the pleural fold strongly calcified. The part of the fold 
which lies next to the gills is not calcified but appears as a thin trans- 
parent membrane. The rostrum is much reduced and is present only 
as a short, blunt, median projection. On either side of it is a lateral 
process from the carapace. These are only slightly longer than the 
rostrum, but more rounded. They overlie the uncalcified part of the 
eye stalks and probably serve to protect them from injury. On the 
anterior half of the carapace are two conspicuous transverse grooves, 
the anterior one being the cervical groove and the posterior one, the 
branchial groove. There are numerous smaller grooves which become 
less distinct laterally and posteriorly. That part of the carapace ad- 
joming the first abdominal tergum is quite smooth. 

The posterior part of the carapace is noticeably wider than the 
anterior, giving the animal a streamlined appearance. This shape may 
be of use in cuttmg down the drag of the water if the animal happens 
to be digging in during the outwash of the wave. Once buried, this 
same shape may serve as a wedge to give the crab a firmer position. 

Terga of Abdomen. The abdomen (Fig. 2) is distmctly segmented 
and flexible. It is tapered posteriorly, ending with the telson which is 
a rather narrow triangular structure. The telson and the sixth segment 
are bent under the body when at rest, and in the case of an oviferous 
female serve to cover and protect the eggs. The tergum of the first 
segment is a small crescent shaped piece which occupies a rounded 
notch in the posterior edge of the carapace to which it is partially 
fused. In addition there is a tough tissue which attaches the first tergum 
firmly to the ventral side of the carapace. The terga of the remaming 
segments are convex, covering their respective sternum and the basal 
joints of the abdominal appendages. The second abdominal somite is 
four to five times broader than long and is characterized by large wing- 
like pleura. The anterior margin of the tergum is almost straight but 
the posterior edge has a curved sinus into which the tergum of the 
third somite fits. The third, fourth, and fifth terga are similar in form 


Functional morphology of Emerita analoga Bi 


corn. 


Fig. 4, eye stalks. Dorsal view. x 3; fig. 5, antennule, left. Lateral view. x 3; fig. 6, 
gill chamber of Emerita analoga. Lateral view, with carapace and branchios- 
tegite removed. x 3; fig. 7, same as above with the gills removed to show the 
calcification of the lateral. body wall. x 3; fig. 8, several annuli from the flagellum 
of the antenna. The hairs have been removed to show their points of attachment. 
x 12; fig. 9, diagrammatic transverse section through the flagellum of the an- 
tenna. x 3; fig. 10, antenna, left. Lateral view. x 3; fig. 11, hairs from the antenna. 
Lateral view. x 12 [A. outer row; B. second row; C. third row; D. fourth row; 
EK. median row; F. portion from the first or outer row hair, looking into the trough 
between the barbs]. 


52 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


but much smaller in size than the second. The tergum of the sixth 
somite is as long as broad and has a wide rounded lobe which extends 
posteriorly and articulates with the telson. 

Sterna. The sterna in the region of the thorax are reduced to bars 
between which the walking legs are attached. The succeeding sterna 
are joimed to each other by a median bar. From each sternum there 
extend inward two apodemes. These afford a certain degree of rigidity 
to the body as well as providing attachment of the extrinsic muscles of 
the legs. Those sterna to which the mouth parts are fastened form a 
more continuous plate (Fig. 1) although some parts of it are very 
slightly calcified and may on first observation give the impression of 
transparent membranes. 

The sterna of the abdominal region are calcified around the points 
of junction with the pleopods, but otherwise, except for thin transverse 
bars, appear only as transparent membranes through which the in- 
testine may be seen. The amount of calcification and the width of these 
bars vary in different individuals. The length of time since changing 
the shell is a significant factor. The ventral surface of the telson is 
membranous except for a narrow calcified border. The structure of 
the telson will be taken up in more detail in a later section. 

The sterna of the abdomen are rarely exposed to any extent and do 
not afford a vulnerable spot as might be the case in animals which 
characteristically swim by flexing the abdomen. In the normal position, 
the tip of the telson reaches the base of the first pair of walking legs. 
It shields the under side of the body against abrasion while the animal 
is burrowing. 

Eyes. The eye stalks of Emerita analoga have four segments, the 
terminal (fourth) one being the longest and most slender and bearing 
at its enlarged tip the cornea (Fig. 4). Near the basal end of the third 
segment there is a small round opening covered by a delicate trans- 
parent membrane and bordered proximally by a tuft of long hairs. 
The function of this opening was not determined. Part of the second 
segment and its articulation with the third segment are very thinly 
calcified. It is this region which is covered above by the lateral-anterior 
projections of the carapace. The muscles and articulations of the eye 
stalks are so arranged that movement is almost entirely in the sagital 
plane. Lateral movement of the stalks is limited to about two milli- 
meters. 

The elevation and lowering of the eyes have been noted while 
watching these creatures in their normal environment. When the 
crab is at rest and not actively feeding, the eye stalks protrude above 


Functional morphology of Emerita analoga 53 


the sand about two-thirds the length of the terminal segment. The 
cornea is so situated that there is vision in all directions, thus ap- 
parently making it possible to observe the approach of enemies with 
the minimum exposure of the body. However, visual stimulation, 
such as the approach of foreign objects does not usually cause any eye 
movement or other reaction unless there is actual contact with some 
part of the animal itself. The slightest touch upon the stalks or the 
antennules causes the eyes to be flattened against the sand, and an 
impact in the proximity will produce the same reaction. This leads 
one to suspect that visual-muscular coordination is not as well de- 
veloped as might be expected. 

Antennules. The antennules consist of a protopodite of three seg- 
ments to which are attached two flagella (Fig. 5). The lateral one is 
the longer and is made up of thirty-five to forty-five short annuli. 
Two distinct rows of hairs approximately at right angles to each other 
extend from the dorsal and ventral sides of the flagellum. The ventral 
row is continuous for the whole length but the dorsal row may stop 
five or six annuli from the end. The median flagellum is shorter and 
consists of sixteen to twenty-one annuli. It also bears two rows of hairs 
approximately at right angles with each other. The hairs are rather 
spinulose in nature but do possess a few short barbs. The exact charac- 
ter of the hairs of the antennules has been observed to be somewhat 
variable between different individuals. 

The opening for the otocyst occurs on the dorsal side of the coxo- 
podite. It is encircled by a row of short spmulose hairs. On the second 
segment of the protopodite there is a short lateral process, the ap- 
pendicular scale, which is characteristically covered with long plumu- 
lose hairs. The distal segment of the protopodite is slightly compressed 
laterally. There are dense rows of spinulose hairs on the dorsal and 
ventral surfaces. 

The functioning of the antennules is an important factor in the 
ability of the sand crab to maintain a respiration water supply while 
buried. This will be described later in the discussion of respiration. The 
hairs from each flagellum of the two antennules interlock forming a 
rectangular channel down which water flows. Sand or other large 
particles are strained out so that a minimum of foreign material reaches 
the gill chamber. 

Antennae. The protopodite of the antenna has the usual two seg- 
ments, the second of which is enlarged and provided with three 
prominent spines on its distal border (Fig. 10). The coxopodite bears 
the opening for the green gland. The endopodite is composed of a 


eee aaa... 


5A Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


peduncle of three large segments and a long flagellum of numerous 
small annuli (165-185 in an adult female). The division of the pe- 
duncle into three segments allows a wide range of movement. Im- 
mediately below and lateral to the basal segment of the endopodite 
there is another small segment which articulates distally with the 
second segment of the endopodite and proximally with the basopodite. 

From each of the annuli of the flagellum arise nine hairs arranged 
in longitudinal rows so that they form a “‘v” shaped trough. The two 
outer rows are made up of hairs which are long, slender, and curved 
sharply inward at their distal ends. The inner rows become succes- 
sively shorter, with the median unpaired row being the shortest of all 
(Fig. 11a-e). The long hairs of the outer rows have two lines of 
numerous fine barbs perpendicular to the main stem and at an acute 
angle with each other so that they form a small “‘v” shaped trough 
(Fig. 11f). The hairs forming the second row are stouter and have 
short, stubby barbs arranged quite irregularly. Near the base of the 
hair, the barbs are longer and much more slender. The rest of the 
inner rows are made up of hairs which are similar in structure but 
smaller in size. 

The hairs of the median row are spine-like and have no barbs. The 
hairs of the flagella are specialized for filtering out the food. As the 
water flows into the trough between the hairs and the barbs, the stout 
inner rows remove the larger particles in suspension and allow the 
rest to pass on to the slender outer rows. These, with their many fine 
branches, complete the filtermg. By taking up so much of the force 
of the water and removing large particles, the strong mner rows of 
hairs protect the outer ones from being broken or pushed apart. 

This arrangement of the parts is of particular significance in rela- 
tion to the function of the flagella. As mentioned before, Emerita ana- 
loga uses plankton for its source of food, and the antennae are the 
means for securing these small organisms. As a wave washes in the 
antennae are extended. The movement of the water through the hairs 
of the flagella as the wave flows out causes the small particles to be 
collected. From time to time the antennae are whipped downward and 
wiped off by the mouth parts. This action has been observed to be 
repeated as many as eight times during a single wave. The two flagella 
are not brought down simultaneously, but separately and with no 
definite alternation. The act of lowering the flagella includes a rotation 
of 180° in that part in such a way that there is constantly a pressure 
into the trough of hairs, thus keeping the collected food from being 
washed away. 


Functional morphology of Emerita analoga 


Or 
Or 


Fig, 12, first maxilla, left. x 3; fig. 13, mouth and mandibles. Ventral view. x 8; 
fig. 14, second maxilla, left. x 3; fig. 15, different types of hairs found on Emerita 
analoga. x 12 [ A. from the scaphognathite; B. from the endopodite of the uropod; 
C. from the margin of the telson; D. from the dactylopodite of the first walking 
leg; E. from the antennule; F. from the coxopodite of the first maxilla; G. from 
the meropodite of the first walking leg]; fig. 16, first maxilliped, left. x 3; fig. 17, 
second maxilliped, left. x 3; fig. 18, third maxilliped, left. x 3. 


56 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


When the animal is not feeding, the antennae are folded together 
and coiled up under the protecting cover of the third maxillipeds. In 
such a position only the two segments of the protopodite and the basal 
segment of the peduncle are visible. The hairs of the flagella are turned 
toward the center of the coil. Practically the whole flexion necessary 
for this movement takes place in the articulation between the distal 
segment of the protopodite and the peduncle and in the segments distal 
to this. The three spines of the basopodite point anteriorly. This pro- 
vides a barbed armor for the rest of the antenna as well as for the 
eye stalks which lie adjacent. 

Mouth. The oral opening (Fig. 13) is a longitudinal slit about two 
millimeters in length by one-half a millimeter in width, leading into 
a slightly larger esophagus. In hfe, the surrounding tissue is colored 
deep red. The mouth is bordered laterally by raised membranous 
structures upon the inner surface of which there are usually found a 
series of wrinkles covered by small hairs. The labium is continuous 
with the lateral folds at the posterior end of the mouth. The labrum is 
a large, conspicuous, triangular flap. It is situated a short distance 
anterior to the mouth. It is quite thick and tough but is not at all cal- 
cified. The anterior border of the mouth is formed by a thin mem- 
branous fold which extends backward from the base of the labrum. 
Between the labrum and the sternum, and attached to the latter, are 
two palps. They are derived from an anterior fold of the epistome. Each 
bears a few long plumulose hairs at its apex. On either side of the 
mouth is a rounded elevation from which arise a large number of 
plumulose hairs. The homology of these structures was not deter- 
mined. 

Mandibles. Each mandible consists of two small uncalcified seg- 
ments and a large palpus (Fig. 13). The proximal segment is fre- 
quently fused along its median edge with the raised margin of the 
mouth. The lateral portion of the basal segment gives rise to the large 
flattened palpus which extends forward enfolding the posterior-lateral 
edges of the labrum. The palpus is bordered by a fringe of plumulose 
hairs. The distal segment of the mandible is broad and flat. It is pro- 
duced medianly into a blunt rounded point upon which are a few 
long hairs. The mandible may be able to create a slight current but 
it would be insignificant compared to the action of the other mouth 
parts. The absence of masticatory apparatus is what would be expected 
in correlation with the sand crab’s exclusive diet of plankton. 

First maxillae. The first maxillae (Fig. 12) are composed of a 
protopodite and a small, palpus-like endopodite. The coxopodite lies 


Functional morphology of Emerita analoga 57 


median to the basopodite and immediately ventral to the mouth. It is 
a thin uncalcified lamella bordered by rather stiff hairs. It is con- 
nected with the basopodite by a narrow, crescent shaped extension of 
the base. The median proximal portion of the coxopodite is produced 
into a pointed process from which arise a dense fringe of long plumu- 
lose hairs. The basopodite is composed of two parts. The distal segment 
is approximately the same width as the coxopodite but it is longer and 
bears a tuft of long spmulose hairs at its extremity. The proximal 
segment is narrow and curved. It terminates in an expanded, knob-like 
base. Very long plumulose hairs extend from the convex surface of the 
basal segment. The endopodite is a small pointed lobe arising laterally 
from the basopodite. 

Second maxillae. The second maxillae (Fig. 14) exhibit practically 
the same structure as the first. The coxopodite is divided into two lobes, 
the median of which is similar in form to the coxopodite of the first 
maxilla. The lateral lobe is a narrow, finger-like projection with a few 
hairs at its tip. The basopodite and endopodite are both quite sumilar 
to those of the first maxilla but somewhat larger. The hairs of the 
endopodite are restricted to a few scattered ones on the median edge, 
while on the basopodite, they form a complete fringe around the 
margin. The exopodite, as in other decapods, forms the scaphognathite. 
In Emerita analoga this structure is rather short and broad. It barely 
reaches into the gill chamber (Fig. 6). Around the lateral, convex 
edge there is a row of plumulose hairs (Fig. 15a) which increase in 
length toward both extremities. The function of the scaphognathite in 
regard to respiration will be discussed later. 

First maxillipeds. The first maxillipeds (Fig. 16) are thinly calcified 
and are quite flexible. The protopodite is reduced to a single small 
segment which articulates with the simgle segment of the endopodite 
and with the exopodite. The epipodite typical of other decapods is 
entirely missing in the sand crab. 

The exopodite has two segments. The proximal one is flat and 
elongated, with a truncate distal end and a widened base. A series of 
short plumulose hairs border the lateral margin. A long lobe arises 
on the median edge. The terminal segment of the exopodite is broad, 
flat, and bluntly pointed. It is so articulated that it is able to create 
a current by a flapping action. The long plumulose hairs with which 
it is armed are curved and give the greatest force in the anterior to 
posterior sweep. This is a striking modification from the typical struc- 
ture of the exopodite. Here the flagellum is replaced by a paddle which 


is part of the respiratory mechanism. 


58 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


The long endopodite is ventrally convex so that a movement of this 
part performs a scooping effect. The lateral and terminal margins are 
edged by long plumulose hairs which are so arranged and curved that 
in action they pass the food on toward the mouth. Near the base of the 
endopodite there is a tuft composed of a few long plumulose hairs 
which reach almost to the end of the lobe. The entire convex ventral 
surface is covered by many short hairs. 

Second mazillipeds. The second maxillipeds (Fig. 17) of the sand 
crab also have the protopodite reduced to a single, very small segment 
which articulates with the exopodite and with the endopodite. 

The endopodite has four segments, the first of which is by far the 
longest. All of the parts of the endopodite are better calcified than 
those of the exopodite, but they do not approach the rigidity found 
in the third maxilliped. The second segment is short and articulates 
with the third by a joint which is not terminal but located on the flat 
surface. The possible movements which this allows are similar to those 
of a ball and socket jomt. The third segment is small and cylindrical. 
The terminal segment is also cylindrical but quite elongated. The two 
distal parts are covered by long plumulose hairs scattered over their 
whole surface. The two proximal segments have long plumulose hairs 
forming a dense fringe along their borders. 

The exopodite consists of two segments, both of them being thin, 
flat, and very lightly calcified. The proximal one is about three times 
the length of the distal, and has a broad base articulating with the 
protopodite at its median edge. From the wide base the proximal seg- 
ment of the exopodite tapers to its articulation with the distal segment. 
The latter is flat and oval. It has a row of plumulose hairs along its edge 
which are much longer than those on the basal segment. The exopodite 
of the typical decapod possesses a long flagellum which in the sand 
crab is replaced by a broad paddle. This is correlated with the adapta- 
tion of this part for its function in the respiratory mechanism. The 
distal, oval segment lies in the channel carrying water to the gills. 
It is able to move in an arc at right angles to its flat surface, thereby 
creating a current which passes through the gill chamber. 

The second maxillipeds function also as a part of the feeding ap- 
paratus. The endopodite assists in moving the food toward the mouth. 
The long hairs serve to wipe the collected food from the flagella of the 
antennae as they are drawn over the mouth. The third maxillipeds 
also participate in this function. When an antennal flagellum is 
whipped down, it is enclosed between the second maxillipeds. The 
long tufts of hairs on the second segments of the endopodites of each 


Functional morphology of Emerita analoga 59 


appendage interlock medially and against this the flagellum moves. 
The two distal segments lie over the flagellum. After the movement 
of the antenna is completed, each endopodite moves laterally so that 
the hairs no longer interlace. They then take up the function of 
passing the food along toward the mouth. 

Third mazillipeds. The third maxillipeds (Fig. 18) of Emerita 
analoga lack an exopodite. The endopodite has only four segments in 
place of the usual five. The two segments of the protopodite are small, 
the basopodite being transversely linear to the ventral surface while 
the coxopodite is subrectangular. Along the median borders of each 
grow dense tufts of long plumulose hairs. The basopodite is joined 
immoveably with the ischiopodite. The latter is a broad, flat structure 
having a prominent longitudinal ridge on the dorsal surface. This is 
the main axis of the segment at either end of which the meropodite 
and basopodite articulate. The ventral surface of the ischiopodite is 
covered by scattered groups of short blunt hairs arismg from shallow 
grooves. There are long plumulose hairs on the median and lateral 
borders, those on the former being two to three times longer than those 
on the latter. The meropodite is much smaller in size than the ischio- 
podite. Its anterior-lateral margin is rounded and possesses a large 
tuft of long and short plumulose hairs. The articulation between the 
ischiopodite and meropodite is such that a flexion of this joint causes the 
distal segments to be hidden behind the broad shield of the ischio- 
podite. When the animal is at rest this is the typical position. The two 
terminal segments are narrow and elongated. They bear a number 
of rows of long and short plumulose hairs along their median edges. 

Since the third maxillipeds.are the most exterior of the mouth parts, 
they form a protecting shield. They are much heavier and more 
strongly calcified than any of the other parts. When food is brought 
down by the antennae, the broad ischiopodites of the third maxillipeds 
form a chamber relatively free from currents in which the rest of the 
mouth parts can transport the food to the oral opening. 

Walking legs. The first walking legs (Fig. 19) do not have chelae. 
The seven segments of the leg are reduced in Emerita analoga to six 
by the fusion of the basopodite with the ischiopodite, as is true for most 
Reptantia. The carpopodite and the propodite have spines which assist 
the crab in getting a strong foothold. The surface of all the segments 
are more or less marked by shallow grooves from which short blunt 
hairs arise. The broad flattened dactylopodite is modified to give a 
large area for contact with the substratum. It is bordered by long and 
short stiff plumulose hairs (Fig. 15d). The function of the first legs 


60 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


dactylo. 


pro. 
carpo. 
mero. 
ischio. 


COXxO. 


carpo. < 


conto) 


COxOoO. 


Fig. 19, first walking leg, left. Lateral view. x 2; fig. 20, second walking leg, left. 
Lateral view. x 2; fig. 21, third walking leg, left. Lateral view. x 2; fig. 22, fourth 
walking leg, left. Lateral view. x 2; fig. 23, fifth walking leg. Lateral view. x 2: 
fig. 24, protopodite of the third walking leg of a female, showing the genital 
opening. x 4; fig. 25, coxopodite of the fifth walking leg of a male, showing the 
enital opening. x 4; fig. 26, pleopod from the second abdominal somite. x 3; fig. 


oO 
as 
7 


. pleopod from the third abdominal somite. x 3; fig. 28, pleopod from the fourth 


/ 
abdominal somite, x 3. 


Functional morphology of Emerita analoga 61 


in walking, as well as in digging, is to push. As the animal burrows 
into the sand or moves along the beach, the spines dig in and hold. 
The flat terminal segment may be turned laterally so that it presents 
the greatest surface against the sand when pushing. 

The second walking leg (Fig. 20) also has the basopodite fused with 
the ischiopodite. The propodite retains a sharp projection but the carpo- 
podite does not. The dactylopodite is broad, flat, and modified as a 
hook. The function of the second legs is to assist with the actual digging 
when the crab starts to burrow. The terminal segment is really a scoop 
with which sand is removed from under the animal and thrown out 
anteriorly. 

The third walking leg (Fig. 21) is a replica of the second in all the 
major details. There are a few less hairs on the segments, but this 
character varies in different individuals. The dactylopodite is slightly 
narrower than that of the preceding appendage. The use of this leg is 
the same as that of the second. In the females, the genital opening 
is found on the coxopodite (Fig. 24). 

The fourth legs (Fig. 22) have the same segmentation as the third. 
There are no spines on any of the parts. The dactylopodite is not a hook. 
It is flat and sub-triangular, terminating in a blunt point. The hairs 
on the fourth legs are relatively longer and more numerous than on 
any of the other legs. Although the function of the fourth walking legs 
is digging, their operation is quite characteristic. The leg is turned 
in such a way that the two terminal segments act as scoops and push 
the sand out sideways. This is in contrast to the second and third legs 
im which the dactylopodite is dug m and used as a hook in a posterior- 
anterior direction. Judging by the relative amount of sand which each 
appendage throws, the hooks are apparently more efficient. 

The fifth walking legs are long and filiform (Fig. 23). They possess 
the seven segments typical of the decapod leg but the basopodite and 
ischiopodite show indications of fusion. The protopodite and the dac- 
tylopodite form a sharply toothed chela which is surrounded by a dense 
tuft of barbed hairs. In its usual position this leg is not visible extern- 
ally. It is folded back under the branchiostegite and lies against the 
gills (Fig. 6). When fully extended, it reaches as far anteriorly as the 
posterior end of the scaphognathite. One of the functions of the fifth 
leg is to keep the gills free of foreign materials, especially sand. There 
is not any other structure or appendage which is adapted for this work. 

In the males the opening of the genital pore is on the coxopodite 
(Fig. 25). It can readily be located by the genital papilla. Aside from 
the function of removing debris from the gills, it has been observed 


62 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


that the male crabs make use of the fifth legs m placmg the sperm 
ribbons. It has also been observed that the fifth legs are used by the 
females in arranging the newly laid eggs on the pleopods. 

Pleopods. The females have three pairs of uniramous pleopods 
(Figs. 26-28) on the second, third, and fourth abdominal somites. The 
first and fifth do not have any appendages. The first and second pairs 
of pleopods are more than twice as long as the third pair. They are 
cylindrical and filiform, and are composed of three nearly equal seg- 
ments. This is in contrast to the biramous, multiarticular pleopods of 
the typical decapod. On the sand crab, these appendages are sparsely 
covered with hairs, the proximal ends of which are expanded to fit 
into cup shaped sockets. During the breeding season the egg mass is 
attached to the pleopods by a tough transparent substance. The eggs 
are arranged in branched strings which are held together by the same 
cuticle-like covermg. The males of Emerita analoga have no pleopods. 

Uropods. The uropods (Figs. 1 and 2) on the sixth abdominal somite 
have a long basal segment and two large terminal lamellae. The proto- 
podite is shghtly broader at its distal extremity than at the base. Along 
its lateral margin there are a series of long and short plumulose hairs. 
The exopodite consists of two segments, the proximal one being very 
small. The terminal segment is approximately twice as long as broad. 
It is flattened and oval in outline. Along the anterior margin there are 
several rows of coarse spmulose hairs while at the extremity there are 
longer plumulose hairs. On some individuals jointed hairs (Fig. 15b) 
are not uncommon. The endopodite is about three times as long as 
broad and is armed with hairs in a similar manner to the exopodite. 

Although the sand crabs in their natural environment do not swim 
to any extent, the uropods perform the function of propelling the 
animal when that is necessary. The broad, oval shape of the terminal 
segments is adapted for swimming as well as for burrowing in the 
sand. 

Telson. The telson is triangular in outline (Fig. 1) and in the adult 
female about twice as long as broad. It has a median shallow sinus 
on its proximal edge by which it is articulated with a rounded lobe 
from the tergum of the sixth abdominal segment. A short groove ex- 
tends posteriorly from either side of the sinus. Lateral to the sinus the 
base of the telson is attached to the protopodites of the uropods. The 
dorsal surface of the telson is convex and is bordered by a slightly 
raised, narrow rim which bears a series of short plumulose hairs. 
Around the margin and extending to the tip are several rows of long 
barbed hairs (Fig. 15c). The ventral surface of the telson is practically 


Functional morphology of Emerita analoga 63 


flat and almost entirely uncalcified except for a narrow band around 
the edge. The anus opens in a small depression about one fourth of 
the way back from the tip. 

The telson exhibits a small degree of sexual differentiation. In 
males it is proportionately smaller and narrower than in females. Also 
the frmge of hairs which is so conspicuous on the female telson is 
usually quite sparse on the male. 


HasitTs 


Burrowing. Under usual circumstances the act of digging into the 
sand is a relatively short one. However, the coarseness and firmness of 
the sand, the amount of water present, and the size of the animal all 
affect the time. Unless the animals are purposely dug up and placed 
on the surface, burrowing is rarely observed in the daytime. At night 
when the crabs are much more actively moving about, the burrowing 
process can be frequently seen. 

In all cases observed under normal environmental conditions, 
Emerita analoga has been found to enter the sand backwards with its 
anterior end pointing seaward. Movement on the surface and while 
swimming is also rear end foremost. The direction of burrowing is 
obliquely downward. This method has quite obvious advantages. Be- 
cause the food of these animals is found in the greatest amount in the 
water, it is necessary that the mouth and food capturing devices be 
located near the surface. In order to see, the eyes must protrude above 
the sand. The antennules must have access to a supply of fresh water 
for the gills. 

In the normal habitat, the process of digging is always the same and 
is usually accomplished while the wave is covering the beach. Often 
an animal which is dug up and placed on the surface will wait for the 
next wave to come in before it disappears under the sand. 

The uropods beat rapidly in an anterior-posterior direction. The 
hairs of the terminal segments are so arranged that they exert the 
greatest force in the posterior to anterior sweep. The action of these 
parts serves to stir up the sand on the bottom and mix it with the 
water; rather than actually scooping out a hole, they merely make 
a more fluid region into which the animal pushes. The second, third, 
and fourth pairs of walking legs are adapted for actually digging. It is 
with the aid of these parts that the animal is able to bury itself in hard, 
compact sand. The terminal segment of the second and third pairs 
are modified as hooks, and the fourth pair has flattened terminal seg- 


64. Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


ments. These legs are so jointed that they are able to perform a power- 
ful scooping action. In the still water of an aquarium, the little puffs 
of sand are seen to be thrown out as the crab disappears. The first pair 
of walking legs is used for pushing. The end segment is flattened and 
there are spines on the fourth and fifth joints, all of which give the leg 
a strong purchase. Digging is continued until only the eyes and the 
tips of the antennules are visible. When the crab is completely covered, 
the first legs and the third maxillipeds push against the side of the 
burrow, thus making an open space under the anterior end of the 
animal. This cavity is filled by the water used in respiration. 

It has been determined that the rate of burrowing increases as tem- 
perature decreases. The optimal temperature range for both sexes is 
20 to 25 C, but females are better able to withstand temperature 
extremes (Jeanette Hanby, personal communication). 

Reproduction. Emerita shows a distinct annual breeding cycle, as 
determined by plotting the percent of the monthly sample of females 
bearimg eggs against months of the year. The ovigerous period extends 
from April to October, reaching a maximum in July when 80% of the 
population was ripe. 

Incubation period for animals in the laboratory is between 29 and 32 
days. It is not certain if Emerita berries more than once during the 
breeding season (Boolootian, et al., 1959). 

Gills and respiration. The gill chambers of Emerita analoga are 
covered by the branchiostegites, which may be removed to expose the 
gills (Fig. 6). There are eight pleurobranches and, at the level of the 
first walking legs, ventral half only of a ninth. All of the gills are 
phyllobranchiae. They are all attached at about the same level on the 
body wall, well above the articulation of the legs. They probably did 
not all arise as pleurobranchs but without knowledge of the develop- 
ment it is difficult to class them otherwise. 

The method of obtaining the supply of water for respiration shows 
marked differences from that used by typical decapods. Since the 
animal is usually buried under a layer of sand, it is necessary to trans- 
port the water from the surface down to the gill chamber. One of the 
most obvious specializations found in Emerita analoga is that the 
direction of flow of the respiratory current is from anterior to posterior 
instead of in the posterior-anterior direction typical of other decapods. 
Calman (1909) states in regard to the respiratory current, “‘as a rule 
it sets from behind forward, though it appears that i some Cases, 
especially in decapods which burrow im sand or mud, the direction of 
the current is periodically reversed. . . . The flagella of the antennules 


Functional morphology of Emerita analoga 65 


in some Hippidea and of the antennae in the Corystidae form a long 
exhalent (or inhalent) siphon?) Weymouth and Richardson (1912) 
have noted the reversal of the respiratory current in Emerita analoga, 
while similar observations were made by Garstang (1896, 1897) on 
Corystes and Porturnnus. However, examination of the arrangement 
of the hairs and the articulation of the jomts mvolved in creating the 
current would seem to indicate that for Emerita analoga the anterior- 
posterior flow is the usual one. Many observations on buried animals 
have shown but a few exceptions. Only when the individuals were 
forced to remain above the surface did they exhibit a periodic reversal 
of the direction of the respiratory current. It was found that sand 
crabs in a laboratory aquarium show this change of direction much 
more frequently than do animals in their natural environment. 

When the sand crab is at rest the tips of the antennules protrude 
above the surface of the sand. Upon close observation the in-going 
current may be seen by noting the movement of the fine particles in 
the surrounding water. This water travels down the rectangular 
channel formed by the hairs of the antennules. The function of the 
antennules in this manner is similar to that of the siphons of the 
burrowing molluscs. Both are used to carry fresh surface water to an 
animal living in the substratum. 

As the current leaves the antennules, it passes into a channel be- 
tween the branchiostegite and the body wall. In this passage lives the 
scaphognathite of the second maxilla which drives the water on to- 
ward the gills. The exopodites of the first and second maxillipeds are 
also located in this channel and have a similar function. They both have 
broad terminal segments which are articulated in such a way that they 
are able to perform a sweeping motion. This would give the water an 
additional push in the direction of the gills. After passing over the 
gills, the current of water continues posteriorly and emerges from 
under the carapace. 

The fifth pair of walking legs which are folded inside the gill cham- 
ber have a function in cleaning the gills. When specimens were opened 
for examination, a number of grains of sand were frequently found 
lodged in the filaments. The chelae of the fifth legs are capable of 
prehension and might be used for picking out the sand and any other 
material which might be present. 


SUMMARY 


The sand crab, Emerita analoga (Stimpson), inhabits the sandy 
beaches in the zone covered by the wash of the waves. This environ- 


66 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


ment, while largely free from competing species and supplying a 
plentiful amount of food in the form of plankton, requires a con- 
siderable degree of adaptation. The shifting substratum makes it 
difficult to maintain a firm and stable position and there is danger from 
the shock of the waves and from predatory animals. In order to escape 
these hazards, the sand crab is able to burrow rapidly. The first 
four pairs of walking legs and the uropods are modified for this 
function. The antennae are adapted for filtering food from the water 
and transporting it to the mouth while the animal remains buried. 
The constant wash of the waves makes it unnecessary for these crabs 
to maintain a current of water to bring in food as other plankton 
feeders are required to do. The direction of the respiratory current of 
water has been changed from that of other decapods and flows from 
anterior to posterior. The antennules form an inhalent siphon. There 
are many other less conspicuous modifications which aid the sand 
crabs in taking advantage of their environment as well as in over- 
coming the disadvantages. 


ABBREVIATIONS 
ab.som.1 first abdominal somite gill gill 
ab.som2 second abdominal somite ischio. ischiopodite 
an. anus labi. labium 
ant. antenna labr. labrum 
antul. antennule lat. lip lateral lip 
m. mouth 

apo. apodeme 
aux. pe. auxiliary piece mand. me 
baso. basopodite Tino: se soy ONE 
branch. gr. branchial groove Beas 3 ae maxilliped 
branchio.  branchiostegite ae een eee 
cara. carapace am ; Onn Peek 
carpo. carpopodite E Ss Paes 

i proto. protopodite 
cervic. gr. cervical groove eae Se . 
cits ieeeret. scaph. scaphognathite 
coxo. coxopodite ap. SeSceT 
dactylo. dactylopodite Welk ieleon 
dor.b.w. dorsal body wall Gn uropod 
endo. endopodite wk. lg. 1 first walking leg 
ELO: exopodite wk. lg. 2 second walking leg 
eye st. eye stalk wk, lg. 3 third walking leg 
flag. flagellum wk. lg. 4 fourth walking leg 
gen. op. genital opening wk. lg.5 fifth walking lege 


Functional morphology of Emerita analoga 67 


BIBLIOGRAPHY 
ALBERT, F. 
1883. Das Kaugerust der Dekapoden, Zeitschrift fiir wissenschaftliche Zoologie, 
39: 445. 
BANNER, A. H. ann D. L. McCKERNAN 


1943. A record of Emerita analoga from the Washington Coast. Science, 97 
(2509) :119. 


BENEDICT, J. E. 
1901. The Anomuran collections of the Fish Hawk expedition to Porto Rico. 
Bull. U.S. Fish Comm. for 1900, pt. 2: 131-148. 


BOOLOOTIAN, R. A., A. C. GIESE, A. FARMANFARMAIAN, ann J. TUCKER 
1959. Reproductive cycles of five West Coast Crabs. Physiol. Zool., 32:213-220. 


BRUCE, J. R. 
1928. Physical factors on the sandy beach. Part I. Tidal, climatic, and edaphic. 
J. Mar. Biol. Assoc., 15: 535-552. 


1928. Ibid. Part II. Chemical changes—carbon dioxide concentration and sulfides. 
J. Mar. Biol. Assoc., 15: 553-567. 


BUDDENBROCHE, W. von 

1913. Uber die Funktion der Statocysten im Sande grabender Meerestiere. 
Zoologische Jahrbiicher, Abteil fiir allgemein Zoologie, 33: 441-482. 

CALMAN, W. T. 

1909. A treatise on zoology. London: Adam and Charles Black, 346 pp. 


DANA, J. D. 
1852. United States exploratory expedition. Crustacea, pt. 1: 409. 


GARSTANG, W. 

1896. On the function of certain diagnostic characters of decapod Crustacea. 
Report British Assoc., 1896: 828-830. 

1896. Contributions to marine bionomics. I. The habits and respiratory mecha- 
nism of Corystes cassivelaunus. J. Mar. Biol. Assoc., 4:223. 
1897. Contributions to marine bionomics. IJ. The function of antero-lateral denti- 
culations of the carapace in sand-burrowing crabs. J. Mar. Biol., 4: 396. 
1897. Contributions to marine bionomics. III. The systematic features, habits, 
and respiratory phenomena of Portumnus nasutus (Latreille). J. Mar. Biol. 
Assoc., 4: 402. 

1897. On some modifications in Crustaceans which burrow. Quart. J. Micr. Sci., 
40: 211. 


GERSTAECKER, A. anp A. E. ORTMANN 
1901. Klassen und Ordningen der Tier-Reich, 5, Ab. 2: 1152. 


HOLMES, S. J. 
1900. California stalk-eyed Crustacea. Occ. Pap. Calif. Acad. Sci., 7: 103. 


MEAD, H. T 
1917. Notes on the natural history and behavior of Emerita analoga (Stimpson). 
Univ. Calif. Publ. Zool., 16: 431-448. 


68 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


MIERS, E. J. 
1879. Revision of the Hippidae. J. Linn. Soc. London, 14: 312. 


ORTMANN, A. E. 

1896. Die Geographische Verbreitung der Decapodengruppe der Hippidae. 
Zoologische Jahrbiicher, Syst., 9: 219. 

ORTON, J. H. 

1920. Sea temperature, breeding, and distribution in marine animals. J. Mar. 


Biol. Assoc., n.s., 12: 339-366. 


RATHBUN, M. J. ’ 

1904. Harriman Alaska Expedition, 10: 168. 

1910. Stalk-eyed Crustacea of Peru and the adjacent coast. Proc. U.S. Natl. Mus., 
38: 554. 

SAY, THOMAS 

1817. An Account of the Crustacea of the United States. J. Acad. Nat. Sci. Phila.. 
1: 155-169. 


SCHMITT, WALDO 
1935. Crustacea macura and anomura of Puerto Rico and the Virgin Islands. New 
York Acad. Sct., 15: 125-227. 


SMITH, S. I. 

1873. Report upon the invertebrate animals of Vinyard Sound. Rept. U.S. Fish 
Comm., for 1871-72: 548. 

1877. The early stages of Hippa talpoida. Trans. Conn. Acad., 3: 311-342. 

STIMPSON, W. 


1857. Notices on new species of Crustacea of western North America. Proc. 
Boston Soc. Nat. Hist., 6: 84-89. 


WEYMOUTH, F. W. ann C. H. RICHARDSON, JR. 
1912. Observations on the habits of the crustacean, Emerita analoga. Smith- 
sonian Misc. Coll., 59, (7): 1-13. 


FOSSIL ARTHROPODS OF CALIFORNIA. NO. 25. SILICIFTED 
LEAFHOPPERS FROM CALIFORNIA MOUNTAINS NODULES 


W. Dwicurt Prerce! 


It is with great pleasure that I present the studies of a series of silicified 
jassid (cicadellid) leafhoppers, Order Homoptera, all extracted by 
acids (formic and hydrochloric) from Miocene calcareous petrolif- 
erous nodules collected in the Calico Mountains, San Bernardino 
County, California, by Allison R. Palmer, Mrs. Laura Rouse, Mr. and 
Mrs. Samuel Kirkby, Mrs. Dara Kuznetzoff (as Dara Shilo), Mr. and 
Mrs. John Gibron, Sr., and Dr. Donald Weissman. 

The nodules were found at twelve sites on two sides of the mountain 
mass and probably represent quite a range in age, as there is over 300 
feet difference in altitude, although the tilting of the strata reduces 
the value of present altitude. However, other evidences indicate sites 
Gibron 4 and U.S. 19057 to be the oldest. They are all Miocene. 

For comparison of location and approximate altitude I have tabu- 
lated the material (Table 1). The collection sites are in SE and SW 
quarters of Section 23, and SW quarter of Section 24, R.1.E., T.10.N.; 
and in NE quarter Section 19, and SW quarter Section 18, R.2.E., 
T.10.N., a distance of 214 miles east and west, and about 1 mile, north 
and south. 

The classification of the Jassidae is very confusing inasmuch as 
each authority has used a different system of group classification, The 
head shows tremendous differences, due to the movement of the epis- 
tomal suture, the frons, and the antennae and ocelli. The group has 
been called by various family names, but I have chosen to follow as 
nearly as possible the system of Evans (1946-1947). 

Hitherto, the few fossil leafhoppers reported have been compressed 
by pressure of volcanic ash, or aquatic deposits. The fossils from the 
lake-bed nodules formed in volcanic areas are unique, in that they are 
three-dimensional, siliceous replicas of the insects, often still contain- 
ing traces of the petroleum that was evolved by catalytic action from 
their tissues. As the silica is colloidal, the preservation of form is per- 
fect, and in several specimens even the form of the brain and nervous 
system is clearly outlined. 

The first specimen found was described and illustrated by Palmer 


Research Associate, Los Angeles County Museum. 


69 


70 


Quarter, 


TABLE I 


Sources oF THE Lear Hopper MATERIAL 


Specimen 


Nodule Altitude | Collector 


Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


Species 


Section No. of 
and Range Nodule 
S.W.23.1 Gibron H1704 Gibron Miochlorotettix 
J gibroni gf paratype 
SiE23i1 (G.1) H1820 Gibron Miochlorotettix 
LA115 5743 gibroni 3 paratype 
SE231 | (G1) | H1561 Gibron 9 
a 115 
S.E.23.1 (G.1) H1567 Miochlorotettix 
115 | gibroni Gf paratype 
S.E.23.1 (G.2) H1431 Gibron 
116 
S.E.23.1 (G.3) H1940 Gibron Miochlorotettixz 
117 gibroni ? 
S.W.24.1 19057 5076 Gibron nymph 
S.W.24.1 19057 (H20) Gibron Miochlorotettix 
; 5278 gibroni allotype 
S.W.24.1 | 19057 (H21) | Gibron Miochlorotettix 
5279 gibroni 
S.W.24.1 | 19057 (H22) ‘| Gibron Miochlorotettix 
5280 gibroni 
S.W. 24.1 19057 alae (H23) Gibron Miochlorotettix 
5281 gibroni 
S.W.24.1 19057 (H26) Gibron Miochlorotettix 
5283 gibroni holotype 
S.W.24.1 19057 (H60) Gibron Miochlorotettix 
5296 gibroni 
S.W.24.1 19057 (H70) Gibron Miochlorotettix 
5299 gibroni 
S.W.24.1 19057A H785 Gibron 
S.W.241 19057A H899 Gibron Miochlorotettix 
| gibroni ? 
S.W.24.1 19057A | H1801 Gibron Miochlorotettix 
gibroni ? 
S.W.24.1 19057B H1354 Gibron 
S.W.24.1 19063 (H1216a) Gibron Miomesamia juliae 
5745 holotype 
S.W.241 H1216b Gibron Miochlorotettix 
gibroni ? 
S.W.24.1 19064b USNM | Palmer Euscelis palmeri 
Io 561985 
S.W.24.1 LA 38 854 |10263C R. hirkby fragment 
N.E.19.2 12 1312 2201 | Shilo Protochlorotettix 
calico Q holotype 
N.E.19.2 17F 3723 |28805 2700+- L. Rouse Phlepsius 
weissmanae ¢ holotype 
N.E.19.2 17 2142 |11806 2700-+- L. Rouse Protochlorotettix 
calico impression 
N.E.19.2 KX42b] 5787.‘ {18799 | 2700+ | S.Kirkby | Miochlorotettix 
kirkbyi 
NE192 | KX14 5788 | 2660 S.Kirkby | Miochlorotettix 
kirkbyi 
S.W.18.2 42 2491 [10528 | 2750 Weissman nymph 
4+quarter | 15 28 2400- if | 6 species 
Sections Sites Specimens 2750 collectors 


Silicified leafhoppers from California 71 


(1957), but he did not give it a name, because of the absence of a 
male specimen. In paleontology one cannot wait until we find all diag- 
nostic characters before we assign a name to a species, even though 
the generic position must later be changed, because we need a “han- 
dle” for it. I am, therefore, assigning a name to his specimen, naming 
it im his honor, and have reproduced his drawings so that the species 
can be readily compared to the others found in the nodules. 

These are the first leafhopper fossils preserved by silica replications, 
and they are truly remarkable fossils. The clear transparent nature 
of some of them makes it very difficult to correctly mterpret all 
characters. 

Leafhoppers are not aquatic, although they may visit and breed 
upon aquatic plants growing out of the waters; hence, we might con- 
sider these insects as accidentally preserved, probably bemg drowned 
by the downfall of volcanic liquids or gases. 

There are six distinct species in the series and by the figures it can 
be seen that they are readily separated by the face, the anterior out- 
line of the head, the shape and proportions of frons, and clypeus. ‘Two 
of the species, gibroni and kirkbyi are unusual in the arching of the 
prothorax due to a greatly enlarged scutellum, and both of these and 
calico show a complete median division of the sternal segment preced- 
ing the female genitalia. Nine of the specimens are assigned to one 
species, gibroni, although there is considerable variation in head 
measurements. 

Specimens 5279 and 5787 are interesting in that the brain has been 
differently crystallized and hence is quite apparent in the clear head. 

In Table 2 are listed the: measurements for comparison; and these 
are to form part of the descriptive data. 


Order HOMOPTERA Leach 1815 
Family sassipaE (Cicadellidae ) 
Subfamily EUSCELINAE 
Tribe EUSCELINI 


Palmer (1957) described and figured a female leafhopper, which Dr. 
David A. Young considered to belong to the genus Deltocephalus s.s., 
based on size and general appearances. 

However, DeLong (1926) states: “In regard to venation proper, 
the normal and apparently constant condition in the Deltocephalus 
wing is a series of three anteapical cells, with a costal cell beyond the 
outer anteapical cell” In those wings in which two cross veins inter- 


72 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


vene between the first and second sectors there are two discal cells, 
three anteapical cells, and five apical cells. 

Palmer’s leaf hopper has only one cross ven and consequently two 
discal and two anteapical cells, plus the five apical cells. For this reason 
I feel that the fossil must belong to one of the genera in which its type 
of wing venation is known to exist, as: Euscelis, Eutettex, Chlorotettix. 
Thamnotettix, Phlepsius, and Paraphlepsius. 

We have three other characters to assist in the elimination: (1 ) 
short, broad vertex, not excessively angulate; (2) wide face, with 
frons broad at base, narrowing to clypeus, which is one-half as wide 
as frons at base, and with clypeus wider at apex than base; (2) last 
sternal segment (7th) of female broadly roundingly emarginate, ex- 
posing basal parts of valves and valvifers, and with pygofer at base 
no wider than the emargination. 

All three of these characters are to be found in Euscelis deceptus 
Sanders. E. parallelus (Van Duzee). E. striolus (Fallen), and Tham- 
notettix mellus Sanders. 

Inasmuch as Palmer describes his species as having head slightly 
convex in profile, it seems to fit better into Euscelis than Thamnotet- 
tix. and hence it is here tentatively placed in the genus Euscelis. 


Genus Euscelis Brullé 
Euscelis palmeri, new species 
(Figures 6, 7, 19, and 20) 


Holotype: USNM 561985, the specimen figured and described by Dr. 
Allison R. Palmer (1957), without name, but placed tentatively in 
Deltocephalus. It is with pleasure that I assign it the species name 
palmeri. I repeat his description, adding in parentheses points derived 
from his drawings. 

Type locality: Site 19064b, U.S.G.S. Cenozoic locality, SW 14 Sect. 
24, R.1.E. T.10.N., Calico Mts., San Bernardino County, California. 

Description: Female. Length about 3.2 mm. Crown slightly pro- 
duced forward, sharply rounded anteriorly in dorsal aspect, slightly 
convex in profile; median length about three-fifths width between 
eyes. (Face with proportions of width to length as 23:18; posterior 
margin of eyes at about middle line of facial length; posterior margin 
of face a curve enclosing genae and clypeus. Frons a little more than 
twice as wide at apex as at base. Clypeus one-half wider at apex than at 
base. Lorae not completely defined. Antennae inserted between eyes 
and frons opposite middle of frons). 


Silicified leafhoppers from California 73 


Forewing with discal cells undivided; cross vein dividing costal area 
origimates from (near base of) anteapical area; proximal parts of 
claval veins not preserved; (basal cells, three; discal cells, two; costal 
cell, one; anteapical cells, two; apical cells, four). 

Femur with two distal pairs of spines, and a single more proximal 
spine arranged on knee. Anterior spines in each pair seem more slen- 
der than posterior spines. 

Seventh sternite with broad median indentation in its posterior 
margin that is flat at its anterior end except for a small centrally lo- 
cated notch. This segment has been pulled forward in fossil to expose 
first valvifers of ovipositor, and between these, smooth basal parts of 
first valves. Posterior to these, basal parts of pygofer overlap slender 
basal parts of third valves. Surfaces of third valves and pygofer gran- 
ular. Pygofer also bears strong spines on its posterior half. Tips of 
third valves extend posteriorly beyond apex of pygofer. (Three dorsal 
segments are visible beyond the pygofer ). 


Miochlorotettix, new genus 


Leafhoppers with typical deltocephaline form, but with prothorax 
strongly arched forward; and scutellum occupying the arch, as well 
as extending back between the wings to about the same distance as it 
extends forward. 

At first sight it was thought this might be a deformity of the pro- 
thorax, but identical thorax and scutellum has been found in all 
specimens. 

It is placed tentatively near Chlorotettix, because of the genitalia. 


Miochlorotettix gibroni, new species 


(Figures 1, 2, 15, 16, and 17) 
Type locality: Crystallized fossil leafhoppers, extracted by hydro- 


chloric acid from calcareous Miocene nodules from Calico Mountains, 
SW 1% Section 24, R.1.E., T.10.N., San Bernardino Co., Calif., from 
site 19057 U.S.G.S. Cenozoic locality, collected and extracted by Mr. 
and Mrs. John Gibron, Sr., of Campbell, California. 

Types: Holotype: & specimen 5283 (Gibron H26); allotype 2, 
5278 (Gibron H20); paratypes 5279 (Gibron 21), with tip of abdo- 
men broken, but probably male; 5296 (Gibron H60) head and thorax 
only; 5280 (H22); 5281 (Hi23); 5299 (H70), all from U.S. Geol. 
Site 19057; also, 5743 (H1820), H1561, H1567 from Gibron Site 1, 
near camping site on Mule Canon Drive; H1431 from Gibron Site 2, 


74. Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


Silicified leafhoppers from California 75 


2) 


across the hill on Phillips Drive side; H1704 from Gibron Site 4 far- 
ther west. Other specimens, labeled with (?) are Gibron’s H1940, 
H899, H1801, and H1216b. Types mounted in balsam, deposited in 
Los Angeles County Museum Collection; all other material in personal 
collection, John Gibron, Sr. 

Description: Measurements of all specimens given in Table 2. 

Male. Smaller than female. In the holotype the body is complete, 
with antennae; one anterior leg with femur and tibia; fragment of 
median leg; one entire posterior leg. 

The face is 34 as long as wide; frons wider than clypeus, which is 
more or less quadrate. Antennae with broad basal segment; second 
segment more than twice as long as wide; third joint elongate, slender. 

The prothorax is strongly arched forward, and the concavity of the 
arch is occupied by the anterior half of the scutellum. 

Gibron’s specimen H1704 is a dry mounted male attached on a 
slide by gum tragacanth. It has the six legs mostly intact and has been 
studied by illumination, comparative measurements being made of the 
lengths of the various leg segments using spaces in ocular micrometer, 
52 spaces=1 mm. 


CoMPARATIVE MEASUREMENTS OF THE LEGS 


leg element front legs middle legs hind legs 

coxa 8 13 — 
trochanter ae aglelie 3 = 
femur 30 5 30 
tibia 15 20 62 
tarsus (entire) 14 15 33 
tarsus I 6) — 18 

II 6 = 10 

III 6 _ 10 


The legs are armed with many long spines. The tarsal joints are 
attached before the apex of the preceding joint, and each is terminated 


by two pulvilli. 


Figure 1. Miochlorotettix gibroni, ventral view of ¢ 5283. 

a. Same. Antenna, ¢ 5283. 

b. Same. Caudal segments and hind tibio-tarsus, ¢ 5283. 
Figure 2. M. gibroni, caudal segments, 9 5279. Figure 3. Protochlorotettix calico, 
side view, 9 1312. Figure 4. Same, caudal segments, Q 1312. Figure 5. Mio- 
chlorotettix kirkbyi, caudal segments, Q@ 5787. Figure 6. Euscelis palmeri, caudal 
segments, after Palmer. Figure 7. Same, wing, after Palmer. 


76 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


Silicified leafhoppers from California rah 


Female. The allotype is larger than the holotype male but has the 
same shaped head, and the scutellum occupying the arch of prothorax. 
The specimen is broken in two parts and mounted in balsam. 

This specimen is interesting in the preservation of the brain, which 
is not crystallized in the same manner as the rest of the body. This is 
a great white mass extending the width of the base of the head, with 
long arms reaching into the eyes dorsally with two large lobes sepa- 
rated by a lower median area, and from the lobes extend a pair of fine 
nerves to the anterior margin to indicate the position of ocelli. The 
under and posterior part of the brain occupies the facial portion. 

The female genitalia also set this species aside among the Jassidae, 
in that the last ventral segment is completely divided at base into two 
lobes, broad at base, and acute at apex. The ovipositor extends beyond 
the pygofer. 

The paratypes have the head, thorax, and scutellum of identical 
pattern as described above. 


Miochlorotettix kirkbyi, new species 
(Figures 5, and 10-12) 


Holotype: 2 specimen 5787 collected by Samuel Kirkby in NW 
NE 144 Section 19, R.2.E., T.10.N., Calico Mountains, and dissolved 
from nodule 18799, Site 42B, altitude close to 2700 feet. 

This specimen is placed in the new genus Miochlorotettix because of 
the greatly enlarged scutellum displacing the prothorax forward; and 
also the complete division of the sternite preceding the female genitalia, 
with two separate sternal lobes as in gibroni and calico. However, it 
differs in the frontal curvature of the head being not as sharply angled 
as gibroni; side profile wider than gibroni or calico; face quite distinct 
from all four species included in this paper. 

Length 3.68 mm., which is shorter than the @ s of gibroni or calico. 


Figure 8. Miomesamia juliae, face, 5795. Figure 9. Same, side view of head, pro- 
thorax and scutellum, 5795. Figure 10. Miochlorottetix kirkbyi, face, 5787. Figure 
11. Same, side view of face, 5787. Figure 12. Same, dorsal view of head, prothorax 
and scutellum, 5787. Figure 13. Phlepsius weissmanae, face, 3723. Figure 14. 
Same, dorsal view of head, prothorax and scutellum, 3723. Figure 15. Miochloro- 
tettix gibroni, face, 3 5283. Figure 16. Same, dorsal view of head, prothorax and 
scutellum showing brain, Q 5278. Figure 17. Same, dorsal view of head, pro- 
thorax and scutellum. Figure 18. Protochlorotettix calico, face, 9 1312. Figure 
19. Euscelis palmeri, face, after Palmer. Figure 20. Same, head, prothorax and 
scutellum, after Palmer. 


78 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


Genus Protochlorotettix, new genus 


This genus is close to Chlorotettix, and has a normal triangular scutel- 
lum, but the last sternal segment is completely divided into two plates 
as in Miochlorotettix, which however, has the scutellum extending 
forward into the prothorax. 


Protochlorotettix calico, new species 


(Figures 3, 4 and 18) 


Holotype: specimen 1312, balsam slide mount, crystallized female, 
acid extracted by W. D. Pierce from blue-gray nodule, No. 2201, 
found by Dara Shilo, May 11, 1956, on blue-gray mine dump (LAC- 
MIP 359) altitude about 2700 feet, on lot 13, from NE 14 Section 19, 
R.2.W, T10.N., Calico Mountains, San Bernardino County, Cali- 
fornia. 

Description: Length 4.88 mm. This fine crystallized female still 
shows traces of color. The remarkable replacement of the tissues by 
the silica has preserved even the reticulate and granulate surface 
sculpture of head, legs and abdomen. 

Its measurements are compared in ‘Table 2 with the specimens of 
M. gibroni. 

It differs from Euscelis palmeri by much more angulated projection 
of vertex; clypeus quadrate; seventh sternite posteriorly divided imto 
two lobes; last dorsal segments surpassed by pygofer and valves; the 
last segment shorter. While having several characters similar to 
Euscelis deceptus, it differs from that species also in the more angulate 
head. 

The head (Figures 3 and 18), is strongly arched at a 90° angle 
(measured from tip of front to outer corners of eyes), though apically 
rounded. Ocelli on parietals between eyes and frons on the anterior 
edge of the vertex. Antennae fossae ventral, obovate, on parietal strips 
between eyes and frons, about middle of eyes. Frons slowly narrowed 
to a distinct notch, then suddenly narrowed to width of clypeus; 
marked with six transverse bands as in Euscelis ovatus and deceptus. 
Clypeus oblong, almost the same width throughout as in Euscelis de- 
ceptus. Rostrum elongate, about as long as clypeus. Lorae not well 
defined, but probably terminate at the frontal notches. Genae apically 
acute. 

The legs are not complete. Front coxae widely separated. Middle 
and hind coxae practically contiguous. 

The abdomen (Figures 3 and 4), has pygofers and valves sur- 


Silicified leafhoppers from California 79 


passing the last two dorsal segments. The last dorsal is a small palpi- 
form segment not twice as long as wide. 

The seventh ventral segment is divided at base to form two acute 
lobes, exposing the first valves and valvifers of the base of the pygofer, 
and the narrow parallel lined third valves. These valves exceed the 
length of the pygofer. 

While a little color and a very fine surface granulation is visible, 
there are no setae present on the pygofer. 

Internally the alimentary canal is visible. 

In addition to the type, specimen 2142, on nodule 11806, collected 
by Mrs. Laura Rouse in NE 14 Section 19, is a surface impression 
from which good latex molds disclose the dorsal form of the male 
leafhopper with dimensions as given in ‘Table 2. 


TABLE 2 
DIMENsIoNnsS oF Fossiz. LEar Hoppers 


Head 


Length 


Specimen Dorsal Width Length | Frontal 
Species No. 5 4 mm, 3 
Phlepsius 95° 
weissmanae 
holotype 
Miochlorotettix 
gibroni fel 90° 
holotype o | 92° 
fet 90° 
o Ls 
allotype fe) 91° 
2 — 
4 90° 
5743 2.8+ 0.14 0.98 0.80 92° 
H1704 Gull Bao = oe = 
1567 fot 3.36 0.21 0.88 0.88 — 
1801 fot 4.00 0.327 0.96 | 0.903 = 
? 1216b =a 0.48 1.04 i= 
? 5787 fe) 3.68 0.28 0.92 0.86 90° 
Miochlorotettix 5788 9 3.04 0.192 — 0.96 — 
kirkbyi 
Protochlorotettix 
calico 
holotype 1312 fe) 4.88 — — 1.32 90° 
impression 2142 ret 3.60 | 0.16 0.88 | — 90° 
Miomesamia 5745 4.45 | 0.31 1.05 1.12 90° 


juliae 


80 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


Genus Phlepsius Fieber 
Phlepsius weissmanae, new species 
(Figures 13 and 14) 


Holotype: Crystallized male extracted from Miocene nodule 28805. 
Specimen 3723, collected by Mrs. Laura Rouse, and extracted by 
Mrs. Ruth Weissman and writer. From Site 17F, Lot 333, NE 4 


California. Named in honor of Mrs. Weissman in acknowledgment of 
her great help for several years in these studies. 

Description: Size: 3.5 mm. Head length: 0.16 mm., width 0.80 
mm., face length: 0.88 mm. 

Color: although crystallized, the brown pattern of face markings 
and thoracic and abdominal sclerites makes this an unusual specimen. 

Head anteriorly broadly rounded, not extending much beyond the 
eyes. From dorsal aspect the median length of head is less than half 
the distance between the eyes. Eyes minutely granulate. 

Face elongate, about as long as wide. Frons broad at vertex; gradu- 
ally narrowed to clypeus, almost twice as long as eyes, marked with 
ten transverse colored bars which are interrupted on median line but 
mutually connected in two longitudinal stripes; two and one half 
times as long as clypeus. 

Antennae between eyes and frons at about middle of eyes; three 
basal jomts of right antenna present. 

Oscelli minute, on anterior margin at corner of eyes. 

Mandibular and maxillary plates or genae diagonal from eyes to 
clypeus. Lorae outlined by curved line from tip of clypeus to frons. 
bisecting the diagonal mandibular and maxillary plates. 

Labium divided into two segments. 


Tribe Platymetopiini 
A very unusual insect was extracted by John Gibron, Sr., which is 
unfortunately, not whole, as it lacks the posterior part of the abdomen: 
the legs, except their basal segments; and the right eye, which has 
been cleft off. Still, as it is so different from any of the other leaf- 
hopper material it should be discussed. 

Classification in the Jassidae is very difficult, but species with large 
protuberant eyes are rare. The presence of the ocelli on the anterior 
margin, and the sharp marginal profile of the head place the specimen 
in Euscelinae, the tribe being Platymetopiini, according to J. W. Evans 
1947, or Mesamiini, according to P. W. Oman 1934. 


Silicified leafhoppers from California 81 
Genus Miomesamia, new genus 


Face wide; eyes prominent, large, strongly facetted, and only slightly 
emarginate on lower side facing lorae. Dorsal outline very much as 
in Ulopa reticulata. 

Profile sharp angulate, with ocelli on the anterior margin each 
terminating a ridge, the frontal suture, which passes the antennae 
and originates at the epistomal suture. 

Frons quadrangular, wider at apex than base, medianly depressed, 
rests on epistomal suture, in front of which the beak is in three parts: 
postclypeus, preclypeus, and labium. At the sides of the clypeus are 
the somewhat swollen lorae and the genae. Antennae set at the sides 
of the frontal sutures, opposite the upper corner of the eyes. The pari- 
etals between frons and eyes wide and longitudinally ridged, and along 
the ridges are rows of round pits, probably originally setigerous. 

Crown of head depressed. 


Miomesamia juliae, new species 


Type: specimen 5795 (Gibron #1216), extracted from nodule by 
hydrochloric acid, by John Gibron, Sr., collected at U.S. Geol. Serv. 
Site 19067, in SW 14 Section 24, R.1. E., T10.N., Calico Mountains, 
by Mr. Gibron, and named in honor of Julia Gibron. 

Description: Length of specimen 3.2 mm. Probable total length 
about 4.45 mm. Length of head above 0.31 mm.; width of head 1.05 
mm.; length of face 1.12 mm. 

Before the final mounting of the specimen considerable petroleum 
was drawn out into the balsam. 


AN UNIDENTIFIED LEAF HoprPEerR 


A more fragmentary specimen (859 from nodule 10263), consisting 
of a crystallized thorax, one anterior and one posterior wing, from a 
nodule found by Mrs. Ruth Kirkby at Site 38 (LACMIP 386), in the 
southeast 14, southwest 14. Section 24, R.1.E., T:10.N., in the Canon 
south of that in which D. palmeri was found. The fragment measures 
6.0 mm., and is hence different from either of the preceding, but as it 
has no diagnostic characters, cannot even be assigned to a genus. 


ee —_ —E 


82 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


LirERATURE CITED 
DE LONG, DWIGHT M. 
1926. Type and venation of wings as factors in separating certain Deltocepha- 
loid genera. Ohio J. Sci., 26(1): 42-48. 


EVANS, J. W. 
1946-1947. A Natural classification of leaf hoppers (Jassoidea, Homoptera), 
Trans. Roy. Ent. Soc., London, 96:47-60; 97:39-54; 98:105-271. 


PALMER, ALLISON R. 
1957. Miocene Arthropods from the Mojave Desert, California. Contributions to 
General Geology. U.S. Geol. Surv., Professional Paper 294-G, p. 258. 


A CONTRIBUTION TO THE BIOLOGY OF THE 
GRAY GARDEN SLUG* 


R. O. Arras? and H. H. Crowe? 


Oregon State University, Corvallis 


INTRODUCTION 


The gray garden slug, Deroceras reticulatum (Miller), is a pest of 
considerable economic importance in fields and gardens of the Pacific 
Northwest. The mild winters and generally moist springs of western 
Oregon are particularly favorable to the growth and reproduction of 
this introduced species. Control by the use of standard metaldehyde 
baits has not been satisfactory under all conditions of weather, or when 
certain crops or horticultural practices are involved. Toxicological 
studies have been hampered by the difficulty in maintaining a labora- 
tory culture of test animals. This paper documents some aspects of 
slug biology and describes techniques developed during an attempt to 
rear the gray garden slug under laboratory conditions. 


PARASITES OF SLUGS AND THEIR CONTROL IN THE LABORATORY 


Most workers have had to depend for their test animals on field- 
collected slugs which usually are abundant only in the fall and spring. 
Several workers (Meggitt, 1916; Lovett and Black, 1920; Reynolds, 
1936; RKozloff, 1956a, 1957) reported that they could not keep slugs 
alive in the laboratory long enough to do comprehensive studies. Sivik 
(1954) reported a technique for obtaining eggs from slugs by con- 
fining them in wooden boxes containing moist soil and covered with 
gauze, but he did not state how long he could maintain a healthy stock 
culture. This method was tried, but high mortality occurred within a 
two week period. A modified version of this technique, however, was 
used in subsequent studies as a standard method of rearing slugs for 
limited lengths of time. Plastic boxes (refrigerator vegetable crisper 


1Approved for publication as technical paper number 1479, Oregon Agricul- 
tural Experiment Station. Part of a thesis presented in partial fulfillment of the 
requirements for the degree of Master of Science in Entomology at Oregon State 
University. Accepted for publication January 2, 1963. 


2Now with the Research Department, Niagara Chemical Division, Richmond, 
California. 


’Associate Professor of Entomology. 


83 


84 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


chests), provided with two screen-covered ventilation holes in the 
cover and a layer of moist soil or vermiculite, proved to be convenient 
containers for this purpose. 

Examination of the bodies of dead and dying slugs in the rearing 
boxes revealed the presence of fungi, bacteria, nematodes, and pro- 
tozoa. Steam sterilization of the soil in the containers failed to reduce 
the slug mortality, indicating that these organisms were being intro- 
duced into the boxes with the field-collected slugs themselves. It was 
not the purpose of this study to determine the pathogenicity of the 
parasitic organisms found, but to seek ways of eliminating them as 
possible causes of slug mortality. 

The fungus associated with dead slugs was identified as a species 
of Fusarium. A fungus found attacking slug eggs, and preventing 
their development, was identified as a species of Arthrobotrys. To 
suppress spread of the fungi and bacteria in the rearing containers, 
water containing 200 parts per million of Agri-mycin 100 (strepto- 
mycin 15% and terramycin 1.5% )* combined with 200 parts per 
million of captan fungicide (N-trichloromethylthio tetra-hydroph- 
thalimide) was added to the soil. The transfer of bacteria and fungi 
from eggs to newly hatched slugs was prevented by dipping the eggs 
in an aqueous solution containing the same amount of Agrimycin 100 
and captan. This procedure also prevented bacterial and fungal growth 
during the incubation period of the eggs. 

Slug mortality continued in the rearing containers, however, and 
the dead and dying slugs were found to be infested with nematodes 
and protozoa which had apparently not been eliminated by the above 
mentioned treatment. The nematodes, determined as Rhabiditus c.f. 
lambdiensis Maupas, Panogrolaimus spp., and Diplogaster spp., were 
usually found beneath the mantle of the slugs and when present in 
large numbers, were occasionally observed protruding from the dorsal 
surface or in the respiratory orifice. All stages of the nematodes could 
be found in slugs at any one time and were also observed free-living 
in the soil of the rearing containers. Transfer of protozoa and nema- 
todes from the parent slugs to their offspring can be accomplished by 
way of the eggs. Both nematodes and protozoans were observed on 
the moist surfaces of the eggs. Attempts to sterilize the eggs with 
chemical solutions, however, were not successful. Egg proteins were 
apparently denatured at very low concentrations of various alcohols 
and of other chemicals tested. 


‘Obtaimed from Charles Pfizer & Co., Inc., Brooklyn 6, N.Y. 


Biology of the gray garden slug 85 


A treatment which was effective in eliminating the nematodes from 
the infected slugs themselves rather than from the eggs was developed. 
Juvenile slugs were freed of their nematode parasites by confinement 
on a special agar-base medium over a two-week period of time. The 
complete medium was compounded by combining three separate 
fractions. The first fraction consisted of agar, 20 grams, combined with 
Bel-Ais salt mixture, 100 milliliters, in 200 milliliters of water. The 
second was a mixture of powdered skim milk, 50 grams; dextrose, 
200 grams; and bacto-yeast extract, 10 grams; in 400 milliliters of 
water. The third fraction consisted of boiled potatoes, 200 grams, m 
400 milliliters of water, of which only the extract was used. These 
three fractions were autoclaved separately at 15 lbs. per square inch 
pressure for 25 minutes. Agri-mycin 100, 2.68 grams, and methyl- 
p-hydroxybenzoate®, 0.375 grams (or a final concentration of 
0.025%), were added after the autoclaving process. The fractions 
were then thoroughly mixed. At a concentration of .05% of methyl- 
p-hydroxybenzoate, there was approximately 45% mortality, mostly 
of young slugs. At .025%, all stages of slugs survived. Although slugs 
fed readily on this medium, they gradually lost weight during the two 
week period. Following treatment they were transferred, for other 
studies, to lettuce leaves in the plastic crisper-chest rearing boxes. 

To further test the effectiveness of methyl-p-hydroxybenzoate, 
nematodes were isolated from the slugs and placed in petri dishes con- 
taining the above mentioned medium. All nematodes were dead after 
five hours exposure. Nematodes in check plates, containing the nu- 
trient media without the methyl-p-hydroxybenzoate, increased in 
size and numbers over a period of several days. 

The protozoan, identified:as Tetrahymena limacis (Warren), was 
not eliminated by the methyl-p-hydroxybenzoate treatment. Protozoa 
and nematodes were usually found together in individual slugs, but 
one form would dominate the other in numbers. When the nematodes 
were eliminated from the slugs by the treatment mentioned above, 
the number of protozoans increased greatly. Although Kozloff (1957) 
noted that these protozoans inhabited the slug’s digestive gland pri- 
marily, they appeared to be present not only in the various internal 
organs, but in the subcutaneous tissue as well. According to Kozloff 
(1956b), a gray garden slug is able to support a large population of 
T: limacis without the slug’s normal biological activities being dis- 
rupted. Kozloff concluded that 7: limacis is not very pathogenic. A 


>Obtained from Eastman Kodak Co., Rochester, N.Y. 


86 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


closely related species, Tetrahymena (Paraglaucoma) rostrata (Kahl) 
Corliss, is a histophagic parasite of enchytreid worms, however, and 
infection results in the death of the host (Stout, 1954). If we accept 
Rozloff’s conclusions regarding T. limacis, we would expect greater 
longevity from the nematode-and-fungus-free slugs in the rearing 
containers. Actually, the slugs did live several weeks longer than 
previously, but mortality was still heavy. Moribund and dead slugs 
were found to be heavily infested with the protozoans. The protozoan 
parasites could not be controlled by the methyl-p-hydroxybenzoate 
treatment, and since they appeared to be pathogenic, new treatments 
were sought for their elimination from the slug cultures. 

Cleveland (1925) was successful in eliminating protozoa from 
termites by subjecting them and their hosts to pure oxygen pressures. 
A modified version of Cleveland’s method was tested against 7. 
limacis in the gray garden slug. It was found, however, that an 
oxygen pressure of 42 psi for 16 hours caused 40% slug mortality 
without freeing the survivors of the protozoan parasites. Even though 
eradication of this parasite from its host was not accomplished, slug 
cultures were maintained long enough to collect considerable informa- 
tion on slug biology under laboratory conditions. 


OVIPOSITION AND EcG VIABILITY 


Various parasites, a limiting factor in the rearing of slugs, were found 
to be excreted in the feces. Early in the investigation, slugs were 
placed in plastic crisper-chests on screen trays to facilitate sanitation 
by washing the excrement through the screen at various intervals of 
time. The water remaining in the bottom of the chests maintained a 


TABLE 1 


Oviposition record for five sexually-mature slugs on soil 
at each of five different saturation percentages. 


Observations Numbers of eggs laid at saturation per cents of 

(one day intervals) 10 25 50 75 100 
Ist 0 0 32 41 0 

2nd 0 0 48 72 0 

3rd 0 45 62 36 0 

4th 0 0 0 51 41 

5th 0 56 23 0 0 

Totals 0 101 165 200 41 


Biology of the gray garden slug 87 


high humidity, but the slugs would not lay eggs under these condi- 
tions. When the screen layer was replaced with a layer of damp soil, 
however, the slugs readily deposited ege masses. 

The range of soil moisture content favored for oviposition was in- 
vestigated in the laboratory utilizmg information from the above 
observations. Oven-dried sandy loam soil was placed in jars and the 
moisture levels adjusted to 10, 25, 50, 75, and 100 per cent saturation 
(as determined by the method of Richards, 1954). Mature slugs from 
screen-bottomed plastic crisper-chests were placed in the jars of soil 
for oviposition. Results of the trial (Table 1) indicated that a soil mois- 
ture content of approximately 75% was most suitable for oviposition. 
Carrick (1942) came to a similar conclusion and stated further that 
normal egg development will not take place in soils below 10% satura- 
tion, nor in soils close to 100% saturation. In the field, egg masses of 
Deroceras reticulatum are usually deposited in cavities of the damp 
soil in the top three inches. According to Karlin and Naegele (1960), 
eggs of D. reticulatum can withstand complete submergence for at 
least four days without affecting normal development. The selection 
by slugs of soil for oviposition substantially below 100% saturation 
would tend to place most egg masses in locations not saturated for long 
periods during the rainy western Oregon winters. 

The incubation period for D. reticulatum eggs has been reported to 
range from about 15 to 96 days (Taylor, 1907; Hawley, 1922; Car- 
rick, 1938; Lovett and Black, 1920; and Binney, 1878). Approxi- 
mately 3000 eggs, deposited in the laboratory within a 24 hour period, 
were observed at a constant temperature of 20°C. The minimum 
period for incubation was 11 days, with a maximum of 21 and a mean 
for 60 groups of 50 eggs each of 15.5 days. These observations on the 
variation in development time can be extended to include the develop- 
ment of eggs from a single clutch. Rarely do all the eggs of a single 
egg mass deposited by one slug hatch on the same day. It was further 
observed that eggs of a single clutch that remained unhatched 12 days 
after the initiation of hatching, failed to develop. Microscopic examina- 
tion of these remaining eggs revealed the absence of the sperm body 
described by Carrick (1938) as characteristic of fertile eggs. In some 
clutches approximately half of the eggs were found to be infertile. 

During embryonic development, slug eggs exhibit an increasing 
ability to withstand extremes of temperature. When first laid, the egg 
is very susceptible to freezing and to temperatures higher than 22°C. 
Eggs containing embryos which have almost completed their develop- 
ment can withstand a temperature of 0°C for several days, and after- 


88 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


ward resume development when placed at higher temperatures. This 
fact facilitates the storage of eggs in refrigerators for long periods of 
time. In the present study, approximately 1000 eggs in the advanced 
stage of development, kept in a refrigerator at 4°C for seven months. 
commenced hatching within 24 hours after removal to room tem- 
perature. 

The eggs of D. reticulatum will desiccate and the embryos die unless 
contact is maintained with a moist surface. There are reports in the 
literature of eggs being completely desiccated for weeks (Lovett and 
Black, 1920), or even for years (Binney, 1878), and on exposure to 
moisture again, these eggs regained their spherical form and resumed 
normal development. Carrick (1942) remarks that these reports are 
too fantastic for credence. In this study, several thousand eggs were 
allowed to desiccate in petri dishes, and were kept in this condition at 
room temperatures for five months. At the end of this period, when 
water was added to the filter paper in the dishes, the eggs rapidly 
regained their original form and turgidity, but no embryonic develop- 
ment took place during a two-month observation period. Similar re- 
sults were obtained by Carmichael and Rivers (1932) in desiccation 
studies on the eggs of Limax flavus. 


GrowTH RaTEs AT CONSTANT AND FLUCTUATING TEMPERATURES 


Slug activity in the field is associated with conditions of high humidity 
and this association has led to an assumption that activity is im some 
way induced by damp conditions. Crozier and Pilz (1923) showed, 
however, that the speed of locomotion of Agriolimax campestris Bin- 
ney varied directly with temperature. Dainton (1943; 1954) showed 
that certain changes in temperature, not conditions of high humidity. 
induced locomotor activity. In her experimental work, in which she 
employed a more or less gradual alternation between two constant 
temperatures, she showed that activity was sharply stimulated by fall- 
ing temperatures below about 21°C and by rising temperatures above 
21°C. Activity subsided as soon as the temperature was maintained 
constant at any level. Temperature changes in the reverse direction 
had no such effect. As at constant temperature, activity was low when 
the temperature was rising toward 21°C or falling from above this 
value. Dainton assumed that increased acquisition of food necessarily 
accompanied increased locomotor activity. 

It was of interest in this study to determine whether a substantial 
increase in growth rate could be induced by certain defined tempera- 


Biology of the gray garden slug 89 


ture conditions, since rapid production of mature slugs from eggs was 
of importance. Most experimental investigations on the effect of tem- 
perature on the rate of development of an organism are conducted 
with a series of constant temperatures. Constant temperatures are rare 
in nature and field slugs are certainly exposed to almost constantly 
fluctuating temperatures due to diurnal and seasonal changes, as well 
as to non-periodic changes in weather. In view of this fact, and to test 
Dainton’s assumption regarding increased feeding associated with 
increased locomotor activity, a series of experiments were conducted 
to compare growth rates of D. reticulatum at both constant and fluc- 
tuating temperatures. 

A system for constantly fluctuating the temperature of a rearing 
chamber was devised by utilizing a Fenwal differential expansion 
thermal switch activated by a pair of metal arms attached to the 
rotating disc of a 24 hour electric timer. After the desired maximum 
and minimum temperatures had been preset, this mechanism pro- 
duced one full cycle of continuous temperature change in a 24 hours 
period by activating a Cenco refrigerating incubator.* 

The growth rates of newly hatched slugs were studied by placing 
one-day-old individuals under a constant temperature of 20°C. An 
equal number of slugs were subjected to fluctuating temperature con- 
ditions of 17° to 24°C (approximately 0.6°C change per hour). 

The results after 45 days (Table 2) showed no marked differences 
in mean weights of the slugs under the two temperature conditions. 
The test also demonstrated the great variation in growth rates possible 
for individual slugs. Some individuals exhibited practically no growth 
during this period, since in 100 observations of newly hatched slugs, 
the mean weight had been found to be 1.8 milligrams, the same 


TABLE 2 


Comparison of weight increases of immature gray garden slugs 
reared under two sets of temperature conditions for 45 days. 


Constant Fluctuating 
Temperature Temperatures 
(202 G,) (17°-24° C.) 
Mean (of 60 slugs 24.8 mg. 26.1 mg. 
for each condition) 
Maximum 83.8 mg. 73.0 me. 
Minimum 4.4 mg. 1.8 mg. 


®Central Scientific Company, Chicago, Illinois. 


Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


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Biology of the gray garden slug Q1 


weight as the minimum found under the fluctuating temperature con- 
ditions of this experiment. 

In another trial of the same temperature conditions, very similar 
results were obtained. The weights of slugs used in this second trial, 
and in further studies, were 25 milligrams or greater because of the 
difficulties encountered in handling smaller individuals. The results 
from weekly weighings (Fig. 1) again showed that growth rates under 
the two temperature conditions were not markedly different. How- 
ever, the slugs under the fluctuating temperature conditions were 
observed to be active more of the time and their food consumption 
during the 4 week test period was noticeably greater. 

A third comparison of growth rates was made in which 40 im- 
mature slugs were placed under fluctuating temperatures of 20° to 
27°C and an equal number under a constant temperature of 17°C. 
Mean weights of the slugs were recorded on a weekly basis for four 


weeks. 


Weight In mge 


Time In weeks 


Figure 3. Comparison of growth rates of immature slugs under four different con- 
stant temperatures. 


Q2 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


The results (Fig. 2) indicated a more rapid growth rate for the 
slugs maintained at a constant temperature, in spite of the fact that 
the more active slugs under the higher and fluctuating temperature 
condition consumed almost twice as much food as their counterparts 
at 17°C. It was concluded that, while the subjection of slugs to fluc- 
tuating temperatures increased their activity and food consumption 
(as indicated by Dainton, 1954), this procedure was not promising 
as a means for obtaming rapid maturation of slugs in laboratory 
cultures. 

Growth rates of slugs at four different constant temperatures were 
also studied. Twenty-five slugs were placed in each of four tempera- 
ture controlled cabinets which were held constant at 5°, 10°, 15°, and 
28°C, respectively. The mean weights of the slugs were recorded 


950 Ue 
900 © 
850 
800 
A © 


500 OX. 
_— 
450 a © 
ae @--Maximum weight 


@--Mean weight (25 slugs) 


Weight tna mge 


300 ae 


250 Vi O--Mintmum wei ght 


0 ! 2 3 4 5 6 7 8 
Time In weeks 


Figure 4. Variation in weight-gain by 25 4-month-old slugs held in a fluctuating 
temperature cabinet at 17° to 24° C for eight weeks. 


Biology of the gray garden slug 93 


over a four week period. The results obtained (Fig. 3) showed that the 
slugs at the temperatures of 10° and 15°C were of almost comparable 
weight after the third week. The growth rate was somewhat low at 
5°C. The slugs at 28°C decreased in weight after an initial increase. 

The experimental animals displayed great individual variation in 
growth rates in the above observations. To study this variation, a 
random sample of 25 4-month old slugs was placed in a cabinet at 
fluctuating temperatures of 17° to 24°C. An ample supply of food 
(lettuce leaves) was available at all times. The slugs were weighed 
individually at weekly intervals over an eight week period. The 
maximum, minimum, and mean weights were plotted for comparison 
(Fig. 4). These results, and those shown in Table 2, indicate the tre- 
mendous growth rate variations which can occur in a slug population 
from eggs hatched on the same day. The reason for the decrease in 
weight after the sixth week (Fig. 4) is not known. 


©) “90S eanshant 
temperature 


@ --17-24°C fluctuating 
750 temperatures 


Time tn weeks 


Figure 5. Comparison of weight-loss by starving slugs under constant and fluc 
tuating temperature conditions. 


04 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 
STARVATION STUDIES 


Immature slugs can withstand extended periods without food, pro- 
viding other conditions are favorable. Lovett and Black (1920) re- 
ported keeping nine slugs for one month without food. To test toler- 
ance to starvation, five groups of ten one-week old slugs were placed 
in jars containing moist, unwashed river sand at a constant tempera- 
ture of 20°C. Water (contaming the previously mentioned antibiotics 
to suppress fungal and bacterial growth) was added occasionally to 
maintain moist conditions. Slug mortality in these jars was not com- 
plete until after a five month period. 

The effects of starvation on mature slugs was also investigated, but 
in this case, the loss in weight under two temperature conditions was 
studied. Field-collected slugs were fed on lettuce for one week, then 


Wetght Ina mge 


O. lo? 3 4.556 "7 98 “Oo LOnIiomts 


Time In weeks 


Figure 6. Growth record of a single slug, reared from the fourth to the seventh 
month of life in a fluctuating temperature cabinet at 17° to 24° C, 


Biology of the gray garden slug 95 


divided into two groups of ten each. One group was kept at 20°C and 
the other at a 17° to 24°C fluctuating temperature. The slugs were 
weighed individually at weekly intervals and the mean weights 
plotted after five weeks (Fig. 5) when mortality from disease ter- 
minated the experiment. The results showed a steady decline in weight 
under starvation conditions with the rates of weight loss (slopes) 
under the different temperature conditions being approximately equal. 
This situation was unexpected, since the slugs under the fluctuating 
temperature conditions exhibited the increased locomotor activity 
similar to that observed in previous experiments. 


MAatTuRATION PERIOD 


General observation of gray garden slugs in the field shows that sexual 
maturity is reached considerably ahead of maximum growth. The 
length of time required to attain maximum size naturally varies with 
environmental and other conditions, and may or may not coincide 
with the total length of life. The published accounts of longevity of 
D. reticulatum are fairly consistent, ranging from 18 months to two 
years (Taylor, 1907; Lovett and Black, 1920; Theobald, 1895; Cook, 
1895; and Hawley, 1922). No mention of the weight of the mature 
specimens of this species, however, was found in the literature. 

The ultimate size attainable by this species was indicated in one 
trial on the effects of fluctuating temperatures, in which a single 
specimen survived for 13 weeks. This slug was reared in a 17° to 24°C 
fluctuating temperature cabinet and was weighed weekly from the 
time it was four months old until its death three months later. The 
results (Fig. 6) mdicated that the slug reached maturity when 6 
months of age, weighing approximately 2.2 grams, at that time. The 
mature specimen measured 50 millimeters long and 9 millimeters 
wide when extended, a size considerably larger than that normally 
seen in fields of western Oregon. 


96 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 
ACKNOWLEDGMENTS 


The assistance of the following persons during the course of this study 
is gratefully acknowledged: Dr. G. Dallas Hanna and Dr. A. G. 
Smith, of the California Academy of Sciences, San Francisco, for con- 
firmation of our identification of Deroceras reticulatum (Miller) ; 
Dr. E. N. RKozloff of Lewis and Clark College, Portland, Oregon, for the 
identification of the protozoa; Dr. H. J. Jensen and Dr. Donald 
Murphy of the Botany and Plant Pathology Department, O.S.U., for 
identification of the nematode parasites; and Dr. Mathew Nadaka- 
vakaren of the Botany and Plant Pathology Department, O.S.U., for 
classifying the fungi. 


LITERATURE CITED 


BINNEY, W. G. 
1878. The terrestrial air-breathing mollusks of the United States and the adjacent 
territories of North America. Bull. Mus. Comp. Zool., 4:1-439. 


CARMICHEAL, E. P ann T. D. RIVERS 

1932. The effects of dehydration upon the hatchability of Limax flavus eggs. 
Ecology, 13:375-380. 

CARRICh, ROBERT 

1938. The life history and development of Agriolimax agrestis L., the gray field 
slug. Trans. Roy. Soc. Edinburgh, 59:563-597. 

1942. The gray field slug, Agriolimax agrestis L., and its environment. Ann. 
Applied Biol., 29:43-55. 

CLEVELAND, L. R. 

1925. The effects of oxygenation and starvation on the symbiosis between the 
termite Thermopsis, and its intestinal flagellates. Biol. Bull., 48:455-468. 


COOK, H. H., A. E. SHIPLEY ann EF R. C. REED 
1895. Molluscs and Brachiopods. (The Cambridge Natural History, vol. 3). 
London, Macmillan, 535 p. 


CROZIER, W. J. ann G. E PILZ 
1923. The locomotion of Limax. 1. Temperature coefficient of pedal activity. 
J. Gen. Physiol., 6:711-721. 


DAINTON, BARBARA H. 

1943. Effects of air currents, ight, humidity and temperature on slugs. Nature, 
1511225: 

1954. The activity of slugs. 1. The induction of activity of changing tempera- 
tures. J. Exper. Biol., 31:165-187. 

HAWLEY, I. M. 

1922. Insects and other animal pests injurious to field beans in New York. New 
York. p. 977-999. (Cornell University. Agricultural Experiment Station. 
Memoir 55). 


Biology of the gray garden slug 97 


KARLIN, E. J. anp JOHN A. NAEGELE 
1960. Biology of the Mollusca of greenhouses in New York state. Cornell Agr. 
Exp. Sta. (Ithaca), Memoir 372. 35 p. 


KOZLOFF, EUGENE N 

1956a. Experimental infection of the gray garden slug, Deroceras reticulatum 
(Miller) by the holotrichous ciliate Tetrahymena pyriformis (Ehrenberg). 
J. Proto-zool., 3:17-19. 

KOZLOFF, EUGENE N. 

1956b. Tetrahymena limacis from the terrestrial pulmonate Gastropods, Mona- 
denia fidelis and Prophysaon andersoni. J. Proto-zool., 3:204-208. 

1957. A species of Tetrahymena, parasite in the renal organ of the slug Deroceras 
reticulatum. Proto-zool., 4:75-79 

LOVETT, R. L. ann A. B. BLACK 

1920. The Gray garden slug with notes on allied forms. Corvallis. 43 p. (Oregon 
Agricultural Experiment Station. Station Bulletin 170). 

MEGGITT, E A. 

1916. A contribution to the knowledge of the tape worm of fowls and of sparrows. 
Parasitology, 8:390-409. 


REYNOLDS, B. P. 
1936. Colpoda steini, a facultative parasite of the land slug, Agriolimax agrestis. 
J. Parasit., 22:48-53. 


RICHARDS, L. A. (Editor) 
1954. Diagnosis and improvement of saline and alkali soils. United States De- 
partment of Agriculture Handbook No. 60:83-126. 


SIVIK, FRANK P. 
1954. A technique for slug culture. The Nautilus, 67:129-130. 


STOUT, J. D. 
1954. The ecology, life history and parasitism of Tetrahymena (Paraglaucoma) 
rostrata (Kahl) Corliss. J. Proto-zool., 1:211-215. 


TAYLOR, JOHN W. 
1907. Monograph of the land and freshwater Mollusca of the British Isles. Vol. 2. 
Leeds, Taylor brothers. 312 p. 


THEOBALD, FRED V. 
1895. Mollusca injurious to farmers and gardeners. Zoologist, 19:201-211. 


ty SNE, 
=e 
o = : 
is = 
i 
= i 
\ 
i s 


SURF-RIDING BY THE CALIFORNIA GRAY WHALE 


Notice of the ability of cetaceans to utilize forward wave motion and 
to surf-ride is of interest to workers in the field of hydrodynamics as 
well as animal behavior. 

This ability has been reported in the small odontocete delphinids 
Tursiops truncatus (Montagu) and 7? gilli Dall. For the former, the 
report was made by Caldwell and Fields (J. Mamm., 40: 454, 1959) 
and Norris and Prescott (Univ. Calif. Publ. Zool., 63: 300, 1961), and 
for the latter by Norris and Prescott (loc. cit.). These reports showed 
that the same individuals repeated the performance several times, in- 
dicating that the surf-riding was not an accidental encounter but a self- 
reinforcing form of play behavior. 

We now report similar behavior by the large mysticete, Eschirich- 
tius glaucus (Cope), the California gray whale. 

Although both Scammon (The marine mammals of the north-west- 
ern coast of North America etc., 1874, p. 24) and Caton (Amer. Nat., 
22: 510, 1888) referred to extensive play by this species in the break- 
ers along the outer coast of Baja California, they made no specific 
mention of surf-riding. 

The following behavior has been reported to us by Robert V. Bell, 
leader of a party of salvage divers who spent seven weeks in Baja 
California in the spring of 1958. They were anchored just south of 
Magdalena Bay in the channel which divides Santa Margarita and 
Cresciente Islands. Just west of the latter was a large area of shoal 
water and the divers were anchored about 50-60 yards from these 
shoals. 

The divers reported that almost daily during the seven-week period 
they watched gray whales ridig the breakers coming in over the 
shoal. The divers were particularly interested in the behavior and 
agreed among themselves at the time that the whales were surf-riding 
“Just as a man would”’; 7.e., just in front of the crest of the waves. 

The whales engaging in the activity were adults. It was not noted, 
however, whether the same individuals rode more than once.—Davip 
K. CaLtpweti and Meza C. CaLpwe., Los Angeles County Mu- 
seum, Los Angeles 7, Calif. 


99 


ve 


- 


MOLLUSCS FROM PACIFIC NORTHWEST ARCHAEOLOGICAL 
SITES, 2. WASHINGTON: 45-CA-30, A COASTAL 
SHELLMIDDEN IN THE OZETTE AREA 


Ropert J. DRAKE 


The University of British Columbia 


INTRODUCTION 


Site 45-CA-30 was excavated over a period of six weeks in the summer 
of 1961 (Borden 1962: 611); it is located in the Olympic National 
Park near the northwest corner of Washington (see Fig. 1-D) near 
Ozette in Clallam County. It was investigated under the direction of 
Richard D. Daugherty of Washington State University with support 
from the National Park Service by means of a grant. 

In May of 1962, the reported upon mollusc and barnacle samples 
from the 45-CA-30 excavations were submitted for study and report, 
accompanied by a profile plan of the small shellmidden and a regional 
location map. A letter from Stanley J. Guinn (dtd. 30 May 1962) 
contained a description of the site and an account of molluscan remains 
in relation to factors of its stratigraphy. 

45-CA-30 was shown, by the excavations of 1961, to have been a 
specialized habitation location apparently occupied primarily for sea 
mammal hunting from late prehistoric into postcontact times. ““Whale 
and seal bones were especially abundant” The midden was stratified, 
of about 5 feet in depth, and made up of animal remains and fire 
broken rock. The greater portion of the recovered artifacts was har- 
poon parts and composite fishhook portions. (Borden 1962: 611). 

Size of the shells, as seen in the midden deposit to lessen toward 
the top, seemed to reflect the greatest change over time. Alteration of 
form of a particular species was not evident. The main shell com- 
ponents were of clams and mussels; limpets, chitons, barnacles, sea- 
urchins, and snails being less frequent. There were less deposits of 
limpets, barnacles, chitons, seaurchins, and snails in the upper levels; 
clam and mussel shells, however, persisted. (in litt., 30 May 1962, 
Stanley J. Guinn). 


PROCESSING AND STUDY 


Upon receipt, the material was processed as a study unit in a program 
of investigation of animal remains from archeological sites. The assign- 
ment of reference numbers (usually referred to as “permanent study 


101 


102 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


4 


Ss cm 


OLYMPIC 
ES SS PENINSULA 
C. SCHIZOTHERUS NUTTALLII 


FiG | MARINE BIVALVES 
SITE LOCATION 


Figure 1. Marine bivalve shells from site 45-CA-30, Washington; location of site. 


numbers’’) of the shells was unnecessary as each lot had been num- 
bered and invoiced before shipment. Therefore, references to samples 
follow the numerical designations in the records of the Department of 
Anthropology of Washington State University. The shells and bar- 
nacles were returned to the Department of Anthropology at Pullman. 


Molluscs from archaeological sites 103 


10mm 
foxceece 
A TEGULA FUNEBRALIS : B.THAIS LIMA 


C NERFTUNEA TABULATA D. OLIVELLA BIPLICATA 


eng 
E. ACMAEA PELTA 


iOmm 
Et 
F ACMAEA MITRA 
BiG e2 : MARINE GASTROPODS 


Figure 2. Marine gastropod shells from site 45-CA-30, Washington. 


MAR InE BIVALVE SHELLS 


Protothaca lacinata (Carpenter), 1864; (45-CA-30/8)—Fig. 1-B. 
References: 
Morris 1952: 47-48; pl. 11, fig. 6. 
Palmer 1958: 96-97. 


104. Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 


Remarks: 
Lives near shore, probably mostly in shallow water. 
Mytilus californicus Conrad, 1837; (45-CA-30/11) —Fig. 1-A. 
Reference: 
Morris 1952: 20; pl. 6, fig. 10. 
Remarks: 
Lives mostly in the tidal zone. 
Schizotherus nuttallii (Conrad), 1837; (45-CA-30/5) —Fig. 1-C. 
Reference: 
Morris 1952: 57; pl. 17, fig. 1. 
Remarks: 
Lives buried in mud in the tidal zone. 


MarINE GaAsTROPOD SHELLS 
Tegula funebralis (Adams), 1854; (45-CA-30/14) —Fig. 2-A. 


Reference: 
Morris 1952: 81; pl. 22, fig. 8. 
Remarks: 
Lives in the tidal zone. 
Thais lima Gmelin, 1791; (45-CA-30/11) —Fig. 2-B. 
Reference: 
Morris 1952: 119; pl. 28, fig. 10. 
Remarks: 
Lives near shore, probably mostly in shallow water. 
Neptunea tabulata (Baird), 1863; (45-CA-30/17) —Fig. 2-C. 
References: 
Oldroyd 1927 (vol. 2, pt. 1): 227-228; pl. 18, fig. 4. 
Morris 1952: 126; pl 29; fie 11. 
Remarks: 
Probably lives near shore and mostly im shallow water, but oc- 
casionally deeper than the tidal zone. 
Olivella biplicata (Sowerby ), 1825; (45-CA-30/16) —Fig. 2-D. 
Reference: 
Morris 1952: 134; pl. 30, fig. 20. 
Remarks: 
Lives in sand in the tidal zone. 
Acmaea pelta Eschscholtz, 1833; (45-CA-30/2) —Fig. 2-E. 
Reference: 
Morms 1952.77.15 pl to nose 
Remarks: 
Lives in the tidal zone. 


Molluscs from archaeological sites 105 


5 com 
B. BALANUS CARIOSUS 


10mm 


oe 
C. HAPLOTREMA VANCOUVERENSE 


FIG 3 BARNACLES & TERRESTRIAL 
GASTROPOD 


Figure 3. Barnacles and terrestrial gastropod shell from site 45-CA-30, Washing- 
ton. 


Acmaea mitra Eschscholtz, 1833; (45-CA-30/6) —Fig. 2-F. 
Reference: 
Morris 1952: 71; pl. 19, fig. 4. 
Remarks: 
Lives in the tidal zone. 


106 ~=Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 
BARNACLES AND TERRESTRIAL GASTROPOD SHELL 


Coronula reginae Darwin, 1854; (45-CA-30/42) — a 3-A. 
References: 
Darwin 1854: 419-421; pl. 15, fig. 5. 
Cornwall 1955a: 54-55; illu. 
Cornwall 1955b: 43-44, fig. 35. 
Remarks: 
Lives on whales, mostly killer whales. 
Balanus cariosus (Pallas) 1788; (45-CA-30/12) —Fig. 3-B. 
References: 
Darwin 1854: 273-275; pl. 7, fig. 3a. 
Pilsbry 1921: pl. 20, figs. 3, 6. 
Cornwall 1955a: 22 (allu., fig. 60n p. 15). 
Cornwall 1955b: 26-28; fig. 19. 
Remarks: 
Lives mainly on rocks in or near the tidal zone. 
Haplotrema vancouverense (Lea), 1839; (45-CA-30/18) —Fig. 3-C. 
Reference: 
Pilsbry 1946: 221-225. 
Remarks: 
Lives on land in the general Coast Conifer Forest habitat. 


DiIscussIoN 


Tegula shells were “found in clusters as though they had been dumped 
from a contamer of some sort?” Jegula is utilized as a foodstuff yet; 
along the central California coast, they are collected in pails for boil- 
ing for stews. 

The shell of the limpet Acmaea mitra (Fig. 2-F) had some bryo- 
zoan incrustation in its interior; this would indicate the shell bemg 
again in marine water after the animal had been removed by man 
or other predator. 

Reagen (1917: 17-20) offered a pioneering trilogy for gross chrono- 
logical precontact cultural change in the Ozette-Makah territory. He 
mentions oyster shells bemg an important part of the molluscan re- 
mains in the sites of the general area, especially for what he then 
considered older and oldest periods. Therefore, presence and/or 
absence of oyster shells in 45-CA-30, and related sites, intimate some 
connection with cultural change related to environmental change in 
different portions of the Olympic Peninsula, and perhaps concur- 


Molluscs from archaeological sites 107 


rently. This suggests the possibility that changes of relatively few feet 
in shoreline oscillation, perhaps over a lengthy period of time, could 
bring about enough habitat alteration to seriously distort the food 
gathering economy. 


MarinE Moutuuscs AND BARNACLES 


The 3 bivalve and 6 gastropod molluscs are tidal-zone or close to shore 
(and probably also shallow water) forms. One barnacle, the Balanus, 
would probably be found attached to rocks within tidal limits when it 
lived. If many specimens of the other barnacle, the Coronula, were 
found in the site, their presence would give additional argument for 
a sea mammal hunting phase for the aboriginal beach living location. 


TERRESTRIAL GASTROPOD 


Haplotrema vancouverense is a species distributed from southeastern 
Alaska to northern California and eastwardly in places to the north- 
ern Idaho region. It has been found in aboriginal sites in southwest- 
ern British Columbia along with landsnails of three other genera: 
Allogona, Monadenia, and Vespericola. (An evaluation of landsnail 
remains in British Columbia sites will be the third in the series in- 
cluding the present report.) In 45-CA-30, Haplotrema is reported as 
“found only in sparse layers and always on top of other snails” (77 
litt., S. J. Guinn, 30 May 1962). It is not known if terrestrial snails 
were used as food by prehistoric Indians of the Pacific Northwest; 
they are being currently studied as remnants of the background biotas 
surroundings man’s record in archaeology. Eventually detailed stu- 
dies of nonmarine molluscs in sites (with possible controlled sampling- 
excavation) and including other biological remains similarly con- 
sidered will supplement a body of indications, some most subtle but 
still interlocking, for changes in past environments. 


ACKNOWLEDGMENTS 


Dr. Richard D. Daugherty and Mr. Stanley J. Guinn of the De- 
partment of Sociology and Anthropology of Washington State Uni- 
versity kindly provided the archaeo-zoological material for site 45- 
CA-30. Drafting was by Mrs. Maureen A. Douglas and record-typing 
was by Miss A. C. Templeton of the Department of Zoology of U.B.C.; 
I did the photography. This report represents one of several such 
evaluations supported by Grant G24475 from the Anthropology Pro- 
gram of the National Science Foundation, 1962-1963. 


108 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 2, 1963 
LITERATURE CITED 


BORDEN, CHARLES E. 
1962. Notes and news. Northwest. Amer. Antiquity, 27:609-613. 


CORNWALL, IRA E. 

1955a. The barnacles of British Columbia. British Columbia Prov. Mus., Handbk. 
No. 7; 69 pages, 9 figs., illus. 

1955b. Canadian Pacific fauna: 10. Arthropoda: 10e. Cirripedia. Fisheries Re- 
search Board of Canada; 49 pages, 39 figs. 


DARWIN, CHARLES 

1854. A monograph of the sub-class Cirripedia, with figures of all the species; 
the Balanidae, ... the Verrucidae. ...The Ray Society; [vol. 25;] 684 pages. 
30 pls., index. London. 


MORRIS, PERCY A. 
1952. A field guide to shells of the Pacific Coast and Hawaii. Boston: Houghton 
Mifflin Co.; 220 pages, text illus., 40 pls., glossary, index. 


OLDROYD, IDA SHEPARD 
1927. The marine shells of the West Coast of North America. Vol. II, Pt. 1. Stan- 
ford Univ. Publs., Univ. Ser.: Geol. Sci. Vol. II, Pt. 1; 297 pages, 20 pls., 


index. 


PALMER, KATHERINE VAN WINKLE 

1958. Type specimens of marine Mollusca described by P. P. Carpenter from the 
West Coast (San Diego to British Columbia). Geol. Soc. Amer., Mem. 76; 
376 pages, 35 pls., frntsp., index. 

PILSBRY, HENRY A. 

1921. Barnacles of the San Juan Islands, Washington. Proc. U.S. Nat. Mus., 59 
(2362) :111-115, pl. 20. 

1946. Land Mollusca of North America (north of Mexico). Acad. Nat. Sci. Phila- 
delphia, Mono. 3, Vol. 2, Pt. 1. 

REAGAN, ALBERT B. 


1917. Archaeological notes on western Washington and adjacent British Colum- 
bia. Proc. California Acad. Sci. (4th ser.), 7:1-31, pls. 1-6, maps. 


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DUP cEIN OF THE SOUTHERN CALIFORNIA 
ACADEMY OF SCIENCES 
Vou. 62 JULY-SEPTEMBER, 1963 Part 3 


ATTRACTION OF INSECTS TO EXUDATES OF 
VERBESINA ENCELIOIDES AND IVA AMBROSIAEFOLIA 


E. G. LinsLey anp M. A. Cazier 
University of California, Berkeley 


We have called attention previously to the attraction of insects to exu- 
dations from the stems of Senecio longilobus Benth. (thread-leaf 
groundsel), a species of composite containing alkaloids highly toxic to 
livestock but not yet shown to be significantly so to insects (Linsley and 
Cazier, 1962). Among the insects attracted to the exudate were tene- 
brionid and lagriid beetles; braconid, multillid, pompilid, sphecoid and 
vespoid wasps; and various Diptera, including Sarcophaga. The domi- 
nant species from the standpoint of size and continuity of presence on 
the plant during the period in which it was under observation, was the 
large, black, heavy-jawed longicorn beetle, Stenaspis solitaria (Say). 
One or more males were present almost continuously and usually also 
females as partners in mating pairs. Both sexes chewed on the surface 
of injured areas and lapped up exuding liquids. It was not clear 
whether or not they were responsible for the primary lesion. After 
extensive feeding, the beetles often appeared logy, especially the fe- 
males, and commonly sat on leaves or stems for some time before 
flying away. : 

In the summer of 1962, at a site one mile east of Douglas, Cochise 
County, Arizona, an opportunity was afforded to make a few obser- 
vations on the attraction of insects to exudates from injuries in two 
additional species of composites, Verbesina encelioides and Iva am- 
brosiaefolia. Both plants were growing in and next to a shallow burrow 
excavation on the upper slope leading into drainage dyke and in each 
case the primary lesions were produced by the large scarab beetle, 
Cotinis palliata. In late July a female of the long-horned beetle, Den- 
drobias mandibularis, was observed chewing on an uninjured stem of 
Verbesina but she failed to produce a primary lesion. Attempts to 
artificially produce exuding lesions on the stems of Verbesina by cut- 
ting or scraping were unsuccessful as the tissue dried too rapidly and 
none of the plants with scarab lesions died durmg the period of these 


109 


110 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


observations. The latter was probably because the stems were not 
girdled by the lesions. 

Adults of Cotinis palliata are notorious for their injury to fruits, 
especially figs, peaches and grapes. According to Nichol (1935), in 
the hot climate of southern Arizona, ripe fruit which is attacked will 
be fermenting by the following day, and the beetles return and feed 
ull it is entirely consumed. They also follow fruit injury by Gila 
woodpeckers and other birds. He states that when they attack a whole 
skinned fruit, they utilize the clypeal horn to break the surface, since 
the mouthparts are not suitably developed for tearmg the epidermis. 
They leave a sickening odor where they have been feeding as well as 
a greasy excrement. He found both sexes feeding during all of the day- 
light hours, but mostly between 10 am and dusk. Broken watermelons 
are especially attractive to Cotinis and have been used in orchards as 
traps. During the breeding season the diet is varied with pollen from 
sorghums, maise, grasses, and various herbs and shrubs. Nichol ex- 
perimented with various fermenting baits—those containing sour milk 
or grape or peach juice and water being the most attractive of those 
tested. 


Verbesina encelioides (Cavy.) Benth. & Hook. 


This species, one of the crown-beards, is an erect, many-branched an- 
nual, with numerous flowerheads, canescent stems, and a large tap- 
root. It was the dominant plant for a distance of several hundred yards 
along the excavation, the majority of individuals ranging from 4 to 6 
feet in height. However, injured plants were limited to about a dozen 
specimens in a group of 75-80 examples growing in an area about 
10x24 feet square. Feeding by large scarabs (Cotinis palliata) at the 
base of plants near the ground level, or less commonly on branches in 
the middle of the plant, was noticed in mid-July. The resulting lesions 
produced white exudates for a period of two or three weeks that were 
attractive to a wide range of insects. Nevertheless the attraction was 
selective and by no means representative of the fauna present in the 
immediate area. Aside from Cotinis, the most conspicuous of the insects 
attracted were large, yellow and black long-horned beetles (Dendrobias 
mandibularis ), large blue mud wasps (Chlorion aerarius), large red 
spider wasps (Cryptocheilus severini), and large black ground beetles 
(Calosoma). These species were aggressive and belligerent toward one 
another in varying degrees and thus exhibited varying degrees of 
dominance in different combinations. 

Males of Dendrobias mandibularis were seen feeding on the exudate 


Attraction of insects to plant exudates 111 


as early as July 23, 1962, but sustained observations on their activity 
were not possible until August 16, 1962. At 7:10 am (air temperature: 
79° F), a male was observed feeding at a lesion in nearly horizontal 
stem toward the middle of a plant, in the company of two Cotinis and 
various small flies, wasps and beetles (for hourly counts of the com- 
position of the feeding aggregations, see Table II). He was joined 
shortly by a female which flew in. The male immediately engaged her 
in copulation, holding her alongside the pronotum with the long, broad, 
front tarsi, the middle and hind legs remaining on the substrate. While 
the female continued to feed, the pair was rushed repeatedly by a fe- 
male Chlorion which the male fought off with the antennae and front 
tarsl. 

At 8:16 am the female stopped feeding and broke off copulation, 
flying off after a few moments of preening and cleaning the mouth- 
parts. Thereupon the male returned to feeding, fighting off wasps and 
larger beetles belligerently but largely ignoring the smaller insects. 
The arrival of a third Cotinis immediately precipitated a battle with the 
scarabs butting each other and the cerambycid. The latter was at dis- 
advantage when butted from behind and was knocked off the branch 
several times, only to fly directly back and resume battle. In head-to- 
head combat Dendrobias and Cotinis were more or less stalemated, but 
when the former approached from behind he succeeded in pushing 
each of the scarabs off the branch and keeping them away. 

At 10:14 am, a second (smaller) female Dendrobias flew in and 
started to feed. As previously, the male engaged her in copulation, 
which continued until 11:25 am, when she flew off and attached her- 
self to the underside of a leaf of a Verbesina plant about 25 feet distant. 
Once again, the male became very belligerent toward all of the larger 
insects then present. At 11:52 am (air temperature 93° F) the male 
left the stem and crawled to the underside of a leaf and hung up in 
the shade. The next morning, and each morning during the following 
week the male Dendrobias returned to the exudation site and domi- 
nated it, being joined from time to time by females. One morning, two 
mating pairs were present but when copulation was complete and the 
females had left, the first male succeeded in chasing away the second. 

As has been emphasized previously (Linsley, 1959), the olfactory 
sense is well developed in Cerambycidae. Beeson and Bhatia (1939) 
and others have recorded the rapid response of Hoplocerambyx spini- 
cornis (Newman) to the fresh sap of Shorea robusta, to which adults 
fly over considerable distances. They have been attracted over a meas- 
ured quarter of a mile within five minutes, flymg upwind. The males 


112 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


precede the females in flying to a trap tree, to which as many as 465 
adults have been attracted in a single day. 836 over a period of several 
days. 

Little is known of the food habits of adult Dendrobias mandibularis. 
Schwarz (1904) reported that the species is especially fond of printers’ 
ink and sometimes obliterates the large letters on posters of theatrical 
performances, etc., which are pasted on walls and fences. More details 
on this habit and data on the chemical nature of the ink and glues 
involved would have been of interest in relation to the above observed 
behavior. Chemsak (1958) recorded the attraction of large numbers 
of individuals of both sexes to watermelon rinds in trash barrels (a 
sample contained 32 males and 34 females). Among these were two 
females of the longhorned beetle Eustromula validum (LeConte) all 
that were seen. Watermelon is attractive to other insects, including 
bees (Bohart, 1950) and, as mentioned above, the scarab Coftinis. 
Dendrobias have been collected as they were feeding on the exudate of 
Salix sp. at Holtville, California (Cazier & Ross) and on the sap of 
Baccharis sarothroides i1 Sabmo Canyon, Arizona (Bradt & Cazier ) 
where they were found in association with another Cerambycid. 
Stenapsis verticalis, which was feeding on the same injuries. In late 
June, 1950, a number of D. mandibularis were collected from the 
blossoms of Yucca elata at Don Luis, Arizona (Cazier), but no feeding 
observations were made. They were taken singly and in copulation on 
these flowers. 

At the ground level sites, the Cotinis and Calosoma were dominant. 
usually ignoring attacks from Chlorion, which was dominant in the 
absence of the larger beetles. A maximum of five Cotinis were found 
feeding at a single lesion at the same time on Verbesina and six Calo- 
soma were feeding on or were in the immediate vicinity of a lesion 
on Iva. Although the smaller insects were often dislodged in battles 
between the larger forms, they flew or crawled back immediately and 
apparently would not leave until satiated. At the time when the sample 
reported in Table I was collected, most of the ground level lesions were 
drying up, and the residual species composition was not fully represen- 
tative of the total group attracted. 


Attraction of insects to plant exudates 113 


TABLE I 
Numbers and kinds of insects feeding at hourly intervals 
from exudations from base of Cotinis-injured Verbesina encelioides 
during morning of August 16, 1962* 


Insects 853 07 30 LO SOT 1-30 
Sarcophaga prohibita (Diptera, Sarcophagidae) 3 3 
Calosoma peregrinator (Coleoptera, Carabidae) 1 
Aeolus mellillus (Coleoptera, Elateridae) 
Statira sp. nr. pluripunctata (Coleoptera, Lagriidae) 
Dasymutilla gloriosa (Hymenoptera, Mutillidae) 
Dasymutilla klugit (Hymenoptera, Mutillidae) 
Notogramma purpurata (Diptera, Otitidae) 
Chlorion aerarius (Hymenoptera, Sphecidae ) 
Euxesta sp. (Diptera, Tephritidae) 
Polistes major castaneicolor (Hymenoptera, Vespidae) 
Priocnemioides t. texanus (Hymenoptera, Pompilidae ) 
Dasymutilla chrysocoma (Hymenoptera, Mutillidae) 
Polistes fuscatus centralis (Hymenoptera, Vespidae ) 2 
Largus cinctus (Hemiptera, Pyrrhocoridae) 1 1 
Asterocampa leilia (Lepidoptera, Nymphalidae) 1 


me tot 


PrRrPrP bb a 
bo 


Perr bb to lb 
ee 


*Among the insects observed at the exudations of this plant during the week 
prior to August 17 but not represented in the above samples were: Carpophilus 
lugubris (Coleoptera, Nitidulidae), Cotinis palliata (Coleoptera, Scarabaeidae), 
Euphoria fascifera (Coleoptera, Scarabaeidae), Megacyllene antennata (Coleop- 
tera, Cerambycidae), Dendrobias mandibularis (Coleoptera, Cerambycidae), 
Opius sanguineus (Hymenoptera, Braconidae), Zonosemata vittigera (Diptera, 
Tephritidae) and several undetermined small flies (Diptera). 


TABLE II 
Numbers and kinds of insects feeding at hourly intervals from exudations 
from Cotinis-injured stem near middle of plant of Verbesina encelioides 
during morning of August 16, 1962 (7:20 a.m. air temperature: 80° F’) 

Insects 7:20 8:20 9:20 10:20 11:20 
Algarobius prosopis (Coleoptera, Bruchidae ) 
Centrinaspis hospes (Coleoptera, Curculionidae) 
Cotinis palliata (Coleoptera, Scarabaeidae) 
Dendrobias mandibularis (Coleoptera, Cerambycidae ) 
Undet. small flies (Diptera) 
Priocnemioides t. texanus (Hymenoptera, Pompilidae) 
Chlorion aerarius (Hymenoptera, Sphecidae) 
Opius sanguineus (Hymenoptera, Braconidae ) 
Sarcophaga prohibita (Diptera, Sarcophagidae ) 1 
Asterocampa leilia (Lepidoptera, Nymphalidae) 
Cryptocheilus severini (Hymenoptera, Pompilidae ) 
Rygchium guerrero (Hymenoptera, Vespidae ) 
Zonosemata vittigera (Diptera, Tephritidae ) 
Dyscrasis hendeli (Diptera, Otitidae) 1 
Euptoieta claudia (Lepidoptera, Nymphalidae) 1 
Notogramma purpurata (Diptera, Otitidae) 1 


PBPrPrPbWbWNAOD 
— Aron 
bo 4 wor Dd bv 


PS eS LS) 9) 
ps ess es Ss 


114. Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


The following insects were found feeding on exudates of Verbesina 
between July 23, 1962 and August 21, 1962: 


Order Hempitera 
Family Pyrrocoridae 
Largus cinctus (Herrick-Schaeffer)— widely distributed in southwestern 
United States and northern Mexico. Adults reported to feed on berries and 
fruits. 


Order Lepidoptera 
Family Nymphalidae 
Asterocampa leilia (Edwards)—Known from Texas to southern Arizona and 
Mexico. 
Euptoieta claudia (Cramer )— occurring in southern United States, Mexico 
and Central America. 


Order Coleoptera 


Family Carabidae 
Calosoma (Camedula) peregrinator Guérin-Méneville—Ranging from New 
Mexico to southern California. Adults known to feed on cutworms and 
other caterpillars. 


Family Elateridae 
Aeolus mellillus (Say)—Southwestern United States. Adults nocturnally 
active and attracted by light. 
Family Lagriidae 
Statira sp. nr. pluripuncta (Horn)—Representative of a large genus of near- 
ly 450 described species in the American tropical and subtropical regions, 
of which six occur in southwestern United States, three in eastern and cen- 
tral United States. 


Family Scarabaeidae 
Cotinis palliata (Gory)—A common species of Mexico and the southern 
United States; injurious to fruits, etc. (see above). 
Euphoria fascifera LeConte—A rarely encountered species known from Baja 
California, northwestern Mexico and southern Arizona. 


Family Cerambycidae 
Dendrobias mandibularis mandibularis Serville—Ranging from Baja Califor- 
nia and the mainland of Mexico to southern Texas, New Mexico and Ari- 
zona. 
Megacyllene antennata (White)—Baja California and northern Mexico to 
Texas, New Mexico, Arizona and southern California. 


Family Bruchidae 
Algarobius prosopis (eConte)—Occurring from Texas to southern Califor- 
nia, northern Mexico and Baja California. 


Family Curculionidae 
Centrinaspis hospes (Casey )— Our material is all from southern Arizona, 
Sonora and Chihuahua. 


Attraction of insects to plant exudates 115 


Order Diptera 
Family Sarcophagidae 
Sarcophaga prohibita Aldrich—Described from Kansas and widely distributed 
in southwestern United States. 
Protodexia hunteri (Hough)—A western North American species repeatedly 
reared from grasshoppers (Aldrich, 1916). 
Family Otitidae 
Notogramma purpurpata Cole—A rare species described from the Gulf of 
California. 
Dycras:s kendeli Aldrich—Known to us only from a few examples froim 
Texas. 
Family Tephritidae 
Euxesta sp. 
Zonosemata vittigera (Coquillett)—A fruit fly living in the fruit of Solanum 
elaeagnifolium (Cazier, 1962). 


Order Hymenoptera 
Family Braconidae 
Opius sanguineus (Ashmead )—A species widely distributed in eastern North 
America. Locally a parasite of the larvae of Zonosemata vittigera (above). 
a Solanum fruit fly (Cazier, 1962). 
Family Multillidae 
Dasymutilla chrysocoma Mickel—Our material is all from Arizona. 
Dasymutilla gloriosa (Saussure)—A species of the desert areas of southwest- 
ern United States and northern Mexico. 
Dasymutilla klugti (Gray )—Also a species primarily limited to southwest- 
ern United States and northern Mexico. 
Family Vespidae 
Polistes fuscatus centralis Hayward—Occurring in the southwestern United 
States, northern Mexico, and Baja California. 
Polistes major castaneicolor Bequaert—Our material is all from southera 
Arizona and New Mexico and northern Mexico. 
Rygchium guerrero (Saussure )—Western Texas to southern Arizona, north- 
western Mexico and Baja California. 
Family Pompilidae 
Priocnemioides texanus texanus (Cresson )—Our material is from southwest- 
ern United States and adjacent areas of northern Mexico. 
Cryptocheilus severini Banks—Widely distributed in lower regions of cen- 
tral and southern United States and northern Mexico. 
Family Sphecidae 
Chlorion aerarius Patton—A widely distributed North American wasp. 


Verbesina encelioides var. exauriculata occurs from Kansas to south- 
western United States and northern Mexico. According to Blake 
(1951) it is said to have been used by the Indians and white pioneers 
for boils and skin diseases. The Hopi are also reported to bathe in water 
in which the plant has been soaked to relieve the pain of spider bite. 


116 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


Heal, Rogers, Wallace and Starnes (1950), in tests with various in- 
sects, found that aqueous extracts of the flowerheads and of the stems, 
leaves, and flowers were very toxic to American cockroaches when in- 
jected into the blood stream, but German cockroaches and milkweed 
bugs were unaffected after immersion in the extract. Alcohol, petro- 
leum ether, and chloroform extracts of the whole plant were toxic to 
black carpet beetle larvae but not to German cockroaches, milkweed 
bugs, confused flour beetles, and larvae of the webbing clothes moth 
and Aedes mosquitoes. The related V. virginica Linnaeus, in the form 
of powdered stems and leaves did not prove toxic to southern army- 
worms, mealworms, and southern beet webworms (Bottger and Jacob- 
son, 1940) nor larvae of the European cornborer (Jacobson, 1953). 
However, the last author found combined petroleum ether, ethyl 
ether, chloroform, and alcohol extractives toxic to codling moth larvae 
but not houseflies. No evidence of toxicity was observed among the in- 
sects feeding on the exudates from the Verbesina lesions. However, 
their behavior suggested an overwhelming response to the attractive- 
ness of the exudate. 


Iva ambrosiaefolia Gray 


This plant, one of the marsh elders, of which about fifteen species are 
known—all North American, is a shrub-like herb with inconspicuous 
green flowers. In the site under study, individuals were widely scat- 
tered along the wash among the dominant Verbesina encelioides. Only 
a few exhibited Cotinis injuries, but these were overrun with insects 
of many kinds. The most prominent of these was a plant about 4 ft. 
high and 5 ft. in diameter. 

The following insects were observed on a plant exuding liquids from 
Cotinis mjuries: 


Order Hemiptera 
Family Pyrrocoridae 
Largus cinctus (Herrick-Schaeffer) (See above) 
Order Neuroptera 
Family Mantispidae 
Climaciella occidentis (Banks) 
Order Coleoptera 


Family Carabidae 

Calosoma (Camedula) peregrinator Guérin-Méneville (See above) 
Family Lagriidae 

Statira sp. nr. pluripuncta (Horn) (See above) 


Family Nitidulidae 


Attraction of insects to plant exudates ANG 


Carpophilus lugubris (Say)—The widespread and notorious “dusky sap 
beetle” which has long been known to be attracted to fermenting and de- 
caying plant materials. On sweet corn plants, sites injured by other insects 
are very attractive to adults and infestations increase when damage by 
other insects is present (Harrison, 1962). 

Family Scarabaeidae 
Cotinis palliata (Gory) (See above) 
Family Cerambycidae 
Megacyllene antennata (White) (See above) 


Order Diptera 
Family Asilidae 
Beameromyia macula Martin—Our material is all from southeastern Ari- 
zona. 
Family Sarcophagidae 
Sarcophaga prohibita Aldrich (See above) 
Protodexia hunteri (Hough) (See above) 
Family Lauxaniidae 
Camptoprosopella sp. 
Family Otitidae 
Notogramma purpurata Cole (See above) 
Family Tephritidae 
Euxesta sp. 
Zonosemata vittigera (Coquillett) (See above) 


Order Hymenoptera 

Family Mutillidae 
Dasymutilla chrysocoma Mickel (See above) 
Dasymutilla gloriosa (Saussure) (See above) 
Dasymutilla nogalensis Mickel—Our material is all from southern Arizona 

and northern Mexico. 

Family Vespidae ; 
Polistes fuscatus centralis Hayward (See above) 
Polistes major castaneicolor Bequaert (See above) 

Family Pompilidae 
Cryptocheilus severini Banks (See above) 


Hemipepsis ustulata ustulata Dalbohm—Central and southern United States 
and Mexico 


Priocnemioides texanus texanus (Cresson) (See above) 
Family Sphecidae 
Chlorion aerarius Patton (See above) 


118 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


TABLE III 
Numbers and kinds of msects found at hourly intervals 
at exudations at base of Cotinis-mjured Iva ambrosiaefolia 
during morning of August 19, 1962 
Insects 731) “8:1 Poe lO15. ele 


Calosoma peregrinator (Coleoptera, Carabidae) 6 i 7 + s 
Cotinis palliata (Coleoptera, Scarabaeidae) 4 4 5 5 5 
Misc. small flies (Diptera) 4 6 9 7 10 
Statira sp. nr. pluripunctata (Coleoptera, Lagriidae) 3 3 1 1 1 
Hemipepsis u. ustulata (Hymenoptera, Pompilidae) 3 2 1 3 1 
Chlorion aerarius (Hymenoptera. Sphecidae) 3 4 4 2 3 
Carpophilus lugubris (Coleoptera, Nitidulidae) 2 2 

Dasymutilla gloriosa (Hymenoptera, Mutillidae) 2) 1 1 
Sarcophaga prohibita (Diptera, Sarcophagidae) 2 1 2 2 1 
Cryptocheilus severini (Hymenoptera, Pompilidae) 1 2, 1 1 1 
Polistes fuscatus centralis (Hymenoptera, Vespidae) 1 3 2 2 
Dasymutilla nogalensis (Hymenoptera, Mutillidae) 1 

Beameromyia macula (Diptera, Asilidae) * 1 1 1 1 2 
Climaciella occidentis (Neuroptera, Mantispidae) * 1 

Megacyllene antennata (Coleoptera, Cerambycidae) 1 1 1 1 
Priocnemioides t. texanus (Hymenoptera, Pompilidae) i 1 

Polistes major castaneicolor (Hymenoptera, Vespidae) 1 1 1 
Dasymutilla chrysocoma (Hymenoptera, Mutillidae) 1 1 

Zonosemata vittigera (Diptera, Tephritidae) 1 2 1 
Camptoprosopella sp. (Diptera, Lauxaniidae) 1 1 

Largus cinctus (Hemiptera, Pyrrhocoridae) 1 1 1 1 
Euxesta sp. (Diptera, Tephritidae) 1 


*Neither of these species was observed to feed on the exudate, but their presence on t] 
plant suggests the possibility that they may have been attracted by the secretion. 


Iva ambrosiaefolia occurs from western Texas to southern Arizona 
and Northern Mexico, blooming from May to October. The stems are 
pubescent and the plant aromatic, but we have been unable to find 
reference to its toxic properties, if any. The related /. xanthifolia Nutt.. 
according to Blake (1951), induces dermatitis in some persons after 
contact, and the pollen is a cause of hay fever. Heal, Rogers, Wallace 
and Starnes (1950) applied extracts of the upper parts, leaves, and 
flowers of this last species and of J. azvillaris Pursh. and I. frutescens 
Linnaeus to German and American roaches and to milkweed bugs. 
without toxic effect. 


PrepaAtors Upon ATTRACTED INSECTS 


During the course of these observations the remains of D. mandibularis 
and C. palliata were found in the early morning beneath the plants on 
which they had been seen feeding during the preceding day. Both 


species were observed at night on or near the exuding lesions and 


Attraction of insects to plant exudates 119 


when these were near the ground the beetles evidently fell prey to some 
nocturnal species of rodent. What appeared to be mouse droppings 
were intermixed with the hard remains of the beetles. C. palliata was 
also found “sleeping” high up on Conyza coulteria, Prosopis juliflora, 
Iva ambrosiaefolia and Verbesina encelioides. D. mandibularis was 
found at night on the leaves and flowerheads of Verbesina encelioides. 
No remains of the third large beetle (Calosoma) were found although 
these beetles remained overnight near the lesions. Their repugnant 
odor or ability to run rapidly may account for their apparent immu- 
nity to rodent attacks. 


SUMMARY 


(1) The attraction of various insects to exudates from living plants 
of Verbesina encelioides and Iva ambrosiaefolia was observed during 
July and August in southeastern Arizona. 

(2) The primary lesions from which the exudates flowed were made 
by the scarab beetle Cotinis palliata. 

(3) The exudates were selectively attractive and the assembled in- 
sects were not a representative cross-section of the immediate local 
fauna. About 30 species were observed at Verbesina, about 25 at Iva. 

(4) The exudation sites were usually dominated by large insects 
which were antagonistic to one another. At various times and at vari- 
ous sites the dominant insect was the longhorned beetle Dendrobias 
mandibularis, or the scarab beetle Cotinis palliata, or the ground beetle 
Calosoma peregrinator, or the sphecid wasp Chlorion aerarius. Smaller 
insects were tolerated by these, but were frequently dislodged in 
battles between them. 

(5) Insects remaining overnight at lesions near the ground were 
subject to predation—presumably by rodents. 


ACKNOWLEDGMENTS 


The authors are indebted to Marjorie Statham, American Museum of 
Natural History, for the photographs which accompany this article. 
Insects were identified by Frank Cole, University of California, Ber- 
keley and H. J. Reinhard, Agricultural and Mechanical College of 
Texas (Diptera), P D. Hurd, University of California, Berkeley (Acu- 
leate Hymenoptera), Paul J. Spangler, T: J. Spilman, and R. E. 
Warner, United States National Museum (Coleoptera), and R. L. 
Usinger, University of California, Berkeley (Hemiptera). Plants were 
identified by Margaret S. Bergseng, Herbarium, University of Cali- 
fornia, Berkeley. 


120 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


Figure 1 (upper). Frothy exudate from base of Cotinis-injured plant of Verbesina 
encelioides, August 16, 1962. 


Figure 2 (lower), Female of Dasymutilla gloriosa feeding on exudate. 


Attraction of insects to plant exudates ADA 


Figure 3 (upper). Butterfly (Asterocampa leilia) feeding on exudations from base 
of Cotinis-injured plant of Verbesina encelioides. Background insects include the 
bug, Largus cinctus, and miltogrammine flies. 


Figure 4 (lower). Calosoma peregrinator feeding on exudations. 


122 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


ee ~ 


Figure 5 (upper). Mating pair of Dendrobias mandibularis, the female feeding on 
exudates from Cotinis-injured mid-area of Verbesina encelioides, August 16, 1962. 
Background insects include Sarcophaga prohibita (large fly), Centrinaspis hospes 
(weevil), Algarobius prosopis (bruchid), and Opius sanguineus (braconid wasp). 


Figure 6 (lower). Mating pair of Dendrobias mandibularis with newly arrived 
female on opposite side of stem feeding on exudates. Background insects include: 


Sarcophaga prohibita (large fly), Protodexia hunteri (small fly), and Centrinaspis 
hospes (weevil). 


Attraction of insects to plant exudates 1 


to 
Oo 


Figure 7 (upper). Female of Megacyllene antennata, newly arrived on Cotinis- 
injured Verbesina plant. 


Figure 8 (lower). Cotinis palliata (scarab) and Centrinaspis hospes (weevil) 
feeding on exudates of Verbesina encelioides. 


ee 


Figure 9 (upper). Chlorion aerarius feeding before attempting to displace mat- 
ing pair of Dendrobias mandibularis. Background insects include Sarcophaga 
prohibita (large fly), Protodexia hunteri (small fly), Centrinaspis hospes 
(weevil), and Euzxesta (tephritid fly). 


Figure 10 (lower). Chlorion aerarius and Dendrobias mandibularis (male) 
feeding after the former had attacked mating pair (above) and chased off the 
female. Male longicorn, although feeding, adopted aggressive attitude toward 
Chlorion and ultimately chased it away. 


to 
Ov 


Attraction of insects to plant exudates 4 
| 


Figure 11 (upper). Insects feeding at exudations at base of Cotinis-injured plant 
of Iva ambrosiaefolia, August 19, 1962. A butterfly (Asterocampa leilia), mud 
dauber wasp (Chlorion aerarius), and numerous flies, including Sarcophaga 
prohibita and Protodexia hunteri, are represented. 


Figure 12 (lower). Cotinis palliata feeding at base of /va ambrostiaefolia. Back- 
ground insects include the fly Sarcophaga prohibita. 


126 «Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


ee eee 


= nn 


Figure 13 (upper). Insects feeding at exudates from base of Cotinis-injured Iva 


ambrosiaefolia. Included are: the flies: Sarcophaga prohibita, Protodexia hunteri, 
and Notogramma purpurata, and the wasps Cryptocheilus severini and Polistes 
fuscatus centralis. i 


Figure 14 (lower), Calosoma peregrinator and Chlorion aerarius feeding at 
exudates. 


Attraction of insects to plant exudates 127 


Figure 15 (above ), Cotinis palliata feeding on stem of Iva ambrosiaefolia, August 
19, 1962. 


Figure 16 (below). Elytra of Cotinis palliata and rodent droppings beneath 
Cotinis-injured plant of Iva ambrosiaefolia, suggesting that the scarabs are subject 
to nocturnal predation by rodents. 


128 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


Figures 17 and 18. Polistes fuscatus centralis (above) and Climaciella occidentis 
(below), wasp and wasp-like neuropteron, on foliage of Iva ambrosiaefolia, 
August 19, 1962. 


Attraction of insects to plant exudates 129 


LITERATURE CITED 


ALDRICH, J. M. 

1916. Sarcophaga and allies in North America. Say Found., Ent. Soc. Amer., 
1:1-301, 16 pls. 

BEESON, C. FE C. ann B. M. BHATIA 


1939. On the biology of the Cerambycidae of India, Burma and Ceylon. Indian 
Forest Records (n.s.), Ent., 5:1-235. 


BLAKE, S. E 
1951. Compositae, in: Kearny, T. H. and R. H. Peebles, Arizona Flora. Univ. 
Calif. Press, pp. 829-971. 


BOHART, G. E. 

1950. Observations on the mating habits of halictid bees. Pan-Pac. Ent., 26:34-35. 

BOTTGER, G. T. ann M. JACOBSON 

1950. Preliminary tests of plant materials as insecticides. Bur. Ent. Plant Quar., 
E-796: 1-35. 

CAZIER, M. A. 


1962. Notes on the bionomics of Zonosemata vittigera (Coquillett), a fruit fly on 
Solanum (Diptera: Tephritidae). Pan-Pacific Ent., 38:181-186. 

CHEMSAK, J. A. 

1958. An attractant for two species of Cerambycidae. Pan-Pacific Ent., 34:42. 

HARRISON, E P. 


1962. Infestation of sweet corn by the dusky sap beetle, Carpophilus lugubris. ]. 
Econ. Ent., 55:922-925. 


HEAL, R. E., E. F ROGERS, R. T. WALLACE anp O. STARNES 
1950. A survey of plants for insecticidal activity. Lloydia, 13:89-162. 


JACOBSON, M. 

1953. Insecticidal plants. U.S. Dept. Agr., Bur. Ent. Plant Quar., Div. Insecticide 
Investigations, Spec. Rep. 26: 1-132. 

LINSLEY, E. G. i 

1959. Ecology of Cerambycidae. Ann. Rev. Ent., 4:99-138. 


LINSLEY, E. G. anp M. A. CAZIER 

1962. A note on the attraction of Stenaspis solitaria (Say) and other insects to 
Senecio longilobus, a range plant highly toxic to livestock. Canadian Ent., 
94:745-748, figs. 1-2. 


NICHOL, A. A. 

1935. A study of the fig beetle, Cotinis texana Casey. Univ. Ariz. Agr. Exp. Sta., 
Tech., Bull., 55:157-198. 

SCHWARZ, E. A. 

1904. Food habits of longicorn beetles. Proc. Ent. Soc. Washington, 6:21-22. 


NOTES ON THE BARNACLE LEPAS FASCICULARIS 
FOUND ATTACHED TO THE JELLYFISH VELELLA 


Several years ago while the Allan Hancock Foundation vessel Velero 
IV was occupying station 2792-54, about 7.2 miles WNW of Palos 
Verdes Point, San Pedro Channel, California, the surface of the sea 
was observed to be littered with floating barnacles. Two quarts of these, 
obtained by dip net, were preserved by the writer and brought back 
to the Foundation laboratory, where they were identified as Lepas 
fascicularis Ellis and Solander 1786, a well-known pelagic species. 
Further examination revealed that the barnacle float in most cases was 
attached to the underside of the float of Velella a pelagic jellyfish fre- 
quently encountered. 

A review of the early literature reveals that this phenomenon was 
not unknown, though there have been no records since 1876. Darwin 
(A monograph on the fossil Lepadidae, 1851) discussed the cement 
glands and ball floats of Lepas fascicularis at great length and briefly 
mentioned the occasional attachment to Velella. Willemoes-Suhm 
(Philos. Trans. Roy. Soc., 167:131-154, 1876) in his work on the de- 
velopment of Lepas fascicularis, collected on the Challenger expedi- 
tion between Japan and the Sandwich Islands, also mentioned that the 
cypris-like larvae attach to the dead floats of Velella. Though a few 
records of this peculiar phenomenon have been recorded in the past 
it is felt that a photograph which clearly shows the morphology and 
size of L. fascicularis and its position in relation to Velella will be of 
value since other published illustrations are lacking. 

Frequently three or more barnacles have their peduncles imbedded 
in one common float. Of the twenty-one floats collected, with their 
numerously attached barnacles, sixteen definitely contained Velella, 
two more appeared to have Velella, and the remaining three were 
small single barnacle-float combinations which contained no visible 
traces of Velella. In attaching the cirriped larvae fasten to the under 
side of the Velella float. In the process of development they secrete a 
cement substance which forms a yellowish-tan vesicular ball ranging 
up to one-and-one-half inches in diameter. In most cases these balls 
were below the Véelella float, leaving the chitinous concentric air 
chambers and erect sail exposed. In others, however, the cement ball 
passed over the basal portion of the float covering it along with the 
sail. The accompanying photograph (Fig. 1) shows five Lepas fasct- 
cularis attached to the under side of Velella (in the central mass). Of 
twelve additional barnacle ‘‘colonies” that were later found washed 


130 


Barnacles attached to jellyfish 131 


ashore at Pismo Beach, California, all contained small Velella im- 
bedded in their floats. 

Lepas fascicularis, however, is a truly pelagic animal which is not 
dependent on foreign floating material for its existence. When float- 
ing objects such as Velella are not present for attachment the barnacle 
secretes a ball of frothy material from its cement glands to keep it 
afloat. 

The size of this ball is increased as the barnacle grows. In spite of 
its float L. fascicularis is like other barnacles in that it has no power of 
locomotion in the adult stage, and therefore is a victim of oceanic cur- 
rents. The barnacle feeds on microscopic plankton and may in turn 
serve as food for larger marine animals. 

The capitulum size of the L. fascicularis collected ranged from 24 
mm. to 42 mm. in length, and from 8 mm. to 30 mm, in width. An- 
other pelagic barnacle of smaller size, Lepas anatifera Linn., was also 
attached to the capitulum and floats of L. fascicularis.—Jens W. Knud- 
sen, Dept. of Biology, Pacific Lutheran University, Tacoma 44, Wash- 
ington. 


ae ae oe a oe wT i) a hifi ca ry elpa Nl NE 


Figure 1. Five Lepas fascicularis are shown with their peduncles imbedded in a 
common float that is attached to the underside of the jellyfish Velella, Velella can 
be seen in the central mass characterized by the concentric rings of its air cham- 
bers and by the diagonal sail that extends from the lower left to the upper right- 
hand corner. Several specimens of L. anatifera are attached to L. fascicularis. 
Allan Hancock Foundation photograph. 


COMPARISON OF BOCCARDIA COLUMBIANA BERKELEY 
AND BOCCARDIA PROBOSCIDEA HARTMAN 
(ANNELIDA, POLYCHAETA) 


KerruH H. Woopwick 
Fresno State College! 


INTRODUCTION 


Boccardia columbiana and B. proboscidea are morphologically separa- 
ble on only one recognizable characteristic; the first segment has a 
conspicuous notopodial setal fascicle in the first species and a greatly 
reduced one in the second species. Boccardia columbiana Berkeley 
(1927) first described from Vancouver Island, British Columbia, 
occurs in shaly rock where it drills galleries. Boccardia proboscidea 
Hartman (1940) first described from central and southern California 
occurs in minute burrows penetrating shale and limestone reefs. The 
descriptions of the two species are nearly identical morphologically 
and ecologically, differing mainly in geographical location, the first in 
latitudes 49° to 50° N, the second in 34° to 39° N. This geographical 
isolation has been eliminated, however, because B. proboscidea has 
been noted northward to Oregon (Hartman, 1944; Hartman and 
Reish, 1950) and Vancouver Island (Berkeley and Berkeley, 1950, 
1952). Hartman (1940: 385) had suggested for Boccardia probos- 
cidea, ““. . . its range may extend north to Puget Sound, Wash” 

Since 1927, B. columbiana had been reported only from the original 
locality; it is herein recorded south to central California. Its distribu- 
tion overlaps that of B. proboscidea and their ranges are nearly iden- 
tical. It is significant also that these species occur together in the shale- 
limestone and in non-shaly habitats. 


PURPOSE 


Boccardia columbiana and B. proboscidea are similar in range, habi- 
tat, and morphology. It is the purpose of this paper to review similar- 
ities and emphasize the few differences in their morphology, range, 
and habitat. As an aid to identification, a key (see below) distin- 
guishes all Boccardia species known from the Pacific coast of the 
United States and Canada. 


1Rresno, California 


Comparison of two species of Boccardia 133 


ACKNOWLEDGMENTS 


It is a pleasure to acknowledge the assistance of graduate students 
Walter R. Hopkins and William Delton Shirley. This study was aided 
by a grant from the National Science Foundation (NSF G-17990). 


MorPHOLOGY 


Boccardia Carazzi, 1895, is a genus of Spionidae, Polychaeta, Anne- 
lida. It belongs to the polydorid spionids in which a modified fifth seg- 
ment contains enlarged and specialized setae. A pair of palpi used in 
feeding is located anteriorly and branchiae begin on segments ante- 
rior to the modified fifth segment. Bidentate hooded hooks appear first 
in the neuropodium of segment 7. 

Boccardia columbiana and B. proboscidea are similar in that they 
have a rounded prostomium; this with its caruncle extends back to the 
posterior margin of segment 3 (Figs. 1 and 2). Both species may have 
pigmentation laterally along the prostomium and at the edges of the 
palpal grooves. Branchiae are present on segments 2, 3, 4, —, 6, and 7 
and posteriorly but the last few segments lack them. Hooded hooks 
begin on segment 7. Segment 5 is comparable in being well developed, 
larger than preceding and succeeding segments and modified. It lacks 
branchiae, anterior dorsal setae, and setal lobes, but has a double row 
of specialized setae (Figs. 3 and 4). They are of two kinds, a heavy 
falcate spine alternates with a bushy-topped spine (shown in original 
descriptions). Berkeley (1927, Pl. I, Fig. 6) showed an additional kind 
not found in any of the specimens examined by me. Its appearance 
suggests super-imposition of the two kinds of setae. No differences 
were found between the specialized setae of the respective species. 

Late larval forms, early adults, and mature specimens have com- 
parable lateral pigment spots on segments 7 and 8. These spots are 
located between notopodia and neuropodia; they are pronounced in 
younger forms and still visible on mature adults. 

The two species are different in that adult Boccardia columbiana 
specimens have an average length of 12.0 mm. and a maximum of 
15.0 mm.; adult B. proboscidea are more than twice as large, they 
measure 30.0 and 35.0 mm. In B. columbiana the first notopodium is 
well developed with many fine capillary setae spread out in a fan- 
shaped arrangement; their distal ends extend forward beyond the an- 
terior limit of the prostomium and peristomium (see Fig. 1). The setae 
of segment 1 are best seen under reflected light and after the palpi 
have been removed from the specimen. In B. proboscidea the notopo- 


134 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


2 ‘ 


Figure 1. Boccardia columbiana, anterior end (x70) 


Figure 2. Boccardia proboscidea, anterior end (x50) 


dium is weakly-developed, it has only a few short setae and a small 
post-setal lobe; its neuropodium is also poorly developed (see Fig. 2). 

Specimens of both species were taken from piling material at Cayu- 
cos, central California; they measured 6.0 to 8.0 mm. long. Boccardia 
proboscidea was more heavily pigmented in the region of the pros- 
tomium and also on the dorsal surface of segments 2 and 3. Boccardia 
columbiana showed only one of 11 specimens with a heavily pig- 
mented prostomium and none had dorsal pigment on segments 2 and 
3. Generally, the largest B. proboscidea from the piling material had 
the greatest amount of pigment; however, specimens (15.0 to 17.0 
mm.) from the back bay marina mud in Morro Bay showed very little 
pigmentation even along the prostomium. The forms thus are similar 
im general pigmentation but the amount and distribution may vary 
between and within the species according to age, habitat, and possibly 
method of preservation. It has not been possible to ascertain a definite 
pattern of pigmentation for all members of either form. 


3 
Figure 3. Boccardia proboscidea, photomicrograph of modified setae of 
Segment 5 (x500) 


Figure 4. Boccardia proboscidea, modified setae of Segment 5 (x500) 


Boccardia columbiana is anteriorly attenuated with the peristomium 
tapered towards the tip of the prostomium. Boccardia proboscidea has 
a squared-off appearance with a broader, blunter peristomium. The 
appearance of the anterior end varies, however, and is more useful in 
comparing mature adult forms than younger stages. 


GEOGRAPHICAL AND ECOLOGICAL DISTRIBUTION 


Boccardia columbiana has been reported previously from only British 
Columbia; its range is here extended to central California. Boccardia 
columbiana has been found at Monterey, Cambria, Cayucos, and 
Santa Barbara, all in California. Berkeley (1927) reported it usually 
boring in shaly rock. It is now reported in a number of habitats in- 
cluding sandy material in and on wood pilings, sand between barnacle 
tests on pilings, sand tubes in an algal holdfast from mid-tide (worms 
had penetrated and eroded the algae in many places), sponge from 
the underside of rock in low tide, between closely applied branches of 
the holdfast of a kelp, Postelsia palmaeformis Ruprecht, surface of 
rock in close spatial relationship with Phragmatopoma californica 


136 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


(Fewkes), coralline algae (Lithophyllum sp.), and galleries bored in 
shells of several different kinds of gastropeds. 

Boccardia columbiana was found in Tegula funebralis (Adams) 
shells inhabited by Pagurus samuelis (Stimpson) and Purpura foliata 
Martyn and Olivella biplicata (Sowerby) inhabited by Pagurus gran- 
osimanus (Stimpson). It was also found in Acanthina spirata (Blain- 
ville) and O. biplicata mhabited by PR samuelis. In one specimen of 
A. spirata from Santa Barbara it was associated with two other poly- 
dorids, B. proboscidea and Polydora ciliata (Johnston) ; in an O. bipli- 
cata also from Santa Barbara it was associated with Polydora com- 
mensalis Andrews, P. ciliata, and Polydora limicola Annenkova. Boc- 
cardia columbiana also occurred in borings in shells of living Diodora 
aspera (Eschsholtz) and with B. proboscidea in Jaton festivus Hinds. 
It was further found that B. columbiana was present in T: funebralis 
shells inhabited by the gastropod itself, in empty shells, and in shells 
containing hermit crabs. 

Boccardia proboscidea has been described from California, Oregon, 
and British Columbia. Its distribution is extended here only to other 
areas of California including Santa Barbara, Goleta, Cayucos, Fort 
Bragg, and Humboldt Bay. It occurred in sandstone reefs with pholids 
and sipunculids at Santa Barbara and in similar material at Goleta in 
the warm water of a high tide pool. At Fort Bragg it was found in 
crevices in graywacke sandstone as a co-habitant of narrow ledge tide 
pools with a copepod, Jigriopus californicus (Baker). 

Boccardia proboscidea has also been found in soft sandy mud at Bal- 
lona Creek near Venice, Los Angeles County, and more recently from 
mud in the boat slips at Morro Bay and from mud flats of Humboldt 
Bay. It also was found in mucous tubes in sand at the base and among 
the branches of upright branching coralline algae, on the surface of 
intertidal rocks with Phragmatopoma californica, in holdfasts of Mac- 
rocystis integrifolia Bory and Phyllospadix scouleri Hooker, on the 
sandy bottom of inner Santa Barbara harbor in areas of considerable 
pollution, and at Cayucos and Santa Barbara from piling material 
which included Balanus spp. and Mytilus spp. as the dominant ani- 
mals. At Goleta it was found at the base of large Mytilus californianus 
Conrad growth on the outer side of a rocky intertidal reef. Many speci- 
mens were found in pink coralline algae (Lithophyllum sp.) which 
encrusts intertidal rocks at Cayucos. Boccardia proboscidea erodes the 
algae in forming its tube or nestles in pockets between lamellae of the 
corallme where it gathers sand and silt. It may gather with other poly- 
dorids to form pockets of worms. Boccardia columbiana and B. tricuspa 


Comparison of two species of Boccardia 137 


Hartman are associates here; the former has the same habits as Boc- 
cardia proboscidea but the latter erodes and drills the coralline, pro- 
ducing cleaner burrows with no concentrations of associated sand and 
silt. 

Boccardia proboscidea has also been found in gastropod shells in- 
habited by hermit crabs. It occurs with Pagurus samuelis in Tegula 
funebralis and T: brunnea shells at Cayucos, Jaton festivus and Acan- 
thina spirata shells at Santa Barbara, and with Pagurus granosimanus 
in Olivella biplicata shells at Cayucos. Present with this species in 
Tegula brunnea was Boccardia columbiana and Polydora ciliata. In 
some cases B. proboscidea merely nestles in the damaged apex of the 
gastropod shells but in others it erodes and drills the shell material. 

A living Mytilus californianus from rocks on the ocean side of the 
breakwater at Santa Barbara Yacht Harbor contained a B. proboscidea 
nestled in a pocket produced by a fracture between lamellae of the 
shell. The worm tube opened at the external edge of the shell follow- 
ing the spatial pattern characteristic of the blister worm, P. websteri 
Hartman, which occurs in oysters. 


SUMMARY AND DIscuSsSION 


Boccardia columbiana and B. proboscidea have been found in the 
Eastern Pacific at British Columbia in the north, and central Califor- 
nia in the south and at intermediate localities. Both occur in sandstone 
crevices, piling material, algal holdfasts, coralline algae, surface of 
rocks with Phragmatopoma californica, and in gastropod shells. They 
occur in living gastropod shells, in shells inhabited by hermit crabs, 
and in empty shells. Thus, the worms do not have an obligatory com- 
mensal relation with the hermit crabs. 

Although both species occur farther north in open surf regions, Boc- 
cardia columbiana is the dominant form in this habitat. Boccardia 
proboscidea was the dominant form in piling material from the quiet 
waters inside Santa Barbara Yacht Harbor. Boccardia columbiana has 
not been found in the sand-mud flat environment in which B. probo- 
scidea flourishes at Ballona Creek, Morro Bay, and Humboldt Bay. 

Morphologically the two species differ in the arrangement of setae 
of segment 1; this is the only characteristic consistently usable. Cer- 
tain pigmentation patterns and the general aspect of the anterior end 
may be of use in preliminary sorting of mature adults, but in general 
the younger adults of B. proboscidea resemble more the features 
ascribed to B. columbiana than those of their own adults. 

An unresolved problem of morphology associated with plankton 


—— 


138 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


studies is the determination of the stage in development at which the 
notosetae of segment 1 assume the adult appearance. If this occurs only 
at settling, free-swimming larval forms of the two species would be 
indistinguishable. Boccardia columbiana has not been investigated as 
to its development and larval characteristics. Hartman (1940, 1941 ) 
gave information on the development of Boccardia proboscidea. 

Nine species of Boccardia are known from the north eastern Pacific; 
they comprise about two-thirds of the world’s known species of the 
genus (Hartman, 1959: 375). Of the nine, six were originally de- 
scribed from this region. The north eastern Pacific, thus is unlike the 
Atlantic coast of the United States where members of Polydora are 
dominant and the central Pacific which has a greater representation 
of Pseudopolydora species. A key to the nine species of Boccardia is 
given below. 


Key to BOCCARDIA Sperctes From THE NorTHEASTERN PACIFIC 


1. Modified 5th segm.—branchiae lacking ant. toSegm.5................. 2 
la. Modified 5th segm.—branchiae present ant. toSegm.5...... (Boccardia) 3 
2. Neuropodial hooded hooks begin on Segment 7 ...............-. Polydora 
2a. Neuropodial hooded hooks begin on Segment 8 ........... Pseudopolydora 
3. Specialized setae of Segment 5—of one type .....................-005- 4 
3a. Specialized setae of Segment 5—of two types ................... 00005. 6 
4, Anterior branchiae on Segments 2, 3,-,-,-,7.......... Boccardia redeki 
4a, Anterior branchiae on Sesments 2,3, -,-— 6,7 .....:. 54.0.2 eee 5 
5. Posterior notopodial hooks present .........0.00...00see Boccardia uncata 
5a. Posterior notopodial hooks absent .................... Boccardia truncata 
6. Setae of Segment 5—Falcate and tridentate ........... Boccardia tricuspa 
6a. Setae of Segment 5—Falcate and bushy-topped ....................... 7 
7. Prostomium rounded... Joi. 05 0 galie ces wees 2 nee eee 8 
7a. Prostomatum: bifid). 2.30 wiles. ie ees wale ees eee 2 ose 9 


8. Notosetae of Segment 1— long, in fan-shaped group .. Boccardia columbiana 


8a. Notosetae of Segment 1- short, not so disposed ...... Boccardia proboscidea 

9. Posterior hooded hooks mainly falcate ................ Boccardia basilaria 
9a. Posterior hooded hooks bidentate .................. Joe ee 10 
10. Notosetae present on Segment 1 ...................... Boccardia natrix 
10a. Notosetae absent on Segment 1 ................. Boccardia polybranchia 


Comparison of two species of Boccardia 139 


LITERATURE CITED 


BERKELEY, E. 
1927. Polychaetous annelids from the Nanaimo district. 3. Leodicidae to Spioni- 
dae. Canad. Biol. Ottawa, Contr, n.s., 3:405-422. 


BERKELEY, E. anp C. BERKELEY 

1950. Notes on Polychaeta from the coast of western Canada. Polychaeta Seden- 
taria. Ann. Mag. Nat. Hist., ser. 12, 3:50-69. 

1952. Canadian Pacific Fauna, Polychaeta Sedentaria, No. 9 b (2). Toronto: Fish. 
Res. Board Canada, pp. 1-139. 


HARTMAN, O. 

1940. Boccardia proboscidea, a new species of spionid worm from California. 
J. Wash. Acad. Sci., 30:382-387. 

1941. Some contributions to the biology and life history of Spionidae from Cali- 
fornia. Allan Hancock Pacific Exped. Rpts., 7:289-324. 

1944, Polychaetous annelids from California, including the description of two 
new genera and nine new species. Allan Hancock Pacific Exped. Rpts., 
10(2):239-318. 

1959. Catalogue of the polychaetous annelids of the world. Allan Hancock Found. 
Publ., Occ. Pap., no. 23, 628 pp. 


HARTMAN, O. ann D. J. REISH 
1950. The marine annelids of Oregon. Oregon State Coll., Monograph Ser., pp. 
1-64. 


NEW SPECIES OF HYPERODES JEKEL AND A KEY TO 
THE NEARCTIC SPECIES OF THE GENUS. 
(COLEOPTERA: CURCULIONIDAE)! 


Wix~uiAM D. StocKToN 
Long Beach State College 


INTRODUCTION 


In the course of an extended study of the weevils of the genus Hyper- 
odes, the species described below were recognized as being new to 
science. I had planned to publish the descriptions of the new forms as 
a part of a longer paper on which work is not yet completed, but at 
the request of fellow workers, who wish to have the names available 
for reporting data of economic importance, I have decided to publish 
descriptions at this time. To facilitate studies by workers who may 
have to deal with members of this somewhat difficult group, I am also 
including a key to those species which have been encountered in North 
America north of Mexico. 


ABBREVIATIONS 


The following abbreviations have been used in the present paper to 
indicate the locations of specimens examined: 


(USNM) Collection of the United States National Museum 
(ELS) Collection of Dr. E. L. Sleeper 

(WDS) Collection of the author 

(CU) Collection of Cornell University 

(CARN ) Collection of the Carnegie Museum 

(HFH) Collection of Mrs. H. F Howden 


DESCRIPTIONS OF NEw SPECIES 


Hyperodes wallacei, new species 


Holotype: Rostrum shorter than prothorax, stout, broad, flattened 
above; tricarmate, densely clothed with dull grayish-yellow scales, 
bearing four rows of short, stout, setae; underside with filiform scales, 
apex peaene a few long bristles. Antennae reddish brown, segments 
one and two of funicle subequal, the first stouter, other forenio seg- 
ments moniliform, gradually becoming broader toward the club, all 


1Biological Sciences Paper No. 8, Long Beach State College. 


140 


New Species of Hyperodes 141 


the segments with a few fine bristles; club oval, covered with fine 
dense pubescence, bearing scattered longer bristles; obscurely three- 
segmented. Scrobes deep, slightly widened posteriorly, upper margins 
impinging against upper edge of eyes. Head with many round and a 
few strap-shaped, yellowish metallic scales. Frontal fovea moderate, 
ocular lobes feeble. Prothorax wider than long, sides moderately 
rounded, slightly constricted at apex, densely clothed with yellowish 
brown scales which are minutely granulose; a median and two lat- 
eral vittae of paler scales, all these vittae rather vague and ill-defined; 
an indication of a depressed line, almost a sulcus, along median vitta; 
a scattering of short, stout, clavate setae on the disc of the prothorax. 
Scutellum oval, clothed with tiny, white scales. Elytra slightly emar- 
ginate at base, densely clothed with yellowish-brown scales, larger 
than those of prothorax; striae well-impressed, punctures deep, re- 
mote; intervals convex, even-numbered ones more so, these with a row 
of stout, clavate setae; apices of elytra separately rounded and pro- 
longed. Pro- and mesothoracic sterna densely clothed with small fili- 
form scales medially, giving way to round scales at the sides; metas- 
ternum with round scales laterally, glabrous medially, coarsely and 
densely punctate and with numerous fine hairs. Abdominal sternites 
coarsely punctured, with fine hairs arising from the punctures. Apex 
of last sternite of male not impressed. Legs rather densely clothed 
with round scales, sparsely pubescent; tibiae denticulate and seti- 
gerous within, with feeble mucro. ‘Tarsi densely pubescent beneath, 
sparsely so above. 

The female of this species is unknown. 

Notes and Discussion: Holotype, male, (USNM), labelled as fol- 
lows: “Dunellon, Fla., Jun. 12, 1939, Oman’? One paratype, male, 
(CARN) ; three paratypes, males, (HFH) ; one paratype, male, (CU) ; 
two paratypes, males, (USNM); two paratypes, males, (ELS), 
labelled as follows: “Alachua Co., Fla.; 27-vii-54; H. V. Weems, Jr.; 
taken at light”: Total length, 3.9 mm.; pronotal length, 0.9 mm.; 
pronotal width, 1.2 mm.; elytral width, 1.7 mm.;: “Monroe Co., Fla.; 
v-1-53; N. J. and E. L. Sleeper, Collrs:’: Total length, 3.8 mm.; pro- 
notal length, 0.8 mm.; pronotal width, 1.0 mm.; elytral width, 1.8 
mm.; one paratype, male, (WDS), labelled as follows: “Swan Quar- 
ter, N. C.; vii-22-1953; W. M. Kulash; light trap” Total length, 3.7 
mm.; pronotal length, 0.7 mm.; pronotal width, 1.0 mm.; elytral 
width, 1.5 mm. The range in total length for all specimens which I 
have had the opportunity to examine is from 3.7 mm. to 4.7 mm. 

This species bears certain superficial resemblances to both H. alter- 


142 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


nata and H, annulipes. and agrees in many respects with the pub- 
lished description of H. peninsularis. However, the short, stout beak 
of the present species is distinctive, as is the general habitus of the 
weevil itself, and I see no choice but to establish a new species for the 
individuals I have studied which seem to belong together in agreeing 
with the above description. 

I am glad to have the opportunity to express my appreciation for 
the generous cooperation of Dr. George E. Wallace of the Carnegie 
Museum throughout the course of my studies on Hyperodes by dedi- 
cating this species to him. 


Specimens Examined: (USNM) 3 (CU) 1; (HFH) 3; (ELS) 2; 
(WDS) 1 


Distribution: Florida—Dunellon; Gainesville; Miami; Paradise 
Key; Pensacola: Monroe Co.; Alachua Co. 


North Carolina:—Swan Quarter. 


Biological Notes: Yaken in light trap at Swan Quarter, North Caro- 
lina by W. M. Kulash, also at ight by H. V. Weems, Jr. in Alachua 
Co., Florida. Swept from grass by N. J. and E. L. Sleeper in Monroe 
Co., Florida. 

It will be noted, as is true in many new species of insects, that not 
too much is known of the biology of this species, nor of the other species 
considered in this paper. 


Hyperodes texana, new species 


Holotype: Rostrum shorter than prothorax, stout, flattened above, tri 
carimate, the median carina the strongest; densely clothed above with 
round scales, densest toward the base; four rows of erect, yellowish 
setae, two on each side of the median carina; anteriorly the round 
scales give way to dense, metallic, yellow, semi-recumbent, filiform 
ones; apex with a few strong bristles; ventral surface with fine pu- 
bescence. Antennae reddish brown, scape clavate, segments one and 
two of funicle subequal, the first stouter; remaining funicular seg- 
ments moniliform, widening gradually toward the club, with fine 
pubescence and a few stronger bristles; club oval, finely pubescent, 
obscurely three- segmented; scrobes rather deep, slightly widened pos- 
teriorly, upper edge impinging against top of eye. Head densely 
clothed with yellow, metallic, strap-shaped scales; frontal fovea deep. 
receiving the median carina of rostrum. Prothorax wider than long, 
sides broadly rounded, densely covered with round, yellowish scales 


New Species of Hyperodes 143 


which are minutely granulose; a median and two6 lateral vittae of 
paler scales; disc of prothorax with scattered, rather short, slender 
setae; ocular lobes moderate; scutellum oval, covered with tiny white 
scales. Elytra moderately emarginate at base; densely scaly, the scales 
brownish yellow in color; striae well impressed, punctures deep, rather 
remote; intervals moderately convex, each with a row of short, slender, 
slightly clavate setae; tips of elytra conjointly rounded. Thoracic sterna 
densely scaly, both round and filiform whitish scales, restricted to 
the lateral edges of the metasternum; metasternum and abdominal 
sternites coarsely, densely punctured, and with fine, pale, sparse pu- 
bescence; last four abdominal sternites lighter in color; apex of last 
sternite of male with a vague, broad impression. Legs light reddish 
brown, sparsely pubescent; femora with a ring of yellowish scales dis- 
tally; tibiae more strongly setose, with scattered round scales, denti- 
culate and setigerous within, with rather a strong mucro. ‘Tarsi den- 
sely pubescent beneath and with fine, pale, sparse pubescence above. 

Allotype: Similar in almost all respects to the male holotype, the 
chief visible difference being that the apex of the last sternite of the 
female is deeply foveate and emarginate. 

Notes and Discussion: Holotype, male, and allotype, female, 
(USNM). The two specimens are mounted on separate points on the 
same pin which also bears the followimg labels: “McAllen, Texas; 
III-2-45; Fraser & Ball; reared in parsley; 45-8847" Eight paratypes 
(USNM). One paratype, male, (WDS), labelled exactly as is the 
holotype and allotype. Total length, 4.5 mm.; pronotal length, 1.0 
mm.; pronotal width, 1.2 mm.; elytral width, 1.8 mm. One paratype, 
female, (WDS), labelled exactly as is the male paratype. Total length, 
4.3 mm.; pronotal length, 0.8 mm.; pronotal width, 1.1 mm.; elytral 
width, 1.8 mm. I have named this species in deference to the fact that 
most of the specimens I have examined have come from the state of 
Texas. It bears superficial resemblances to echinata, sparsa, and rotun- 
dicollis, but the male genitalia are distinct. The size range in specimens 
I have examined is from 4.0 to 4.7 mm. in total length. 


Specimens Examined: (USNM) 108; (WDS) 2. 

Distribution: Louistana—Deeville; Violet; Chalmette; Meraux; New 
Orleans. 

Texas—McAllen; Mission; San Juan; Weslaco; Santa Maria. 


Biological Notes: This species has been reared from parsley at Violet, 
Louisiana; Deeville, Louisiana; and McAllen, Texas. It has been found 
in dill roots and stems at Mission, Texas, and on celery stems at San 


144 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


Juan, Texas. It has been reared from carrot roots at McAllen, Texas, 
and was collected from dead egg-plants (leaves) at New Orleans, 
Louisiana. Miss Warner of the U.S. National Museum informs me 
(personal communication ) that a species she determined as Hyperodes 
texana has been reported to be doing considerable damage to carrots in 
the lower Rio Grande Valley of Texas. 


Hyperodes hoodi, new species 


Holotype: Rostrum shorter than prothorax, broad, stout, flattened 
above, tricarinate, the median carina strongest, densely clothed with 
round brownish scales and lighter colored filiform ones; four rows of 
slender, erect, rather inconspicuous setae; underside with stiff, yellow, 
hair-like scales, a few longer bristles at apex. Antennae reddish brown, 
segments one and two of funicle subequal, both slightly clavate, other 
funicular segments with numerous rather strong bristles; club slender, 
oval, clothed with fine pale pubescence. Scrobes deep, suddenly 
widened posteriorly, upper margins impinging against upper part of 
eye; frontal fovea deep, receiving the median carina of the rostrum. 
Head densely clothed with yellowish, strap-shaped scales. Prothorax 
slightly wider than long, sides broadly rounded, ocular lobes weak, 
apex slightly compressed; densely clothed with rounded dull-brown 
scales; a median and two lateral vittae of paler scales; setae sparse. 
slender, inconspicuous; scutellum oval, clothed with tiny white scales. 
Elytra emarginate at base; striae well-impressed, punctures deep, re- 
mote; intervals slightly convex, each with a row of slender, dark, semi- 
recumbent, mconspicuous setae; tips conjointly rounded; elytra den- 
sely clothed with dull-brown scales, mottled with patches of paler ones, 
all the scales mmutely granulose. Thoracic sterna densely clothed with 
small, dirty-looking scales, giving way to round ones laterally on the 
metasternum. Abdominal sternites densely and finely punctured. 
with a fine, pale hair arising from each puncture; apex of last sternite 
of male not impressed. Femora sparsely haired, with a band of dull 
brownish scales distally; tibiae more densely setose, without scales, 
denticulate within. Tarsi densely pubescent beneath, with rather stiff, 
fine bristles dorsally. 

Allotype: The apex of the last abdominal sternite of the female is 
marked with a broad, rather indistinct concavity. Otherwise the sexes 
are very similar. 

Notes and Discussion: Holotype, male, in the collection of (USNM). 
The specimen bears the following label: “BRIT. AMER”. Allotype. 


New Species of Hyperodes 145 


female, (USNM), same label as the holotype. Seven paratypes, 5 
males, 2 females, (USNM). One paratype, male, (WDS), total length, 
4.7 mm.; pronotal length, 1.0 mm.; pronotal width, 1.2 mm.; elytral 
width, 1.9 mm.; same label as all other type material. I am unable 
to restrict the type locality at the present writing. Range is from 4.2 
mm. to 4.9 mm. in total length. 

This species resembles both sparsa and echinata in general habitus, 
but the male genitalia seem to prove conclusively that it is distinct. 

It is with great esteem that I dedicate this species to my good friend 
and wise counselor in many a taxonomic difficulty, Dr. J. Douglas 
Hood of Cornell University. 


Specimens Examined: (USNM) 9; (WDS) 1. 
Distribution: “British America’: 


Biological Notes: Nothing is known of the biology. 


Hyperodes dietrichi, new species 


Holotype: Body surface black and shining. Rostrum about as long 
as prothorax, tricarinate, the median carina strongest, coarsely punc- 
tured and rugose, slightly narrowed at middle, wider at base and apex; 
bearing long slender setae which are dark on the dorsal portion of the 
rostrum and light-colored and finer on the ventral surface; numerous 
fine, yellowish, recumbent, filiform scales scattered over the dorsal 
surface; a few long, pale bristles at apex; rostrum without round 
scales. Antennae reddish brown, the scape clavate, segments one and 
two of funicle subequal in length, the first stouter, remaining seg- 
ments of funicle subequal, moniliform; club slender, oval, and coy- 
ered with fine, short hairs, obscurely three-segmented; all the an- 
tennal segments bear scattered, rather strong bristles; upper margin 
of scrobe impinges against upper margin of eye; scrobes well-defined, 
scarcely widened posteriorly. Head coarsely, densely punctured, the 
punctures bearing very fine pale hairs; head without strong setae; 
frontal fovea profound, receiving median carina of rostrum. Pro- 
thorax with sides and base strongly rounded, slightly wider than long, 
apex slightly constricted; coarsely, confluently punctate, bearing many 
slender, recumbent hairs and numerous erect, slender, bristle-like 
setae; a vague median and stronger lateral vittae of yellowish round 
scales, the scales minutely granulose. Scutellum small, oval, covered 
with tiny pale scales. Elytra slightly emarginate at base; striae well- 
impressed, the punctures large and close-set; mtervals slightly convex, 
each with a row of long. slender, bristly setae; elytra rather sparsely 


146 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


covered with pale yellowish scales, these thicker laterally and on the 
declivity; tips of elytra conjomtly rounded. Thoracic sterna coarsely 
punctate, with sparse, small, dirty-yellow scales and numerous hair- 
like setae. Abdominal sternites rather coarsely punctured, bearing 
sparse, coarse pubescence, both punctures and pubescence finer on 
terminal segment; segments three and four narrow; apex of last ven- 
tral segment not impressed. Femora darker than tibiae, bearing sparse. 
fine, pale hairs; tibiae reddish brown, more heavily setose than femora, 
denticulate and setigerous within, with rather strong mucro, hind 
tibiae with a brush of long, flying hairs. Tibiae concolorous with tarsi, 
latter densely pubescent beneath. 

Allotype: The female has at the apex of the last abdominal sternite 
a small, well-marked fovea and lacks the brush of long flying hairs on 
the hind tibiae. Other characters agree very closely with those de- 
scribed for the male. 

Notes and Discussion: Holotype, male, (ELS, no. 73), labelled as 
follows: “San Magarita, C. Z., Panama’’; “V-15-46”; “E. L. Sleeper, 
Collr” ‘Total length, 3.2 mm.; pronotal length, 0.7 mm.; pronotal 
width, 1.1 mm.; elytral width, 1.6 mm. Allotype, female, (ELS). 
labelled the same as the holotype; total length, 3.3 mm.; pronotal 
length, 0.8 mm.; pronotal width, 0.9 mm.; elytral width, 1.4 mm. 
One paratype, female, (ELS), labelled as follows: “San Blas, Nayarit. 
Mexico”; “VII-14-60”: ““R. B. Loomis and J. Maris, collectors?’ Total 
length, 3.2 mm.; pronotal length, 0.8 mm.; pronotal width, 0.9 mm.; 
elytral width, 1.4 mm. One paratype, male, (USNM), labelled as fol- 
lows: “Mexico, II 13 45, Laredo, Tex’; “H. R. Cary, on orchid plant. 
45-3653”: ““Hyperodes sp., LLB °45” Total length, 3.5 mm.; pronotal 
length, 0.9 mm.; pronotal width, 1.1 mm.; elytral width, 1.6 mm. 

Iam unable to match my specimens with any of Champion’s species, 
described in the “Biologia Centrali-Americana?’ nor with any form 
in the Nearctic fauna of the genus, so that I have little hesitation in 
considering this to be a new species. I am herein restricting the type 
locality to San Magarita, C. Z., Panama. I felt that it was wise to dis- 
cuss the present species in this paper, along with the other, undoubted. 
Nearctic species, since it is very probably represented in other collec- 
tions in the United States. 


Specimens Examined: (USNM) 1; (ELS) 3. 
Distribution: Texas—Laredo. 
Mexico—San Blas; PANAMA—San Magarita, C. Z. 


Biological Notes: Dr. Sleeper (personal communication) took the 


New Species of Hyperodes 147 


holotype and allotype on orchid flowers at San Magarita. The female 
paratype (ELS) was taken with blacklight apparatus at San Blas. The 
male paratype (USNM) was taken on orchid at Laredo. 

It gives me great pleasure to dedicate this handsome species to my 
kindly mentor and good friend, Dr. Henry Dietrich of Cornell Uni- 


versity. 
Key To THE NEarcrTIC SPECIES OF Hy perodes 

1. Rostrum at least as long as or longer than prothorax .................. Y 
1a. Rostrum shorter than prothorax, stout, flattened above ............... iS) 
2. Second segment of funicle appreciably longer than the first, latter clavate, 
CODES MAIC Cmaps er ese scons os a'e¥e specs: 2a. opens loposeahpassia els. srapstey scheme spel guaicracs one 3 
2a. Second segment of funicle at most only slightly longer than the first, the 
lattersslender, but stouter than the second).......................-«-- 5 
3. Prothoracic scales moderate in size; elytral setae inconspicuous ......... 4 
3a. Prothoracic scales large; pronotum cribately punctured; elytral setae con- 
SPUCUOUSMMIP II Ga ae vai cich serene wurmiene a miele teu wt Sets evaunlereh a cuore aie cuaiate cry ptops 
4, Prothorax sub-cylindrical, sides almost straight; scales not metallic delumbis 
4a. Prothorax sub-orbicular, sides broadly rounded; scales of dorsum with 
TATE EA UM CPNUISLE LMU NAR Sie i ones ae pea a cee cteme oe aes ae ace Acne sane decepta 
ome alitermaterelytralvintervals setose -.. 5... seco sete net bose ne 6 
5a. Each elytral interval with a row of setae, at least on the declivity ........ 8 
OmElyvinalgsetaevlong. moderately clavate 5. .).-...2:-.0.c50+asse>seaue- 7 
6a. Setae short, strongly clavate, semi-recumbent ................. annulipes 
(merothoraxawith sides broadly rounded .>.2....552..--0.0.-- 4: alternata 
7a. Prothorax with sides almost straight; femora ringed with a band of large, 
ocelllatemsilivieny-wihite’scales.:. 4212. u0- Ges sess etek eee peninsularis 
8. Vestiture rather sleek, smooth-looking; scales with metallic or pearly 
QSREIE a0. 0,8 sbhar tac eheRO RCC Reece ene ep A RRA aoe AR tees Aa net eae 9 
8a. Vestiture rough; scales not metallic or nacreous .................000-- 10 
9. Prothorax moderately rounded; scales with metallic luster ...... hy perodes 
9a. Prothorax with sides very broadly rounded; scales of head, rostrum, and legs 
WANE CAE LASTOR. wi.5 a8 cus wideviueyina cB eons aes elersls: Cita eee a cea grypidioides 
HO Mleaccerspecies; over Om, im length) 30. 8s. so. s- laramiensis 
hOamesrialller-wesssthan\ orm: 3.8) .0) ao sees os a oe ote Sle el oe eee 11 
11. Prothorax with sides almost straight; ventral surface of abdomen densely 
SOAK . o's 08 AREA RS A ae SN op Rta sear a tee aoe MS tect SR eit vitticollis 
11a. Prothorax with sides broadly rounded; ventral surface of abdomen not coy- 
enedevitinderse:.scallesi sc)... 1ehec a Ree eas Seseetey Als evel cians chao a cere hentai a 12 
12. Opaque; very dull and dark-colored; scales very rough-looking; setae very 
COMSTINC UO USED sey tay te yoees) Siehalatleit coded Seis ust Sten llench whch oaaeel Meg au nee: Seog tens: SAS horni 
12a. More shining, not as dull or dark; scales more smoothly arranged; setae not 


GCOMSPLCWOWS: pany 2b Shela seen eRe SEs Aree Lous choca tale SSSA rath Oe RRP ARES 13 


148 


Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


. Setae moderately clavate; head thickly clothed with small scales, paler than 


those of prothorax...% 2346) .0086 Anode st seen cle eee montana 


. Setae slender, not clavate..... 00. 6.0. cs. 5552 oes 6 ee oe 14 


. Median vitta of prothorax composed of a double line of yellowish, rather 


shining scales (5 ./.i0' «cscs a0 ails sige oe oes «a dorsalis 
. Median. -vitta obsolete... ¢.4.0: 025.5 14)s4.0 seas cs see poseyensis 
. Body clothed with squamiform scales; not pubescent on the dorsal surface 16 
. Body clothed above with coarse pubescence .......... 7.3 eee 30 
. Upper edge of scrobe directed toward middle of eye .................. 17 
. Upper edge of scrobe directed toward top of eye ..................... 22 
. Elytral scales very tiny, not concealing the surface; surface very shining; 

elytral setae small, almost imvisible...............) eee 18 
. Elytral scales not tiny; elytral setae conspicuous ..................... 19 
. Rugosities of rostrum with scattered punctures ................. lodingi 
. Rugosities smooth, impunctate .................... ses eee carinata 
. Each elytral interval setose; elytra densely scaled .................... 20 
. Alternate intervals setose; elytra sparsely scaled ................ latinasa 
.. Frontal fovea ‘absent. . i .cciesic de jsein o on te Bele eee obtecta 
. Frontal fovea present . 0. 6.0660. 6s ccd sees ees a ee 21 
. Elytral setae stout, dense, clavate ....-.........-2 50 gue hirtella 
. Elytral setae slender, sparse, not clavate ...................... echinata 
. Surface of body very shining, not densely scaly; prothorax coarsely punc- 

tate; hind tibiae of male with a brush of long, flying hairs ............. 23 
. Surface densely scaly; prothorax with a granular appearance; male with- 

out a brush of long hair on the hind tibiae ......:.. 9. 242. sere eee 24 
. Alternate elytral intervals setose; setae clavate ................. humilis 
. Each elytral interval setose; setae long, slender, bristle-like ....... dietrichi 
. Alternate elytral intervals with setae; apex of each elytron separately pro- 

longed) ncsass 6 ego cdatee wha ane Sinker eos otra oe 25 
. Each interval setose; elytra conjointly rounded ...................... 26 
. Setae rather long, slender, slightly clavate; head clothed with filiform, yel- 

Jowish. scalés: ...i:02 .300@ nantes Manes We fe tate ene sparsa 
. Setae short, stout, strongly clavate; head clothed with both round and fili- 

form scales’ ..02 cic geblea awa ase Spas) pd oe one wallacet 
. Setae long, slender bristle-like ...... 2.0.0... o..:: 02s eRe novella 
. Setae shorter; body densely scaly .............55.. 2 Gee eee 27 
. Prothorax with sides almost straight; setae stout, rather conspicuous; elytral 

intervals sub-costaté ¢ 0.5 633.056 co bees oe aitalelasle eee obscurella 
. Prothorax with sides broadly rounded; setae slender, inconspicuous; intervals 

at most moderately convex... 6. bcc 0 Sel oe oe © oo nach eer 28 
. Scales of head fine, hair-like, sparse .........2. +... #2 eeeEeeee rotundicollis 
. Scales of head strap-like, dense ..........2.).-... +: see eee 29 


. Color light yellowish-brown; prothoracic vittae conspicuous; elytral setae 


moderately strong: <..../c0veenen voeaies ee oe ee eee texana 


. Color dull, dark brown; vittae and setae inconspicuous .............. hoodi 


30. 
30a. 
Sie 
31a. 


32. 
32a. 


New Species of Hyperodes 149 


Pubescence very sparse; surface of body shining ..................... 31 
Pubescence rather dense; surface of body dull, opaque ................ 32 
Larger; median prothoracic carina strong .................... anthracina 
Smaller; median carina variable, never very strong, sometimes obsolete; 
scrobes not greatly widened posteriorly ......................4. porcella 
Prothoracic carina short, abbreviated before and behind ....... maculicollis 
Carinaventine® linear =. 0... 26 Vals es oa Vo oe dea een eo bees as californica 


ACKNOWLEDGMENTS 


It gives me great pleasure to acknowledge, in addition to the apprecia- 
tion expressed above under the individual species, the generous en- 
couragement and unstinting help of my colleagues of the Biology De- 
partment of Long Beach State College, particularly Dr. Elbert L. 
Sleeper, Dr. James H. Menees, and Dr. Richard B. Loomis. I am also 
most grateful to Dr. David K. Caldwell of the Los Angeles County Mu- 
seum for his aid during the preparation of the manuscript of this paper 
for publication. 


NEW INFORMATION ON THE STRUCTURE OF PERMIAN 
LEPOSPONDYLOUS VERTEBRAE—FROM AN 
UNUSUAL SOURCE? 


PETER PauL VAUGHN 


University of California, Los Angeles 


INTRODUCTION 


Records of the variety of vertebrate fossils from given horizons and 
localities are frequently exaggerated in their proportions of large ani- 
mals. Skeletal elements of very small vertebrates are much less com- 
monly found in the usual prospecting and quarrying operations than 
are large, conspicuous elements. The reasons for this are obvious. 
Small, delicate elements are undoubtedly not readily preserved, espe- 
cially durmg the deposition of coarse sediments; but an even more 
nmportant reason is that the collector is likely to overlook the smaller 
elements in the usual field and laboratory procedures. This bias has 
been largely overcome in recent years through the development of 
various methods of washing and screening (see McKenna, 1962). An- 
other solution of this problem has not been employed to the full extent 
of its possibilities; this method is the careful search for skeletal ele- 
ments in coprolites. 

This method has a number of advantages. First, coprolites are usu- 
ally large enough to be easily seen and collected, and the collector’s 
eyes do not have to be focused for small elements. Second, the predator 
or scavenger to whom the coprolite pertains has concentrated and asso- 
ciated the elements. This could provide a valuable means of acquiring 
more thorough knowledge of animals that are otherwise known from 
only scattered, isolated elements. Palaeoniscoid fishes, for example, are 
often known only from such remains, frequently only from scales. 
Coprolites packed with scales offer the opportunity to study variations 
in scale structure in individual fossil fishes. One can, of course, rarely 
be sure that any given coprolite represents only one meal, that is, one 
individual animal eaten, but consistent association of the same kinds 
of parts in a number of coprolites can lead to confidence that these 
parts represent one kind of animal. 

Perhaps the most immediately interesting use of coprolites is m the 
search for new faunal elements. This report presents new information 


‘This study was supported by National Science Foundation grant NSF G- 
12456. 


150 


Permian Lepospondylous Vertebrae 151 


that was acquired in this way on the structure and occurrence of Per- 
mian lepospondylous amphibians. 


LEPOSPONDYL REMAINS IN A COPROLITE 


The Welles quarry near Arroyo de Agua, Rio Arriba County, north- 
ern New Mexico is a locality from which parties from the University 
of California, Berkeley have obtained many well preserved parts of a 
number of vertebrates (see Langston, 1953). This quarry is in the 
Cutler formation and is of early Permian (Wolfcampian) age. The 
Welles quarry, along with two nearby quarries, seems quite clearly to 
be part of a lake deposit. Its fauna includes pleuracanth sharks, at least 
one kind of palaeoniscoid fish, several genera of labyrinthodont am- 
phibians, the cotylosaur Diadectes, and two genera of pelycosaurian 
reptiles; there are also small pelecypods and a number of plants. In 
addition, there are a great number of coprolites, some of which were 
collected by a party from the University of California, Los Angeles in 
the summer of 1960. 

The coprolites are of many sizes, from as small as 1 cm. long to 
larger than 11 cm. long. There are a variety of shapes: fusiform, 
twisted, irregular. Some consist of spirally wound laminae, some of 
irregularly arranged but faintly concentric laminae. Almost all con- 
tain hard parts of vertebrates. Some contain palaeoniscoid scales, often 
along with small bones and bits of bones. Some contain parts of teeth 
of pleuracanth sharks. Some contain only unidentifiable fragments of 
bones. One at least, UCLA VP 1648, contains among other items a 
small vertebra with a urodele-like haemal arch. 

Before preparation was begun, this coprolite (No. 1648) was about 
70 mm. long. It was somewhat flattened, with an elliptical cross- 
section of about 33 mm. by 13 mm. The ends and sides were rounded. 
A polished transverse section shows irregularly concentric lamination. 
There are many small, fragile bones scattered throughout the copro- 
lite. These are difficult to prepare, but successful development of a 
selected few elements was carried out with the use of fine insect pins 
with points ground to sharp, beveled edges. 

A number of vertebral parts are present. One is a fairly complete 
vertebra a little more than 3 mm. long from the anterior end of the 
centrum to the posterior end of the neural arch. The transverse diam- 
eter of the centrum is slightly less than 1 mm. near the anterior end. 
The neural arch is rounded above, and the neural spine is represented 
by only a faint ridge. The transverse process arises from the neural 


i eayyey . 
OA ea EI A Cabal 
REINS we ds 


Figure 1. A caudal vertebra found in a coprolite from the lower Permian of New 
Mexico. UCLA VP 1648. A, dorsal, B, right lateral, and C, ventrolateral views. In 


the ventrolateral view, only the right side and distal end of the haemal arch can 
be seen. Unshaded areas represent matrix. x20. 


Permian Lepospondylous Vertebrae 153 


arch in a broad base whose center lies about 1 mm. posterior to the 
front end of the centrum; it extends directly laterally for about 0.5 
mm., narrowing to a single costal articulatory facet slightly less than 
0.5 mm. in diameter. The structure of the sides and bottom of the 
centrum is not clear. Much of the anterior part of the neural arch is 
missing, and a natural cast of the relatively large neural canal is there- 
by exposed. On the right side, it can be seen that the neural arch broke 
away from the centrum along a smooth line that curves lateralwards 
near the anterior end of the vertebra. This smooth line of parting is 
shown also by a centrum—lacking the neural arch completely—else- 
where in the coprolite. This indicates the presence of a neurocentral 
suture. 

The most interesting element in the coprolite is the caudal vertebra 
illustrated in Figure 1. The centrum of this vertebra is about 3 mm. 
long; the posterior zygapophyses were damaged in preparation, but 
the distance from the anterior edge of the prezygapophyses to the pos- 
terior edge of the postzygapophyses, about 3.5 mm., was measured 
prior to the damage. The neural arch is swollen to an almost circular 
cross-section, greater in diameter than the centrum except at the ends 
of the vertebra. The neural spine is only a slight ridge, commencing 
about midway along the neural arch and becoming more prominent 
posteriorly. The dorsal part of the neural arch flares gently to become 
broader from side to side in the regions of the zygapophyses. The pre- 
zygapophyses have horizontal, slightly concave articular facets and 
are connected to one another via a semicircular notch. As may be seen 
in the illustration, the right prezygapophysis was chipped along its lat- 
eral edge in preparation. Because of breakage, details of structure of 
the postzygapophyses cannot: be described. The smooth concavity of 
the intervertebral notch for the spinal nerve can be easily made out on 
the right side between the postzygapophysial area and the centrum. 
There is no costal facet; presumably this vertebra occupied a fairly far 
posterior position in the caudal series. Faint traces of the neurocentral 
suture can be seen, especially near the anterior end. The lateral sur- 
faces of the centrum, unlike the surfaces of the neural arch, are not 
swollen; a cross-section midway through the centrum would show flat 
sides. The centrum is somewhat pinched midway in its length and 
flares at either end to a width approximately equal to that of the 
neural arch. The ventral surface of the centrum is a broad, shallow 
trough. Passing away insensibly from the ventrolateral edges of the 
centrum is a haemal arch. The base of this arch is broad and extends 
farther anteriorly along the centrum than it does posteriorly; the 


154, Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


distal end of the arch is narrower and lies about midway along the 
length of the centrum. 

The caudal vertebra that has been described is very similar to caudal 
vertebrae in living urodeles. ‘Taking a caudal vertebra of a urodele as, 
say, Cryptobranchus for comparison, the only differences are that in 
Cryptobranchus there is a short neural spine jutting posterodorsally 
beyond the postzygapophysial area and there is a spike-like postero- 
ventral continuation of the haemal arch. The latter difference may not 
be real; on the ventral surface of the rounded distal end of the haemal 
arch of the vertebra in the coprolite can be seen a small area of break- 
age that may indicate the former presence of such a spike-like distal 
process. 

There are other elements besides vertebrae in the coprolite. These in- 
clude at least one rib that seems to have both capitular and tubercular 
facets. There are also a number of what appear to be limb bones and 
parts of limb bones. One of these seems to be half of a propodial, broken 
away from the missing half at the narrow waist of the shaft, presum- 
ably about midway in the length of the original bone. The preserved 
half is about 4mm. long, is about 1.4 mm. thick near its articular end, 
and is about 0.6 mm. thick at its broken end. 

The general similarity of all the various vertebral parts in the copro- 
lite to one another and the general fitness in size of all the elements 
to one another make it seem likely that they all came from one animal. 


DiscussIoNn 


Romer (1945) has classified the urodeles and apodans (gymno- 
phionans) along with a number of highly problematical groups of 
Carboniferous and Permian vertebrates as members of the amphibian 
subclass Lepospondyli. The order Microsauria, in particular, has pre- 
sented thorny problems in phylogeny. The microsaurs have been con- 
sidered variously as possible relatives of the urodeles, of the apodans, 
and of at least the captorhinomorph suborder of cotylosaurian reptiles 
(see Vaughn, 1962). The major uniting feature of the Lepospondyli 
is the presumed common possession of “‘lepospondylous” vertebrae 
that arise, as in living members of the subclass, by direct formation of 
the centra through ossification around the notochord with little or no 
preformation in cartilage. The lepospondylous vertebra shows no sign 
of the division into intercentral and pleurocentral parts that character- 
izes the labyrinthodont amphibians and that is still evident in the ver- 
tebrae of reptiles and even mammals, although Williams (1959) feels 


Permian Lepospondylous Vertebrae 155 


that the centrum of the living lepospondyls can be equated with the 
major central element, the pleurocentrum, of living amniotes. 

In labyrinthodont amphibians, reptiles and mammals, the haemal 
arches (chevrons) of the caudal region are outgrowths of intercentra. 
In urodeles, contrariwise, the haemal arches are outgrowths of the 
centra and there is no sign of intercentra. It is known that such a “cen- 
tral” haemal arch was present also in the members of the Carbonifer- 
ous and Permian lepospondylous order Nectridia, an order that in- 
cludes the well known Diplocaulus, but the nectridian haemal arch is 
quite different from that of urodeles in that the distal portion is long 
from front to back and flattened from side to side, usually with verti- 
cal fluting along the sides; frequently, the nectridian haemal arch is 
constricted at its base, so that it is fan-shaped in lateral aspect. In nec- 
tridians the neural arch in the caudal region looks much like the 
haemal arch, the total effect being a flattened tail that was obviously 
of use in sculling locomotion. Except for its “central” position, the 
haemal arch of the vertebra in the Welles quarry coprolite is not at 
all like that found in nectridians; it is very much like that seen in 
living urodeles. 

To date, the only lepospondyl of any kind reported from the Per- 
mian of New Mexico is the microsaur Pantylus, and this is known 
from only a smgle mandible found at the Anderson quarry near Ar- 
royo de Agua (Langston, 1953). This is in marked contrast to the 
number of lepospondyls found in approximately contemporaneous 
lower Permian deposits in northcentral Texas, whence are known: 
two genera of Nectridia, a genus of the Lysorophia, and a half dozen 
genera of the Microsauria, among them Pantylus (Romer, 1950a, 
1960). The very small size of the vertebrae in the coprolite from the 
Welles quarry makes it seem highly unlikely that they might pertain 
to Pantylus. Further, Williston (1916) reported caudal vertebrae with 
intercentral haemal arches found in close association with parts of 
three skulls of Pantylus in Texas. As Romer (1950b: 640) has noted, 
“Tf the caudal vertebrae figured by Williston are actually those of 
Pantylus they are of considerable theoretic interest in bearing haemal 
chevrons:’ The theoretic interest stems from the intercentral position 
of the chevrons, a condition that would tend to remove microsaurs from 
the close relationship they have been thought to have with the other 
orders classified under Lepospondyli. It must be noted, however, that 
Case (1929) failed to find any trace of intercentra in Ostodolepsis, pre- 
sumably closely allied to Pantylus, and that there is no good evidence 
for intercentra in other microsaurs. Steen (1938) reported poorly pre- 


156 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


served intercentral haemal arches in one specimen of the Pennsyl- 
vanian microsaur Microbrachis, but, as Romer (1950 : 633) has re- 
marked, “Since . . . they do not appear in other individuals of this or 
in related forms, and since some confusion with caudal ribs is not im- 
possible in a small and poorly preserved specimen, judgment may be 
suspended on this feature’’ 

Granted that the evidence for intercentral haemal arches in micro- 
saurs is not good, neither is there any reason for believing that they 
had ‘‘central”’ haemal arches; indeed, there is some indirect evidence 
that they did not have such arches. Gregory, Peabody and Price 
(1956) studied microsaurian skulls and postcranial parts from the 
lower Permian fissure deposits near Fort Sill, Oklahoma. The bones 
in these deposits are rarely found in articulation, and referral of iso- 
lated vertebrae to any particular genus or even larger group is diffi- 
cult. The microsaurian atlas seems to be represented by several strik- 
ingly urodele-like specimens (op. cit.: 43) The dorsal vertebrae that 
seem to belong to microsaurs differ from those of urodeles in that the 
transverse processes arise from near the front of the neural arch and 
bear only one articular facet, in that there is a neurocentral suture, 
and in other, less obvious ways. In these same fissure deposits were 
found a number of caudal vertebrae that were referred to the micro- 
saur Cardiocephalus with “great hesitation” (op. cit.: 46-47) because 
“The only features that distinguish these from known caudal verte- 
brae of [the cotylosaur] Captorhinus are their smaller size and the 
absence of any trace of the transverse fissure permitting autotomy of 
the tail. . . °° Nevertheless, in spite of the uncertainty in identification 
of these vertebrae, it must be noted that no caudals were found with 
any trace of ‘central’? haemal arches. For this reason, as well as the 
caution prompted by Williston’s report of intercentral chevrons in 
Pantylus, it would seem to be unwise at this time to suppose that the 
vertebrae in the Welles quarry coprolite represent a microsaur. 

This leaves us with no known order to which the vertebrae in the 
coprolite may be confidently assigned. That these vertebrae are of a 
basically lepospondylous nature would seem to be clear from the na- 
ture of the haemal arch. Of the groups of Paleozoic lepospondyls be- 
sides microsaurs: (1) in the Nectridia, as has already been pointed out. 
the haemal arches look quite different, and nectridian vertebrae do 
not have a neurocentral suture; (2) there is no evidence of haemal 
arches in lysorophians, in at least Lysorophus and Megamolgophis 
the neural arch is composed of two separate lateral halves, and the 
limbs in these animals were apparently much reduced (3) aistopods 


Permian Lepospondylous Vertebrac 157 


were limbless and had ribs with peculiar processes quite unlike the 
ribs seen in the coprolite. Of living lepospondyls: (1) in the vertebrae 
of apodans the transverse processes are placed far forward on the 
neural arch, but there are special “‘infrazygapophyses” that provide 
additional articulatory surfaces along the ventral parts of the centra, 
and besides, caudal vertebrae are almost nonexistent in the apodans 
and when present do not have haemal arches; (2) the vertebrae of 
urodeles are nectridian-like in the placement of the transverse process 
midway along the centrum and in that the transverse process usually 
is inclined posteriorwards and has two articular facets; (3) neither 
apodan nor urodele vertebrae have neurocentral sutures, but this, as 
Gregory, Peabody and Price (op. cit.) point out, may not be signifi- 
cant. The known vertebrate fauna of the Permian is, of course, proba- 
bly only a small sample of the fauna actually living at that time, and 
the sampling error is undoubtedly even greater when only the smaller 
forms are considered. It is entirely possible that the animal represented 
in the coprolite belongs to some lepospondylous group otherwise un- 
known. It may also represent a much-modified variant of one of the 
known Paleozoic groups. It may even be seen eventually to help cor- 
roborate the opinion of Gregory, Peabody and Price that the urodeles 
were derived from nectridian-like forms, but fossils of connecting 
forms are needed to establish this. 

Even if we put aside the problem of ordinal assignation of the form 
represented by the bones in the coprolite, this animal is still of consid- 
erable interest in that it shows such a strikingly urodele-like haemal 
arch in the Permian, in that it adds to our picture of the early Permian 
fauna of New Mexico, and in that it demonstrates the possible rewards 
of diligent search for skeletal ‘elements in coprolites. 


158 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


ACKNOWLEDGMENT 


The drawings that illustrate this paper were made by Miss Susan 
Ahrend. 


LITERATURE CITED 


CASE, E. C. 
1929. Description of a nearly complete skeleton of Ostodolepis brevispinatus Wil- 
liston. Contrib. Mus. Paleont, Univ. Michigan, 3(5):81-107. 


GREGORY, JOSEPH T., PEABODY, FRANK E., anp PRICE, LLEWELLYN I. 
1956. Revision of the Gymnarthridae, American Permian microsaurs. Bull. Pea- 
body Mus. Nat. Hist., 10:1-77. 


LANGSTON, WANN, JR. 

1953. Permian amphibians from New Mexico. Univ. California Publ. in Geol. 
Sci., 29(7) :349-416. 

McKENNA, MALCOLM C. 

1962. Collecting small fossils by washing and screening. Curator, 5(3):221-235. 

ROMER, ALFRED SHERWOOD 

1945. Vertebrate Paleontology. Univ. Chicago Press, 687 pp. 

1950a. The upper Paleozoic Abo formation and its vertebrate fauna. Soc. Vert. 
Paleont. Guidebook for northwestern New Mexico, 4th Field Conf., pp. 47-55. 

1950b. The nature and relationships of the Paleozoic microsaurs. Amer. J. Sci., 
248: 628-654. 

1960. The vertebrate fauna of the New Mexico Permian. New Mexico Geol. Soc. 
Guidebook of Rio Chama Co., 11th Field Conf., pp. 48-54. 


STEEN, MARGARET C. 
1938. On the fossil Amphibia from the gas coal of Nyrany and other deposits in 
Czechoslovakia. Proc. Zool. Soc. London, ser. B, 108(2):205-283. 


VAUGHN, PETER PAUL 

1962. The Paleozoic microsaurs as close relatives of reptiles, again. Amer. Midi. 
Nat., 67(1):79-84. 

WILLIAMS, ERNEST E. 

1959. Gadow’s arcualia and the development of tetrapod vertebrae. Quart. Rev. 
Biol., 34(1):1-32. 

WILLISTON, SAMUEL WENDELL 


1916. The osteology of some American Permian vertebrates. II. Contrib. Walker 
Mus. Univ. Chicago, 1(9) :165-192. 


PROCEEDINGS OF THE ACADEMY 


The Southern California Academy of Sciences met nine times during 
the fiscal year 1962-1963. Eight of these meetings were held in the 
Auditorium at the Los Angeles County Museum, and the Annual 
Meeting was at the University of Southern California. The section on 
Earth Sciences held several meetings at different schools in the area. 

The following talks were given at the monthly meetings. The Bot- 
any Section, on September 21, 1962, presented Dr. D. Foard of 
U. C. L. A. who spoke on the “Developmental Anatomy of the Camel- 
lia Leaf” The Invertebrate Zoology Section was in charge October 19, 
1962, when Dr. Irwin M. Newell, Univ. Calif., Riverside, gave a talk 
“Biology and Systematics of the Giant Red Velvet Mites of the Desert?” 
On November 16, 1962, the Experimental Biology Section presented 
Dr. Richard M. Straw, of Los Angeles State College, who discussed 
“Cytomegalovirus (Human Salivary Gland Virus) in Tissue Culture 
Cells” The Vertebrate Zoology Section heard Dr. Charles A. Mc- 
Laughlin of the Los Angeles County Museum talk on ““The Mammals 
of Chad, Africa’ January 18, 1963. Professor Emma Lou Davis, 
U.C.L.A., reported on the “Migration of the Prehistoric Mesa Verdes 
Peoples,’ sponsored by the Anthropology Section, on February 15, 
1963. The Earth Sciences Section presented Dr. Theodore Downs of 
the Los Angeles County Museum, who talked about the “Fossil Verte- 
brates of the Anza-Borrego Desert;’ on March 15, 1963. The Junior 
Academy on April 19, 1963, presented three students who discussed 
their studies. 

The newly elected Board of Directors and Advisory Board met on 
May 3, 1963, to hear reports and elect officers for 1963-1964. 

The third Annual Meeting was held at the University of Southern 
California, on May 18, 1963. More than 120 persons attended the Sci- 
entific Sessions, including a special symposium with eight papers, and 
the general session with 18 papers in four sections. The student par- 
ticipants were judged for two student prizes which were presented at 
the Annual Dinner for the first time. 

The papers presented were: Symposium “Adaptation to Cold” Dr. 
Harry Sobel, Chairman and organizer. 1. ‘““Some Comparative Aspects 
of Cold Acclimation in Birds and Mammals.’ Dr. Rowand R. J. Chaf- 
fee (speaker) and Dr. Wilbur W. Mayhew, University of California, 
Riverside. 2. “Fatty Acid Metabolism in Cold Exposure and Cold Ac- 
climation”’ Dr. Edward Masoro, University of Washington, Seattle. 
3. “Some Physiological Aspects of Cold Acclimatization” Dr. John 


159 


160 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 3, 1963 


Patrick Mehan, University of Southern California. 4, “Physiological 
Role of Brown Fat?’ Dr. Robert E. Smith, University of California, Los 
Angeles. 5. “Progressive Reduction of Low Temperature Induced 
Fibrillation Point by Hypothermia Exposure”’ Dr. Dale L. Carpenter, 
Marquart Corporation. 6. “Profound Hypothermia in the Rabbit” Dr. 
David Norman, North American Aviation. 7. “Changes in Guinea 
Pigs Exposed Continuously to Temperatures of 2-4°C for One Year’ 
Dr. Harry Sobel, Veterans Administration Hospital, Sepulveda. 8. Dr. 
G. S. Bajwa, Institute for Biological Research, Culver City, presented 
histological findings. 


General Session papers: 


Studies on the Mytilus edulis community in Alamitos Bay, California: Develop- 
ment and destruction of the community. Donald J. Reish, Long Beach State 
College. 

Evolutional Trends in the Chonotrichida (Protozoa, Ciliata). John L. Mohr, H. 
Matsudo, and Y. M. Leung, University of Southern California, and V. L. Greg- 
ory, California Polytechnic College, Pomona. (Presented by Mohr) 

An Ecological Study of the Green Lynx Spider Population of Northwestern Flor- 
ida. Donald C. Lowrie, Los Angeles State College. 

Fortuitous Appearance of Two Exotic Fishes in the California Fauna. Carl L. 
Hubbs, Scripps Institute of Oceanography. 

The University of Southern California Antarctic Program and Some Preliminary 
Results of Work in the Bransfield Strait. Hugh H. Dewitt, University of South- 
ern California. 

Evolution in the Viperfish Chauliodus sp. Jules Crane, Cerritos College. 

New Information on the Structure of Permian Lepospondylous Vertebrae—from 
an Unusual Source. Peter Vaughn, University of California, Los Angeles. 

Heavy Machinery in Archeology. Ruth D. Simpson, Southwest Museum. 

The Transition of Isopods from a Marine to a Terrestrial Habitat. Gary J. Brusca, 
University of Southern California. 

Studies on the Ecology of Planktonic Foraminifera and Radiolaria off the South- 
ern California Coast. Richard Casey, University of Southern California. (Co- 
winner of student award) 

Water balance of the Florida Mouse. Daniel S. Fertig, University of Southern 
California. 

Metabolic Aspects of Temperature Regulation in Lepidoptera. J. E. Heath, Uni- 
versity of California, Los Angeles, and P. A. Adams, University of California, 
Santa Barbara. (Presented by Heath) 

Some Investigations in Cinephotographic and Microphotographic Techniques. 
William R. Stanley, and Wm. J. Bruff, Cerritos College. (Presented by Stan- 
ley) 

Comparative Osteology of the Plethodontid Salamander Genus Aneides. David B. 
Wake, University of Southern California. (Co-winner of student award) 

Winter-time Biota of the Pacific Coast Lagoons of Central Baja California, Mex- 
ico. Robert Eberhardt, Lockheed-California Company. 


Proceedings of the Academy 161 


The late Miocene Mammal Fauna from Camp Creek, Nevada. J. R. Macdonald, 
Los Angeles County Museum. 

Pleistocene Lake Panamint, Panamint Valley, California. Thomas Clements and 
Lydia Clements, University of Southern California. (By title only) 

Late Pleistocene Bird Fossils from the Channel Islands, California. Hildegarde 
Howard, Los Angeles County Museum. 


Seventy-five members and guests attended the Annual Diner 
Meeting which was held at the Faculty Center, University of South- 
ern California. Dr. Herbert Friedmann, Director, Los Angeles Coun- 
ty Museum, presented a talk on “Aspects of Evolution in a Genus of 
Crested Cuckoos”’ 

Fellows elected to the Academy were announced at the Annual 
Dinner. They were: Mr. Cyril EF Dos Passos, Mendham, New Jersey; 
Dr. Herbert Friedmann, Los Angeles; and Dr. Laurence M. Klauber, 
San Diego. 

Co-winners of the student awards were Mr. Richard Casey and Mr. 
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Dr. Theodore Downs, retiring president, relinquished the gavel to 
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office. 

Thanks were extended to Dr. Jay M. Savage, Chairman of the 
Local Committee, and to other faculty members and graduate stu- 
dents in the Department of Biology, University of Southern Califor- 
nia, for their help in conducting a successful Annual Meeting. 


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id 
“BULLETIN OP eTHE” 


Southern California 
Academy of Sciences 


LOS ANGELES, CALIFORNIA 


VoL. 62 OctToBER-DECEMBER, 1963 Part 4 


CONTENTS 


An experimental study of the echolocation ability of a California 
sea lion, Zalophus californianus (Lesson). William E. Evans 
and Ruth M. Haugen 


The number of conenose bugs, Triatoma, infected by one engorge- 
ment on a mouse with Trypanosoma cruzi. Sherwin F. Wood .. 


Birds and Indians in the West. Loye Miller 
Contributions from the Los Angeles Museum—Channel Islands 
biological survey. 37. Brachydont desmostylian from Miocene 


of San Clemente Island, California. Edw. D. Mitchell, Jr. ... 192 


Notes on the ova of six California moths. John Adams Comstock .. 202 


A study of the Zygopinae (Coleoptera: Curculionidae) of America 
north of Mexico, I. Elbert L. Sleeper 


Theodore Payne, 1872-1963. Bonnie C. Templeton 


Index for volume 62 


Issued December 31, 1963 


Southern California 
Academy of Sciences 


OFFICERS 
Richard B. Loomis) ...ja.i: a2 oe oe eo nee ee eee President 
John AL Witte yee. ee IRC 5 cite ae tty RN aa ee a First Vice President 
Peter P. Vaughn... . 2)... say's aie ae en eer Second Vice President 
Gretchen Sibley 00. 202025.) sce Gale «(eas citer eee Secretary 
Toy d Vi. Martin (220.5. .2 eiek. Sec See ee Assistant to Secretary 
W.Dwight Pierce «5 15.15.50 ten he dyn ete ies toate. eee: eer Treasurer 
David K: Caldwell: ye. 60. Sek Se Editor 
DIRECTORS 


David K. Caldwell 
E. Yale Dawson 
Richard B. Loomis 
Lloyd M. Martin 


James A. Peters 
W. Dwight Pierce 
Jay M. Savage 
Gretchen Sibley 


Ruth D. Simpson 
Peter P. Vaughn 
John A. White 


ADVISORY BOARD 


Phillip A. Adams 


Theodore Downs 


Kenneth E. Stager 


A. Weir Bell Richard Etheridge Richard H. Swift 
Russell E. Belous Herbert Friedmann Richard Tedford 
Bayard Brattstrom Hildegarde Howard Fred S. Truxal 
Henry E. Childs, Jr. Charles McLaughlin Louis C. Wheeler 
John A. Comstock Elbert Sleeper Sherwin Wood 


STANDING COMMITTEES 


Finance Library 

Russell E. Belous, Chairman Dorothy E. Martin, Chairman 
Publications Membership 

Fred S. Truxal, Chairman James A. Peters, Chairman 
Conservation Student Committee 

Henry E. Childs, Jr., Chairman Harry Sobel, Chairman 

Publicity 
Theodore Downs, Chairman 
SCIENCE SECTIONS 

Anthropology Experimental Biology 

Russell E: Belous, Chairman Ronald A. Kroman, Chairman 
Botany Invertebrate Zoology 

E. Yale Dawson, Chairman John L. Mohr, Chairman 
Earth Sciences Vertebrate Zoology 


Richard H. Tedford, Chairman 


James A. Peters, Chairman 


The Bulletin is published quarterly by the Academy. Address all communications to the 
appropriate officer at the Los Angeles County Museum, Los Angeles 7, California. 


Printed by Anderson, Ritchie & Simon, Los Angeles, California 


PUGctivwiiN OK THE SOULHERN CALIFORNIA 
ACADEMY OF SCIENCES 


VoL. 62 OcroBER-DECEMBER, 1963 Part 4. 


AN EXPERIMENTAL STUDY OF THE ECHOLOCATION 
ABILITY OF A CALIFORNIA SEA LION, 
ZALOPHUS CALIFORNIANUS (LESSON ) 


Wituram E. Evans! anp RutH M. HAucEn? 
Lockheed-California Company, Burbank, California 


INTRODUCTION 


It has been well established that dolphins, specifically the Atlantic 
bottlenose dolphin (Tursiops truncatus), are endowed with an ex- 
tremely efficient biological “sonar” system (McBride, 1956; Schevill 
and Lawrence, 1956; Kellogg, 1958, 1959; Norris, Prescott, Asa Do- 
rian and Perkins, 1961). During several cruises of the Lockheed-Call- 
fornia Company Research Vessel Sea Quest early in 1962, another 
marine mammal, the California sea lion, Zalophus californianus Les- 
son, was observed to produce at least five different types of sounds 
under water during what appeared to be feeding and investigating ac- 
tivities. A sonagram of one of the more characteristic of these sounds 
is presented in Figure 1. The waveform of these sounds and the be- 
havior associated with their production was indicative of some form 
of echolocation. However, the possible presence of various unobserved 
delphinids, which are known to be sound makers, e.g., the Pacific Pilot 
Whale, Globicephala scammoni and the Pacific whitesided dolphin, 
Lagenorynchus obliquidens, (Schevill and Watkins, 1962), in the 
immediate area made the positive identification of these sounds as 
being of sea lion origin tenuous. However, these field observations 
were recently verified by the observation of underwater sound pro- 
duction by various captive pmnipeds (Poulter, 1963; Schevill, Wat- 
kins and Ray, 1963). 

In order to further verify these observations, especially in reference 
to the use of these sounds for echolocation, the following series of tests 
was conducted. 


1Also Research Associate in Marine Zoology, Los Angeles County Museum. 


“Now with Sea World Inc., San Diego, California 


165 


166 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


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Figure 1. Sonogram of one of several underwater sounds produced by a Califor- 
nia sea lion Zalophus californianus (Lesson). This sound was recorded at a loca- 
tion 1000 yds. off of Santa Barbara Island, California. 


Sea lion echolocation ability 167 


1. A single captive California sea lion was monitored during various 
types of activity to evaluate its underwater sound production ca- 


pability. 


to 


The same animal was trained to retrieve objects made of a chemi- 
cally inert material. It was determined whether or not sound was 
used in conjunction with the detection and retrieval of these objects 
under various conditions of visibility. 


3. The same animal’s vision was completely suppressed by blindfold- 
ing and its navigation and target detection capability observed. 


PROCEDURE 


The experimental animal, a one year old female California sea lion, 
was maintained in a circular steel tank, 26 feet in diameter, 9.5 feet 
deep, with a capacity of approximately 40,000 gals. An 8-foot wide 
work platform used for animal training occupied one side of the tank. 
Although this facility was capable of handling sea water, fresh water 
was used to simplify maintenance. 


Training and Tests. ‘The sea lion used in this study was placed in the 
laboratory tank on December 18, 1962. At the time of procurement 
she had been in captivity three weeks and would eat dead fish. Other 
than this small degree of taming she was essentially untrained. 

From the time of her initial introduction into the tank her sounds 
were periodically monitored using a Massa, M-115 hydrophone, 
Massa, M-185 hydrophone power supply and amplifier, and a Clevite 
Ordnance Type O, hydrophone and a specially constructed pream- 
plifier. All signals were recorded on a Vega, 4-channel tape recorder 
at a tape speed of 7.5 ips. In addition, the animal’s underwater be- 
havior was observed through four, 23-inch underwater viewing ports. 

Rings made of 3g inch plastic tubing, 6.5 inches in diameter and 
filled with air were used for the retrieval targets. These were easy for 
the animal to pick up, and would produce a good echo return. Each 
ring was weighted with lead shot to make it smk. The weight was 
placed at one point on the circumference of the rmg so that it would 
stand up-right on the bottom of the tank. 

The animal was trained to accept the test rings on her training 
stand at first, then retrieve them from the floor of the training plat- 
form. She then progressed to water, retrieving floating rmgs and then 
finally submerged rings. During the water retrieval phase of the train- 
ing, records were kept of her retrieval time, and search pattern. All 
initial trials were monitored acoustically. 


168 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


During the retrieval training, several problems were encountered 
in the animal’s behavior that affected final handling techniques. At 
first the sea lion would watch the target rings, jumping into the water 
as soon as the target was thrown. Not only was insufficient time al- 
lowed for the ring to sink to the bottom of the tank, but also the animal 
could see exactly where the ring was thrown and no search was neces- 
sary. To offset this problem, the trainer held the animal’s attention 
by feeding her pieces of fish while another experimenter threw the 
rings into the water. However, the sea lion then began to respond to 
the sound of the ring hitting the water. Consequently, it was neces- 
sary to employ “false targets.’ As the trainer was supplying the ani- 
mal with small bits of food, talking to her, and putting her through 
various simple tricks (e.g., shaking hands), the other experimenter 
would splash the water by repeatedly throwing in a ring and pulling 
it out by means of an attached string. Thus, responses to false targets 
were not rewarded. However, still another anticipatory response was 
to occur. When the experimenter stopped splashing water, the sea 
lion went after the smking ring. By splashing water for a few trials 
without throwing in a target, this undesirable behavior was eliminated. 

The possibility of changes in retrieval time from trial to trial as a 
function of the food consumed during the trials was considered and 
checked. During one training session, using floating rings, the re- 
trieval times of the initial trials were of the order of 6 seconds. By the 
40th trial, retrieval time had slowed to 20-40 seconds as the animal 
appeared “bored or distracted”? However, during the 50th through the 
80th trials, retrieval times were again in the 5-8 second range. There 
was no noticeable “time” or “food” effect during the test sessions. On 
a test day, the animal generally had been fed only a few fish prior to 
the tests. The maintenance diet was approximately 7.5 pounds of 
mackerel, squid, or white bait per day. 

When the underwater target retrieval behavior was well estab- 
lished, a series of daylight and night test trials was run. The final 
handling technique was as follows. The sea lion was positioned on her 
training stand with her back to the water. The trainer fed her small 
fish (white bait), talked to her, and put her through simple tricks. 
Simultaneously another experimenter splashed the water with a false 
target ring. During the splashing, the target rig was thrown into the 
water. After the target rmg had sunk to the bottom of the tank, (ap- 
proximately 15 sec.), the tramer was signaled. The trainer then sent 
the sea lion into the water by the verbal command “go find it?” The 
time interval from the animal’s entrance into the water until exit onto 


Sea lion echolocation ability 169 


the platform with the rmg was considered the “retrieval time?” Upon 
retrieving successfully, the sea lion was first rewarded with a fish, the 
words “good Penny, and then given the command to mount her 
traming stand were she was given another fish before the next trial. 

The position of the target ring in the tank was varied from trial to 
trial. A grid painted on the bottom of the tank was provided for ease of 
visual monitoring of the ring position and the animal’s search pattern. 
A successful retrieve was one in which the animal brought the rmg 
with her when she emerged from the water onto the platform. If un- 
successful, she was sent again. If repeated unsuccessful attempts were 
made, the sea lion was guided to the correct spot by throwing another 
ring, turning on the lights, etc. Each test consisted of 10 retrievals. 
Constant sound and visual monitoring was maintained throughout 
all tests. - 

The same procedure was followed for both daylight and night trials. 
However, during night trials, tests were run with overhead floodlights 
in the on and off positions. With one 500-watt overhead floodlght on, 
the ambient level directly above the surface of the water ranged from 
0.75 to 1.00 foot candle, as measured with a Weston model 756 illu- 
mination meter. During the “lights off” phase, measurements made 
with the sensing element of the meter placed against the underwater 
viewing ports and on the platform directly above the surface of the 
water indicated ambient light levels of 0.03 to 0.05 foot candle. Dur- 
ing this phase of the test the available light consisted mainly of sky- 
light. These levels were measured with approximately 80% cloud 
cover. Most night tests were made under 100% cloud cover, and hence 
slightly lower illumination. 

Under dark conditions it was apparent that the animal was not 
forced to rely entirely on echolocation, if it did indeed possess such a 
capability. Therefore, attempts were made to completely suppress the 
animal’s vision with blindfolds in a fashion similar to that demon- 
strated by Norris, et al, (1961), with the Atlantic bottlenose dolphin. 
Unfortunately, we could not take full advantage of the blindfolding 
technique since latext suction cups would not adhere to the fur of a 
sea lion in the same fashion as to the smooth skin of a dolphin. After 
several failures with cloth hoods and plastic goggles, a 3-inch wide, 
elastic bandage, 36 inches long, proved to be a successful blindfold. 
This device was wrapped around the animal’s neck, over her eyes, 
and under her muzzle in a double thickness, completely blocking light 
from her eyes, yet not impeding her capability for eating on land or in 
the water or for barking in air. 


170 ~Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


The animal was equipped with a harness and tow line that allowed 
her removal from the tank in the case she became completely dis- 
oriented or displayed some violent fright response. 

The animal was accustomed to the blindfold and to the harness 
separately through the method of successive approximations. She first 
learned to rub the folded blindfold material with her nose and chin, 
then to have the material rubbed over her head. Next, she accepted 
the blindfold being laid over her head and then over her eyes. Finally, 
she allowed it to be wrapped around her head and tied. She was oc- 
casionally given pieces of food during the operation and responded to 
the words “good Penny” by opening her mouth for food. The sea lion 
readily entered the water on command, wearing the blindfold 
wrapped, but not tied, slipped it off and returned. Upon a command. 
she would then retrieve the blindfold for her trainer. Acceptance of 
the harness was similarly accomplished. The animal first learned to 
push the leather with her nose, then to put her head through it. Next 
she learned to present the handler with her flipper to allow the harness 
to be fitted. She accepted both the blindfold and the harness calmly 
and patiently with no display of anxiety. Swimming was not impeded 
by the harness. However, when pressure was exerted on the tow line 
by the handler, the animal appeared to be very emotional. She then 
strained in the harness, dashed about, “porpoised” out of the water, 
and refused to return to her work stand. Some emotional holdover 
appeared occasionally in the trials immediately followmg. Through- 
out the course of these tests the animal did not adjust to being re- 
strained by the line. The complete harnessing and blindfolding pro- 


cedures are illustrated in Figure 2. 


RESULTS 


Underwater Sound Production. After several days of monitoring, it 
became apparent that this particular California sea lion was consider- 
ably less vociferous than a single Atlantic bottlenose dolphin observed 
under similar conditions (Evans and Prescott, 1962). During the 
second week of monitoring, at 1640 hours, the animal was observed 
to produce several bursts of broadband pulses while swimming unim- 
peded a few inches off the bottom of the tank, approximately 15 min- 
utes after a feeding session. These sounds were similar to those pre- 
sumably produced by the California sea lion in its natural environ- 
ment. This type of sound production behavior was observed again on 
subsequent days and was usually accompanied by the same slow swim- 


Sea lion echolocation ability 171 


Figure 2. Various steps in the harnessing and blindfolding of a California sea 
lion, Zalophus californianus (Lesson). 


ming, bottom searching behavior, 15 to 20 minutes after a feeding. 
As observed by Poulter (1963) and Schevill, Watkins and Ray (1963) 
the signals produced by this animal were similar in some ways to clicks 
or pulses produced by delphinids while echolocating. However, in all 
of the pulse series we have observed, the sounds were produced in 


172 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


bursts, rather than in long trains of pulses as are commonly produced 
by the bottlenose dolphin. In addition, the signals produced by this sea 
lion (Fig. 3A) had very little energy above 6 Kcps and none above 8 
Keps, while in those produced by Atlantic bottlenose dolphins (Fig. 
3B) maintained in the same test tank during the early part of 1962, 
considerable high frequency energy was present. No double pulse 
structure, as observed by Poulter (op. cit.) was noted. 


Retrieval Tests. A total of 25 trials was run under the dark condition 
(0.03-0.05 foot candle) ; 18 of these were successful retrievals. The ex- 
perimental animal produced no sound during any of these trials. The 
retrieval time during the “lights on” condition, ranged from 5 to 11 
seconds with an average time of 8 seconds. During the dark condition 
the times ranged from 5 to 22 seconds, with an average time of 15 
seconds. The search pattern used during all test conditions consisted 
of a series of continuous loops. The search pattern was more complex 
during the completely dark condition. There was a consistent tendency 


for the animal to “check” initially the area of the last successful find. 


Blindfold Test. When given the command to enter the water for the 
first blindfold test the animal showed no hesitation. She leaped from 
her training stand into the water, and surfaced in the center of the tank 
(Fig. 2E). When tossed food, she oriented toward the sound of the 
splash, but made no attempt to retrieve the food if it was further than 
2 feet from her mouth. However, food that touched her face or vibris- 
sae was quickly consumed. Occasionally she submerged, sinking slowly 
to the bottom and rising to the surface to breathe. During this first trial, 
when the animal was called back to her traming stand, she swam 
quickly and directly toward her handler, colliding with the edge of 
the platform. She then submerged, floated slowly to the surface, and 
refused to attempt to emerge from the water again. All of her move- 
ments were slow and deliberate. In order to remove the animal from 
the water, slow and gentle pressure was applied to her leash, pulling 
her toward the platform. She was then fed pieces of fish as her han- 
dler unfastened the blindfold. With visual orientation restored, the 
animal voluntarily came out of the water to her training stand. She 
produced no discernible sound during any portion of this test. The ani- 
mal’s behavior during the second blindfold test was similar to that of 
the initial trial, mcluding the lack of sound production. During both 
of the blindfold tests, the animal seemed to prefer staying in the water, 
and at the end of the second test was removed under protest. 


Sea lion echolocation ability 173 


ak i oe 
% ig Vet 


Pe ee f cob ae 
ee Pa i oe 
ors hast oak Bok a jd Dibeeds mauaseed cob ae 


O 0.| O20 OS104 O05) .O6 
TIME (SEC.) 


Figure 3. A. Sonagram of clicks produced underwater by a California sea lion 
maintained in a test tank at the Lockheed-California Company, Burbank Facility. 
B. Sonagram of a train of echolocation clicks produced by an Atlantic bottlenose 
dolphin maintained in the same test tank the previous year. 


174 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


These results, in contrast to those of Norris, et al. (1961) with 
Tursiops, indicated that a total loss of vision seriously impaired the 
sea lion’s navigation and food-finding ability. With continued train- 
ing it is possible that some adaptation to this undoubtedly stressful sit- 
uation might occur. However, it was apparent in these tests that sup- 
pression of the visual sense constituted a more serious problem for the 


sea lion than it did for the blindfolded dolphin. 


Discussion 


The results of this series of tests do not support the theory that the 
California sea lion has a well-developed echolocation ability (Poulter. 
1963). However, much insight ito search behavior, sound produc- 
tion and the visual capabilities of the California sea lion has been 
gained. Also, the techniques developed, and the state of training of 
the animal used, should make it possible to define accurately the vari- 
ous parameters of sea lion target detection behavior, whether they are 
visual, acoustic, or both. The possibility that an echolocation capability 
is a learned behavior, should not be overlooked. 


SUMMARY AND CONCLUSIONS 


1. A single, captive California sea hon, Zalophus californianus (Les- 
son), was observed to produce series of short duration, broadband 
sounds, underwater, while apparently searching for food. These 
sounds appear to be similar to the “clicks” used by the Atlantic bot- 
tlenose dolphin for echolocation signals. 

. The sea lion in captivity does not produce underwater sounds nearly 
as frequently as the bottlenose dolphin, under similar conditions. 

3. Under conditions of darkness (ambient light level, 0.05 foot candle ) 
the sea lion can successfully find and retrieve air filled plastic rings, 
6.5 inches in diameter from 9.5 feet of water, without producing 
sound, 

4. A method was devised to blindfold successfully a California sea lion. 
When blindfolded, the sea lion’s ability to find food and navigate 
was seriously impeded. 


bo 


ACKNOWLEDGMENTS 


We wish to thank Messrs. W. W. Sutherland, R. G. Biel, and J. J. 
Dreher of the Lockheed-California Company for their assistance in 
monitoring during the tests, and Messrs. Arthur Thomas and David 
H. Brown. “Mar ineland of the Pacific Oceanarium, for invaluable sug- 


Sea lion echolocation ability Ais 


gestions on handling and caring for the experimental animal. Also we 
wish to thank Dr. K. S. Norris, Department of Zoology, University of 
California, Los Angeles, Dr. D. K. Caldwell, Los Angeles County Mu- 
seum, Dr. W. E. Schevill, Woods Hole, Oceanographic Institution, and 
Melba C. Caldwell, University of Southern California for their con- 
structive comments on this manuscript. 


LITERATURE CITED 


EVANS, W. E., and J. H. PRESCOTT 
1962. Observations of the sound production capabilities of the bottlenosed por- 
poise: A study of whistles and clicks. Zoologica, 47(2): 121-128. 


KELLOGG, W. N. 

1958. Echo-ranging in the porpoise. Science, 128: 982-988. 

1959. Size discrimination by reflected sound in a bottlenose porpoise. J. Comp. 
Physiol. Psych., 52: 509-514. 


McBRIDE, A. EF 
1956. Evidence for echolocation by cetaceans. Deep-Sea Res., 3: 153-154. 


NORRIS, kK. S., J. H. PRESCOTT, PAUL V. ASA-DORIAN, and PAUL 

PERKINS 

1961. An experimenta! demonstration of echolocation behavior in the porpoise, 
Tursiops truncatus (Montagu). Biol. Bull., 120(2): 163-176. 


POULTER, T. C. 
1963. Sonar signals of the sea lion. Science, 139: 753-755. 


SCHEVILL, W. E., and BARBARA LAWRENCE 
1956. Food-finding by a captive porpoise ( Tursiops truncatus). Breviora, 53: 1-15. 


SCHEVILL, W. E., and W. A. WATKINS 
1962. Whale and porpoise voices, a phonograph. Woods Hole Oceanographic 
Instit. Cont. No. 1320. 


SCHEVILL, W. E., W. A. WATKINS, and CARLETON RAY 
1963. Underwater sounds of pinnipeds. Science, 141: 50-52. 


THE NUMBER OF CONENOSE BUGS, TRIATOMA, INFECTED 
BY ONE ENGORGEMENT ON A MOUSE WITH TRYPANO- 
SOMA CRUZI. In order to test the capacity of one small rodent to 
infect Triatoma p. protracta (Uhler) with trypanosomes, a & 11.5 
em. Mus musculus, Expt. 226, was inoculated intraperitoneally on 
Feb. 1, 1960, with 0.05 ml. of sodium citrate solution contaiming the 
No. 1 dropping of a recently-fed 5th instar Triatoma nymph from 
Griffith Park, Los Angeles (Wood, Exper. Parasitol., 10:356-365, 
1960.) This clear fecal droplet revealed numerous metacyclic Try pa- 
nosoma cruzi Chagas. ‘Tail blood samples of the mouse were negative 
for trypanosomes 14, 16, and 18 days after inoculation. Five trypa- 
nosomes were seen on the 22nd day and one on the 23rd day after in- 
oculation in single drop samples. 

Experimentally infected rodents have been used many times for 
transferring trypanosomes to Triatoma (Wood, loc. cit.). No attempt 
was made here to restrict these bugs to a specific area of the rodent 
for feeding. All bugs were released at the same time in the immediate 
vicinity of the mouse which was confined to a wire cylinder in a cir- 
cular plastic dish, 15.5 cm. in diameter and 7 cm. deep. On the 23rd 
day, two sets of bugs were fed on the experimental mouse. The first set 
of 11 Triatoma (3 3,5 2, 3 fifth instar nymphs) were placed with 
the white mouse for 2 hours. The adults fed to capacity but the 5th 
nymphs were 1; full. These bugs were purposely removed to pre- 
vent further decrease of blood volume in the mouse. They were exam- 
ined for trypanosomes one month later and all except 1 were posi- 
tive, an infection rate of 81.9%. 

The second set of 28 Triatoma (9 third and 9 second instar 
nymphs) were allowed to feed at random until death of the mouse. 
Twenty-four contained blood at the time of removal. These bugs were 
also examined for trypanosomes after one month. Twenty-three or 
82.1% were infected including 8 third and 15 second instar nymphs. 

Since the size and weight of an important California reservoir ro- 
dent, Peromyscus truei gilberti (J. A. Allen), is similar to that of Mus 
musculus, this experiment indicates the percentage of insect vectors 
that could ingest Trypanosoma cruzi during two feedings. This high 
rate of replacement of infected bugs and continued susceptibility of 
Triatoma to infection would explain the widespread distribution and 
continuous recurrence of Chagas’ trypanosome in native reservoir ro- 
dents and hematophagous bugs as demonstrated by field sampling 
(Wood, Bull. So. Calif. Acad. Sci., 41:61-69, 1942. Amer. J. Trop. 
Med., 23:315-320, 1943; Bull. So. Calif. Acad. Sci., 49:98-100, 1950; 


176 


Infection of conenose bugs 177 


Wood and Wood, Amer. J. Trop. Med. and Hyg., 10: 155-165, 1961 ) 
throughout the southwestern United States.—Sherwin EF Wood, Life 
Sciences Department, Los Angeles City College, Los Angeles 29, Calt- 
fornia. 


BIRDS AND INDIANS IN THE WEST 


Love MILLER 


University of California, Davis 


From time to time during the previous ten years I have been asked to 
identify the bird remains taken from Indian middens im western 
America during careful explorations by trained anthropologists. In 
addition there have come to hand numerous bones retrieved by the 
casual digger (including myself) who was inspired only by a simian 
or Juvenile curiosity. It seems worth while now to survey and draw 
together the results of these studies as an addition to the splendid work 
done by Howard (1929) on the great “Shell Mound” at Emeryville 
on the east shore of San Francisco Bay—a site now blotted out by the 
relentless wheels of civilization called “Progress?” 

My own studies have focused carefully upon middens located widely 
as follows: The Dalles, Oregon; several shell-heaps on Puget Sound, 
Washington; four sites along the Missouri River in North Dakota; 
Wilson Butte Cave, Birch Creek Valley, and Weiss Rock Shelter in 
Idaho; Iron Gate site in Siskiyou Co., California; San Nicolas Is. and 
shore sites along the southern California coast. These sites represent a 
great variety of terrain as well as a great variety of Indian tribes—each 
with its special language, food habits, totems, intra-tribal organization 
and religious beliefs. The avian remains from the middens of Oregon, 
Washington, North Dakota and Wilson Butte Cave, Idaho have been 
previously recorded (Miller, 1957, 1960, 1961, and Gruhn, 1961, 
respectively). Those from the Birch Creek Valley and Weiss Rock 
Shelter sites in Idaho are discussed herein. 


ACKNOWLEDGMENTS 


More recently much aid and encouragement have come to me from 
the staffs of the Museums at Idaho State College at Pocatello, Idaho; 
the Peabody Museum of Harvard University at Cambridge, Mass. ; the 
University of Oregon at Eugene, Ore.; and the University of Califor- 
nia at Berkeley. Dr. C. S. Cressman, Dr. Earl H. Swanson, Dr. Alden 
MH. Miller, Dr. W. Wood, Dr. B. Robert Butler, and Dr. Ruth Gruhn 
all have rendered especial service and are extended my sincere thanks. 


178 


Birds and Indians in the West 179 


PreviousLy UNPUBLISHED AVI FAUNAS 


Weiss Rock Shelter (Idaho County, Idaho) 


Buteo jamacensis, Red-tailed Hawk, femur and humerus. These two 
bones represent a large Buteo (probably a single individual and a 
female). They are not B. regalis, bemg too heavy shafted and short. 
Both are much abraded at the ends giving a first impression of having 
been gnawed by small rodents. On close examination, however, the 
humerus shows on the palmar aspect of the area between the bicipi- 
tal and the deltoid attachments some rectilinear scratches, two of 
which cross each other and are followed by two parallel lines to for- 
tuitously produce the Roman numeral XII. Other lines on both sides 
of the deltoid crest strongly suggest that a very fine pointed instru- 
ment had been used to remove the crest. Furthermore on the anconal 
face opposite the distal end of the deltoid crest a hole 10.2 x 6.5 mm. 
in size, has been made. A few faint scratches are discernible at the 
antero-distal margin of this hole. The impression in my own mind is 
that an attempt had been made to produce a whistle but the plan had 
been abandoned before completion. I have examined such whistles 
from other middens but they are generally made from the ulna of 
longer winged birds. A 3 mm. hole appears in the shaft of the femur 
which I can not explain though how it could have been made by ac- 
cident without breaking the shaft is a mystery. No tool marks appear 
on the femur. Both bones show weathering to greater extent than any 
other specimen in the collection. 


Falco sparverius, Sparrow Hawk. Humerus and tibia are the only 
bones of this species retrieved thus far and fortunately their preserva- 
tion is excellent. The left humerus lacks a few chips from the deltoid 
crest, the right tibia lacks the proximal 1/4. The bones show no weather- 
ing and are but lightly stained. Rich color of the tail feathers of this 
little falcon may have appealed to the Indians as ornaments. 


Dendragapus obscurus, Dusky Grouse. This grouse is the most promi- 
nent bird in the collection and the majority of the bones are beauti- 
fully preserved. Specimens vary in color from “old ivory” to a warm 
golden brown. Both sexes of adults and two stages of immaturity are 
represented. In the John Day Basin of Oregon I found this species 
quite abundant in sage country with scattered low junipers among 
gently rolling hills. It is surprising that only two of the nine bones 
are definitely from young birds. 


180 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


Pedioecetes phasianellus, Sharp-tailed Grouse. This grouse is repre- 
sented by a single tarsus, presumably of a female bird. Both ends are 
broken but it closely corresponds except for size with an old male, the 
only recent skeleton available. This grouse is smaller and definitely 
more slender legged than Dendragapus. There appears no possibility 
of confusing the tarsi of the two birds. 


Otus asio, Screech Owl. An incomplete ulna is assigned to the Screech 
Owl after repeated check and re-check under the lens. Only the proxi- 
mal 24 remains and the olecranal process is missing. The size, curva- 
ture, intermuscular lines and facets so far as preserved correspond 
with those of a female specimen of O. asio bendirei from central Cali- 
fornia. The fascinating thing about this specimen is that a tiny round 
hole has been drilled in the shaft on the concave (radial) side. This 
hole is less than 1.5 mm. in diameter and appears perfectly round with 
smooth edges. The distal end of the specimen does not appear to have 
been broken. I am confident that it was purposefully cut as was done 
in removing the olecranal process at the proximal end. Was the 
skilled artisan endeavoring to produce a tiny whistle? Several of the 
whistles examined from other Indian middens had been made from 
the ulnae of long winged species (e.g. Grus), the shaft and the ole- 
cranal process had been cut in similar fashion and the hole located on 
the concave aspect of the shaft. Whatever his purpose the Indian did 
a skillful bit of work. 


Asio otus, Long-eared Owl, tibia and humerus. ‘Iwo species of the 
genus Asio might be expected to occur at times in the area of Weiss 
Rock Shelter. Isolated and broken bones of unknown sex of these two 
owls are difficult to assign specifically. The tibia is fortunately quite 
nearly perfect so it is assigned with confidence to the species A. otus. 
The other specimen is a badly broken humerus which is assigned with 
less confidence to the same species. The Long-eared Owl seems the 
species most likely to be found in country not too far from the rock 
shelter, In my own field experience it has been met in fair numbers 
in sage-juniper country as well as in localized timber along streams 
that cut through semi-desert areas. 


Colaptes cafer, Red-shafted Flicker, two humeri and tarsus. The tar- 
sus and one humerus are practically perfect and are readily identifi- 
able. The second humerus, #42,436, is less perfect but corresponds in 
such characters as are preserved although the shaft is slightly heavier, 
a difference assigned to age or sex of the individual. The congeneric 


Birds and Indians in the West 181 


Yellow-shafted Flicker is less likely to occur in this region and com- 
parison of humeri of specimens of the same sex from the two species 
showed a stouter shaft in the Red-shafted Flicker. It seems proper 
therefore to consider C. cafer as the species represented. Although I 
have known the Flicker to serve as a food item for a number of Cau- 
casians and certainly could appreciate such an interest on the part 
of an Indian, I feel that the feathers of this beautiful bird may have 
had an even stronger influence in making it one of the species oc- 
curring frequently in the bird lists from Indian middens. 


Corvus corax, Raven, coracoid. This practically perfect bone is the 
only representation of this species which occurs in so many other 
middens and cave deposits. It is wide-ranging in open arid country 
where it commonly uses rocky outcrops for nesting sites. The single 
bone came from a mature individual. 


Corvus branchyrhynchos, Common Crow, pelvis, humerus, ulna. The 
Crow is less commonly seen in open country than is the Raven but two 
and probably three individuals are represented in the collection. I 
feel that a sizable assembly of broad leaf trees furnished attraction for 
these gregarious birds somewhere within reach of the Indians of Weiss 
Rock Shelter. Audubon in his Missouri River Journal, speaks of cer- 
tain tribes of Indians along the River as using the crow for food. Such 
use may explain the abundance of its bones in this Idaho midden. 


Pica pica, Magpie, nine bones. I assume that the Magpie here repre- 
sented is the widely occurring Black-billed species since our only other 
species, the Yellow-billed Magpie, is so closely restricted to limited 
areas and habitats in central California west of the Sierra Nevada. Con- 
stant osteological differences between unassociated bones of the two 
species have not been recognized. 

Only one other species, the Dusky Grouse, is represented in such 
great numbers and no immature specimens have been found. Further- 
more, I have found the adult Magpie a wary bird and quite able to 
take care of himself unless his curiosity be excited. I am impressed by 
the Indians’ ability in capturmg numbers of the birds. The species may 
have had a varied appeal—as food, as pets, as a source of attractive 
feathers or as a tribal totem. 


Turdus migratorius, American Robin. A single bone, a complete tibia 
represents this species that, durig one or another season of the year 
may be found in any type of environment below the limit of trees 
from the arctic to Mexico. Furthermore it may occur, at times, in great 


182 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


numbers during the non-breeding season. Its plump body would have 
been an acceptable food item at the camp fire of any group. Its rarity 
and its perfect preservation are the chief interests attached to this 
specimen. 


Sturnella neglecta, Western Meadowlark. Like the Robin, this full 
bodied bird may have added an acceptable item to the Indians’ bill of 
fare. Its presence indicates open country though not necessarily a per- 
manent meadow. Semi-arid country with annual grass dotted with 
low, perennial bushes may be inhabited by Meadowlarks. 


Birch Creek Valley CAvE on Rock SHELTER 
(ca. 125 NNE of Pocatello, Idaho 


Anseriformes, a single coracoid of a medium sized duck represents this 
order. Seemingly the water birds played a very small part in the life 
of the Indians of this area. There must have been no still water or 
marshes of any size within hunting range, else their arts, crafts, cere- 
monials or nutrition would have included some species of water bird 
in greater numbers. 


Falconiformes, a complete and a broken coracoid may have come from 
the same individual of Falco columbarius, the Pigeon Hawk, which 
alone represents the raptorial order. In none of the Indian middens 
previously studied have I failed to find some large species of raptor; 
eagles, hawks, falcons and even vultures. The Birch Creek environ- 
ment surely provided several species of these birds. Did they have no 
attraction for the Indians? With what plumes did they feather their 
shafts? Grouse feathers are too strongly curved. Crows are a bit too 
small for efficient use and large waterfowl seem not to have been 
available. 


Galliformes, one’s first impression is that the whole domestic economy 
of the Birch Creek Indians centered about the grouse family and that 
the only grouse was the Sage Hen. Closer study, however, shows that 
two other grouse and a quail (Oreortyx) played a minor part for the 
Galliformes. 


Dendragapus obscurus, Dusky Grouse, is represented by three hu- 
meri, one femur, one tibia, one tarsus and two coracoids. The male 
humerus is very closely like that of the female Sage Hen, but the shaft 
is stouter and the distal articulation with the forearm operates around 
a more oblique axis, 7.e. the ulnar condyle and the entepicondyle are 


Birds and Indians in the West 183 


extended more distally than the radial condyle. Similarly the femora 
are much alike except that Dendragapus shows a greater degree of 
pneumaticity in the great trochanter when viewed from the anterior 
face. This grouse is found in open timbered country and pine forest 
today, contrasting with the Sage Hen of the treeless sage. 


Centrocercus urophasianus, Sage Hen. This grouse is the dominant 
bird of the entire collection. Forty six (46) humeri, equally divided 
between male and female, are identified with confidence whereas 
other skeletal elements are strangely in the minority—skull 1, sternum 
1, scapula 1, fibula 1, tarsus 1, femur 2, sternum 2, tibia 7, metacarpus 
none at all. There are a few ulnae present, but they are so badly muti- 
latted that they can be assigned merely to “grouse?” Why are the ulnae 
broken away at both ends while many of the humeri are quite per- 
fect? Seven (7) coracoids are assignable to “grouse,” but only two are 
assigned with confidence to Dendragapus and none to Centrocercus. 

The super-abundance of this species emphasizes the openness of 
Birch Creek Valley and the presence of zeric vegetation. 


Bonasa umbellus, Puffed Grouse. Three grouse metacarpi, per- 
fectly preserved, check perfectly with this small grouse. No other bone 
in the entire collection, however, is thus assignable, nor are there other 
Western grouse known to me with which they could be confused. They 
came from adult birds of both sexes. 

Presence of Ruffed Grouse remains in these small numbers suggest 
a limited amount of stream-side tangle of water loving trees, brush 
and vines. 


Oreortyx picta, Mountain Quail. A single coracoid is the sole rep- 
resentative of this species. It is found today in ecologic association with 
Dendragapus rather than with the open country dwelling Centro- 
cercus. 


Strigiformes 
Asio otus, Long-Eared Owl. There are two bones in the collection 
which are assigned to the owls. One is the distal end of a humerus, the 
other an almost complete tarsus of a bird scarcely out of the nest. Both 
are assigned to the species A. otws; the Short-Eared Owl, Asio flam- 
meus, is very Closely similar but in general it is shghtly larger. It must 
be admitted that the specific assignment may be in error. 
Long-Eared Owls today may be found breeding in mesquite desert, 
pine clad mountain canyon or willow thickets along river courses. 
The bird’s presence here holds little interest except to raise the ques- 


184 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


tion “why?” The immaturity of the bone suggests that a young bird 
may have been captured and brought in by a child or by a medicine 
man. It may even have been held in captivity for a while or used in 
ceremonial rites. Some Pacific Coast tribes consider the owls as in- 
carnations of nocturnal spirits of mystic powers. 


Passeriformes 

The remainder of the collection is assigned with a fair degree of con- 
fidence to this order though some of the badly broken bones could be 
those of the flicker, a species so often found in Indian middens. Many 
of the most delicate little bones are as perfectly preserved as though 
prepared by a skillful technician. These latter, I feel confident, reached 
Birch Creek in the stomachs of owls, to be later cast up unbroken and 
beautifully cleaned, as pellets of fur, feathers and bone. The owls, 
however, seem to have used the area only as temporary roosts since 
they left none of their own bones as record. (Aszo does not inhabit 
caves or rock shelters. ) 

The owls can scarcely be held responsible, however, for the presence 
of the wing bones of certain larger passerines which could not be 
swallowed whole but would have been partially dismembered. 


Corvus brachyrhynchos, Common Crow. The crow is represented 
by seven of the characteristic carpometacarpi and by no other bone. 
Certainly no owl could swallow the terminal joint of a crow’s wing 
(owls do not pluck their prey). This species must have been brought 
in by man and, though divers Indian tribes are reported to have used 
the crow as food, the presence of wing bones only is suggestive of ritual 
or decoration as the purpose served in this case. 

For some strange reason these segments of the crow’s wing are all 
broken at either or at both ends though the slender metacarpal II 
may be well preserved. Also the size of some may reach that of a fe- 
male C. cryptoleucus although that species shows positive divergence 
in osteological characters. Possibly the crow’s primary feathers were 
used by the arrow marker for want of better. This might account for 
fracture of the metacarpus in chopping off and bringing home only 
the wing-tips. 


Pica pica, Black-billed Magpie. Like the crow, the magpie has been 
found in many of the Indian middens of the west. I have ascribed its 
presence to the Indians having used its striking plumage in ornamenta- 
tion or having kept it as a pet. Two humeri, one metacarpus and a ques- 
tionable ulna represent it in the Birch Creek collection. 


Birds and Indians in the West 185 


Smaller passerines. Quite a few of the bones mentioned above as 
probable remnants of “owl pellets” are left nameless. Classification 
within the order is based so largely upon characters not registered in 
the single and unassociated bone that I feel much uncertainty as to 
even the family represented. An error once registered upon the printed 
page can never be erased. Therefore, silence seems to be indicated. 


AGE oF MIDDENS 


Materials from many areas that have passed through my own hands 
represent the birds presumably assembled by Indians who had not yet 
come under the influence of Caucasian man; J.e., he was himself a 
“natural species” of wild creature among wild creatures—more or less 
omnivorous but, when possible, in the main a carnivore. Unlike the 
bear, wolf or cougar, however, he had a primate brain that set him 
quite apart from his fellow creatures. His aesthetics, religion, cere- 
monials, his social organizations, interest in pets or games, craftsman- 
ship in clothing, housing, weapon-making and I know not what else, 
all had an influence upon the accumulation of bird bones aside from 
his mere desire for food, though the latter may have loomed large. 

The time represented in these accumulations extends from the near 
present back into that nebulous borderland between the Pleistocene 
and the Recent. At least the horse, camel, mammoth and ground sloth 
survived long enough to “shatter lances” with the big braied bipedal 
nmmigrant from the Old World. Carbon 14 dating accomplished thus 
far extends the period back some 10,000 to 12,000 years (Butler, 
1962) 

In only one case so far have I found an extinct species of bird im 
the midden collections. Among 9,000 bird bones there was a single 
bone of Coragyps occidentalis from the Dalles, Oregon. Other extinct 
species have been reported by my colleagues from caves in New Mex- 
ico and Texas, but the actual association with man has not yet been 
established. 

The association of the extinct Diving Goose (Chendytes) with man 
im the coastal zone of Southern California is also uncertain thus far 
(see Howard, 1949:24). The California Condor (Gymmnogyps cali- 
fornianus), so near extinction but so widely known in late Pleistocene, 
is extremely abundant in the midden at the Dalles, Ore. on the east- 
ern side of the Cascade Range. The Whooping Crane (Grus ameri- 
cana) is but sparsely represented in middens of North Dakota. Notably 
absent from the Dakota middens are the Passenger Pigeon (Ectopistes 
migratorius) and the Carolina Parakeet (Conuropsis carolinensis) al- 


186 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


though Audubon records the latter within a very few miles of the 
Dakota site. 

Aside from the Condor, no species of bird has been identified in these 
studies at a site outside its present day range. 


ARTIFACTS 


The almost ivory-like texture and the highly pneumatic nature of 
certain groups make the bird’s bone attractive to the Indian for several 
purposes. The long bones of deer and mountain sheep approach the 
fine texture of bird bone and I have seen large awls made of them, but 
for more delicate sewing the bird bone serves best. In the North Da- 
kota middens there were well polished awls made from Swan: ulna 1. 
Eagle: ulna 2, radius 4, tibia 1; Crane: radius 1; and Horned Owl: 
humerus 1. The eagle bones were particularly desirable, it would 
seem, because of the lesser degree of pneumaticity. These bird awls 
were, I imagine, used in the making of finer clothing and headdresses 
out of thin skins of small mammals and birds. 

This same fine texture and pneumaticity made the long bones of 
birds useful in fashioning beads and as counters in certain games. The 
radius of larger birds, when cut into segments, made beads already 
perforated for stringing. The fine texture would appeal to the sense of 
touch. 

The best Indian ‘“‘whistle” I have seen was made from the ulna of 
a crane (Grus canadensis ). Both ends had been removed and a smooth 
hole cut near the proximal end on the concave (radial) side. Within 
the cavity of the bone, just below this hole, a partial plug had been 
skillfully modeled to direct a blast of air out through the hole and 
produce a musical tone of great purity closely approximating that of 
the California Pigmy Owl (Galucidium gnoma). Fortunately the 
artisan had used a somewhat asphaltic matrix which had not been re- 
moved by subsequent ground waters so his skillful handiwork had 
been preserved. The length of this whistle was 614 inches. Another 
ulna from this same midden (on San Francisco Bay Shore) had been 
squarely cut at one end though it was irregular at the other. Unfor- 
tunately the plug, if ever present, had not been preserved; but if a 

thumb closed the rough end, one could blow across the smoothly-cut 
end and produce a good tone much as a small boy would blow across 
an empty cartridge case or a pill bottle. This bone was 21% in. long. 
This midden yielded several other fragments of long bone that might 


Birds and Indians in the West 187 


have been designed for making into whistles by a specially music 
minded craftsman. 

Men (or women) of all times and tongues have yielded to the at- 
traction of the feather. Even the bird himself seems in some species 
to be appreciative of his own colorful clothing. The Indian was no 
less susceptible than his kinsman im Hawaii, Africa, New Guinea (or 
New York). So he decorated his headdress, his medicine bag, his 
quiver and I know not what else with feathers. ‘To this aesthetic sense 
I ascribe the presence of several species of bird remains we find in 
Indian middens. Of course the feather itself may have long ago dis- 
appeared, though there are exceptions (I have seen the gaudy feathers 
of macaws preserved for centuries in the dry air of Peru). Sooner or 
later certain of the species might have become totemic either to a tribe 
or to a clan within the tribe. This might lead to a tabu on the one hand 
or to the ritual consuming of its flesh, blood or heart on the other hand. 

Mrs. Joseph Grinnell gives me an interesting account of her hus- 
band’s parents’ experience during the 1870’s in the midwest. Father 
Grinnell was government physician to the Indians in Indian Territory. 
Wild Turkeys abounded in the timber along the water courses whence 
they were frequently brought in the doctor’s game bag to the kitchen 
door. Mother Grinnell went into the back yard to pluck them. There 
she often found eager little brown hands thrust between pickets of the 
fence to clutch the colorful feathers but never could the Indian be in- 
duced to touch any of the meat. It was tabu to the Indians of that area. 
Indians of the cliff-dweller culture kept turkeys in captivity for the 
sake of their feathers, it is reported, rather than as a source of food. 
Powell, in his account of exploration in the Colorado River Basin, 
figures an eagle cage built and used by Hopi-land Indians. I saw a 
magpie kept in a cage by a wandering tribe of Indians in Death Valley, 
California. What but its plumage could have been the attraction in 
this case? 

I feel that attractiveness of their plumage may account for the pres- 
ence of Sparrow Hawk, Flicker, Magpie and perhaps Crow in many 
of the middens that I have examined. It has been reported that the 
scalps of the Acorn-eating Woodpecker in California and the Ivory- 
billed Woodpecker of the Mississippi Valley were used within historic 
time as a medium of exchange between tribes. I am inclined to credit 
the report. 

The wing feathers of large raptors, waterfowl or even the Raven 
probably found use in the fletching of arrows. Skins of pelicans made 
warm body covering. The fluffy plumes of the Rhea, held together by 


188 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


plant fibre made a ight and comfortable blanket for the native of the 
Argentine pampa. Altogether the bird’s feather was an important 
item of interest to the Indian. ‘To me it explains the presence of cer- 
tain bones in the refuse heaps. Among some ten thousand bird bones 
examined, I have seen but two or three that suggested contact with 
fire. Possibly these Indians, like some Esquimaux, ate the bird raw. 


RITUAL 


The owl was sacred to Pallas Athena; the Romans ascribed great sig- 
nificance to the movements of birds. No less seriously did the Ameri- 
can Indian look upon a wide variety of birds. This aspect of ethnology 
warrants a volume in itself. Only surmise is left us in the work with 
midden remains. How tantalizing and how exciting to the imagina- 
tion! The Indians, more than likely, brought with them from the Old 
World a “rootstock” of belief or superstition from which there grew 
through the ages, along with the evolution of linguistic or even phys- 
ical characters, a whole “botanical garden” of beliefs, rituals, tabus. 
totems and heraldry. The picture is so kaleidoscopic today that I ven- 
tured only suggetsion in the discussion of certain species. 


FRAGMENTATION OF BONES 


Three papers have previously appeared regarding my own study of 
the bird remains from middens (Miller, 1957, 1960, 1961). In each 
of these comment was made on the high degree of breakage. This con- 
tmues to puzzle me. Mammal bones were supposedly cracked to re- 
trieve the yellow marrow from the cavity. Bird bones are small and 
in relatively few species is there even a small amount of fat to be found 
—certainly not enough to encourage an Indian to crack the dense and 
very hard tarsus of an eagle or a condor. Yet seldom are these bones 
found unbroken. 

Trampling by bare or moccasined feet would merely press the bone 
into the soft matrix of the midden. Bone-chewing habits of the dog 
family would tend to concentrate upon the ends of the bone where 
articular cartilages and synovial membranes add flavor. Furthermore 
the bone would be abraded instead of merely broken. Excavation of 
the Dalles midden uncovered almost no canid material whereas the 
breakage of bird bone there was most marked. 

On San Nicolas Is. off Southern California, dog remains of two 
strains were extremely abundant; yet im the extensive middens there 
breakage of bird bone was at the minimum. The Birch Creek middens 


Birds and Indians in the West 189 


in eastern Idaho show less of this breakage than does any other main- 
land midden that I have studied. To me it seems to be the result of 
human activity, a force sometimes quite powerful, that was purpose- 
fully apphed. But what was the purpose? Was it ritual? In the case of 
the Sage Hen from Birch Creek midden, which was presumably a food 
item in the main, many of the long bones were almost perfect. 


SUMMARY AND DEDUCTIONS 


1. Birds were of definite interest to the Indians though that interest 
may have varied greatly. 

2. The interest may have been in its flesh as a food item; in its feathers 
for ornament, for flectching the arrow, for warmth in clothing, for 
symbolism; in its bone for making awls, beads, counters or whistles; 
as a tribal or clan totem; religious or ceremonial sacrifice; as a pet; 
Tabu probably loomed large. 

3. The bird fauna retrieved from a midden varied with geographic lo- 
cation, with local ecology, with climatic changes at the same site. 

4. Only one bone out of many thousand represents an extinct species. 
Only one species, the condor, was found outside its present day 
range. 

5. In some cases mineralization had begun. 

6. Maximum age was 10,000 to 12,000 years. 

7. Bones from most middens were highly fragmented. This phenom- 
enon was not explained. 

8. Owls or falcons may have added some bones to a midden. 

9. Falcons, Ravens and Cliff Swallows may have nested in some of the 
caves. 


190 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


TABLE I. 


Bird remains identified from middens of Western Indians. 


xX —= present 

sp. = species 
Gaviidae 
Podicipedidae 
Diomedeidae 
Pelecanidae 
Phalacrocoracidae 
Ardeidae 
Threskiornithidae 
Anatidae 


Cathartidae 
Accipitridae 
Falconidae 
Tetraonidae 
Phasianidae 
Gruidae 
Rallidae 
Charadriidae 
Scolopacidae 
Recurvirostridae 
Phalaropidae 
Laridae 
Alcidae 
Columbidae 
Tytonidae 
Strigidae 
Picidae 
Hirundinidae 
Corvidae 
Turdidae 
Icteridae 
Fringillidae 


(Howard, 1929) 


x many 
sp. 

x 2 sp. 

x +4 sp. 

x 3 sp. 


Oregon 
(Miller, 1957 ) 


Dalles, 


= 
i 
a) 
ron) 
= 
. ‘Ef 
K 3 
~: = as 
SEg= 329 
Syed eb Tr 
-~o sow 
case SA 
x 2 sp. = 
2 sp. = 
x x 
- x 
x many x3 sp 
sp. 
x 2 sp. x3 sp 
x x 
x x 2sp 
>.< —. 
as = 
> 4 — 
x x 
x x 2sp 


(Miller, 1961 ) 


Wilson 


“mM 


~ Kw mK 


Butte, 


Idaho 


(Gruhn, 1961 ) 


Weiss 
Rock, 
Idaho 


Birds and Indians in the West 191 


LITERATURE CITED 
ADUBON, J. J. 
1843. The Missouri River Journal. Reprint by Dover Publ. Co. 


BUTLER, B. R. 
1962. Contributions to the prehistory of the Columbia Plateau. Occ. Pap. Idaho 
State Coll. Mus., 9: 1-86. 


GRUHN, RUTH 
1961. The archeology of Wilson Butte Cave, South Central Idaho. Occ. Pap. 
Idaho State Coll. Mus., 6: 1-202. 


HOWARD, H. 

1929. The avifauna of Emeryville Shell mound. Univ. Calif. Publ. Zool., 32: 
301-394. 

1949. Avian fossils from the Marine Pleistocene of Southern California. Condor, 
51: 20-28. 


KROEBER, A. L. 
1925. Handbook of the Indians of California. Smithson. Inst., Bull. Bur. Amer. 
Ethnol. 78: 1-995. 


MALKIN, B. 
1962. Seri Ethnology. Occ. Pap. Idaho State Coll. Mus. ,7: 1-58, with appendix. 


MILLER, L. 

1957. Bird remains from an Oregon Indian midden. Condor, 59: 59-68. 

1960. Some Indian midden birds from the Puget Sound area. Wilson Bull., 72: 
392-397. 

1961. Bird remains from Indian middens in the Dakota area. Bull. So. Calif. 
Acad. Sci., 60: 122-126. 


POWELL, J. W. 
1895. Exploration of the Colorado River and its canyons. Reprint by Dover Publ. 
Co. 


SWANSON, E. H. 
1961. A preliminary report on the archaeology in Birch Creek, Eastern Idaho. 
J. Idaho State Coll. Mus., 4: 25-28. 


CONTRIBUTIONS FROM THE LOS ANGELES MUSEUM— 
CHANNEL ISLANDS BIOLOGICAL SURVEY. 371. 
BRACHYDONT DESMOSTYLIAN FROM MIOCENE OF 
SAN CLEMENTE ISLAND, CALIFORNIA. 


Epw. D. MrrcHext, Jn. 


Los Angeles County Museum 
and 


University of California, Los Angeles 


The twelfth expedition of the Los Angeles County Museum’s Channel 
Islands Biological Survey visited San Clemente Island, Los Angeles 
County, California, for one week during February, 1941 (Comstock, 
1946: 100). On the last day of the stay, three teeth and a bone frag- 
ment of an extinct mammal were collected by the party, possibly in 
the vicinity of Wilson Cove on the northeast end of the island. These 
teeth represent a comparatively rare brachydont desmostylian. 

Desmostylians were large, supposedly amphibious mammals which 
inhabited both the east and west margins of the North Pacific Ocean 
during the Miocene Epoch. There were probably a number of differ- 
ent types, but the best known and one of the most widely distributed 
genera, Desmostylus, is still considered by many to be a poorly known 
mammal, and its relationship to other desmostylian species can only 
be regarded as tentative. Each cheektooth of desmostylians is made up 
of a number of columns that are joined at the base. In Desrmostylus, the 
height of the columns is great and the base of the crown is not en- 
larged, but in two other genera, Cornwallius and Paleoparadoxia, the 
columns are relatively shorter and the base of the crown may or may 
not form a predominant part of the tooth. 

The following abbreviations are used in this report: CIT, Califor- 
nia Institute of Technology collections now in the Los Angeles County 
Museum; LACM, Los Angeles County Museum; PMBC, Provincial 
Museum of British Columbia; and UCMP, Museum of Paleontology, 
University of California, Berkeley. 


‘All of the previous contributions from the Channel Islands Survey have ap- 
peared irregularly in this Bulletin, beginning in 1939. 


192 


Miocene brachydont desmostylian 193 


Order DESMOSTYLIA Reinhart, 1953 
Family Paleoparadoxidae Reinhart, 1959° 
Paleoparadoxia sp. 


Material. Four specimens are represented in the collection: LACM 
4371a, a nearly complete cheektooth; LACM 4371b, crown and por- 
tion of the root of a premolar embedded in a bone fragment showing 
portion of an adjoining alveolus; LACM 4371c, a bone fragment with 
the basal end of a tusk in place; and LACM 4371d, an indeterminate 
bone fragment (Fig. 1a-1). 


Collector. The specimens were collected by Jack C. von Bloeker, on 
22 February 1941. 


Locality. LACM fossil vertebrate locality 1164: “Wilson Cove, north 
of airport, San Clemente Island, Los Angeles County, California?’ This 
information is from the original label; no further data are available. 
Neither Mr. von Bloeker (the collector) nor Mr. George P. Kanakoff 
(who accompanied Mr. von Bloeker on the collection date) can re- 
member specifically collecting the specimens. Mr. von Bloeker thinks 
that they were collected near the pier at Wilson Cove on the above 
date, but Mr. Kanakoff believes that they may have been collected else- 
where on the island by U.S. Navy personnel. The rock adhering to the 
specimens was a light brown, medium-grained sandstone composed of 
angular to subangular clasts of volcanic rock. Rocks of similar lithol- 
ogy were not located by Mr. Jere H. Lipps and me during a search at 
Wilson Cove on 5 September 1961. The specimens may not have 
come from Wilson Cove. 


Formation and Age. Olmsted (1958) indicated that the unnamed 
Tertiary sedimentary rocks exposed on the island are middle Miocene 
in age. However, he stated (p. 65) that the rocks at Wilson Cove con- 
tain “phosphatized Foraminifera that are questionably of early Mio- 
cene age.’ 


Descriptions. The four specimens hereinafter described may have 
come from a single collecting station, and possibly from a single skele- 
ton since they were apparently collected im limited time, are from un- 
duplicated parts of a skeleton, all have the same patina, similar weath- 

*Designated “Family Paleoparadoxia” by Reinhart (1953), but was a nomen 


nudum until 1959 when the typical genus (Paleoparadoxia Reinhart, 1959) was 
proposed and characterized. 


194 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


ering marks, and had the same type of matrix adhering to them be- 
fore preparation. 


Cheektooth. The nearly complete cheektooth (LACM 4371a) lacks 
the entire root system, and also the pulp cavity wall for about one 
quarter of its circumference. The columns are reduced in size com- 
pared to the underlying inflated crown. There are four main columns, 
with a group of three accessory columns at each end and at least two 
(possibly three, one may have occupied a lateral broken area) small 
columns between the large pair of main columns (see Fig. 1a). The 
columns are set upon a crown which is laterally inflated and bulbous, 
and which has thin walls ventrally. The columns converge toward the 
center of the crown, and each has a small protuberance of dentine at 
the top. No wear is evident on any column. A well defined cingulum 
runs nearly the length of the tooth on one side and seemingly gives 
rise to an additional incipient lateral cuspule on one end (Fig. 1c). A 
ventral view into the pulp cavity (Fig. 1b) discloses seven indenta- 
tions that can be directly correlated with columns (or groups of col- 
umns) of corresponding relative size directly above (asin Desmostylus; 
VanderHoof, 1937: 182). The pulp cavity itself is exceptionally large 
—if the tooth were not broken, exposing a cross-section of relatively 
thin enamel and thick dentine, it might be thought to be the cavity 


TABLE 1 


Measurements in millimeters of crowns of cheekteeth of brachydont 
desmostylians. Comparative measurements taken from VanderHoof, 
1942; Hay, 1924; and Clark and Arnold, 1923. 


Specimen Maximum length Maximum width Maximum height 
; of crown 
Paleoparadozia sp. 
LACM 4371a — 28.0 26.5 
UCMP 45274 93:2, 18.0 16.0 
UCMP 63981 20.0(?) — 13.0 
Paleoparadoxia tabatai 
UCMP 40862 33.0 25.4 20.0 
UCMP 32076 35.4 25.4 16.2 
Cornwallius sookensis 
PMBC 486 (holotype) 33.0 24.0 16.0 
PMBC 491 4.8.5 34.0(?) 23.0 
UCMP 32682 — 28.0 14.0 
UCMP 36079 — 24.0 26.5 


UCMP 36078 36.0 27.5 1725 


Miocene brachydont desmosty lian 195 


occupied by dentine (see cross-sections in VanderHoof, 1937: fig. 14; 
Hay, 1923: fig. 4; Hay, 1924: fig. 1). The ventral border of the pulp 
cavity as preserved is laterally constricted midway from the ends of 
the tooth, which suggests that the tooth may have been double rooted. 
The dentinoenamel junction was probably about 10 mm. below the 
bases of the columns. At the base of a major column (on the broken 
end of the tooth) the enamel is 2 mm. thick. Fine circumferential 
striae (the surface expression of the enamel prism groups of Vander- 
Hoof, 1937: 182; and “Schreger’s stripes” of Reimhart, 1959: 98) are 
distinguishable on the base of the crown and the columns. Somewhat 
larger vertical striae are distributed around the tooth in the region 
between the base of the crown and the base of the columns. A water- 
worn cheektooth (UCMP 64116) found with another tooth referred 
to Paleoparadoxia at UCMP locality V5555 in late Miocene rocks at 
Santa Cruz, California (Mitchell and Repenning, 1963) closely re- 
sembles LACM 4371a in having a well inflated crown base with col- 
umns that converge sharply inward. Dr. Tadao Kamei (in litt. 25 
November 1962), basing his conclusion on a drawing, thinks that 
LACM 437 1a is a left lower third molar. 


Premolar. The premolar (LACM 4371b) is embedded in a badly 
eroded fragment of bone. The remnant of the root of another tooth 
slopes toward the premolar at an angle of 25°. The root of the pre- 
molar is oval in cross-section and is solid. The crown of the tooth is 
thickly enameled. The crown has very small pits irregularly distrib- 
uted over its surface, but these pits may be due to weathering phe- 
nomena. On the tip of the crown is a tear-drop shaped opening in the 
enamel which exposes dentine (Fig. 1d). On one side of the crown 
(Fig. 1e) is a small, flat wear surface. On the side of the crown near- 
est to the second root fragment is a pronounced bulge accentuated by 
a small, irregular pit above it. On the edge of the tooth opposite this 
bulge is a small indentation which may be a second wear surface. 
There is a well marked separation between the enamel of the crown 
and the dentine of the root. Dr. Tadeo Kamei (in litt. 25 November 
1962), basing his conclusion upon a drawing, thinks that LACM 4371b 
is a right first premolar, but that it could be either upper or lower. 


Tusk. The fragment of bone containing the basal tusk fragment 
(LACM 4371c) is poorly preserved, but some information can be 
derived from it. No sutures are evident on the bone. It is dense except 
for the area at the base of the pulp cavity of the tusk, where it is per- 
forated by numerous large nutrient foramina. Only a portion of one 


196 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


Figure 1. Fragmentary teeth and bones of Paleoparadoxia sp. from Miocene of 
San Clemente Island, California. All views 24 natural size except 1 which is 
144 natural size. a—occlusal, b—bottom, and c—side views of cheektooth LACM 
side views of premolar LACM 4371b; f—diagram- 
matic cross section, g—anterior, and h—side views of tusk fragment LACM 
4371c; and i—cross-section of indeterminate bone fragment LACM 4371d (which 


4371a; d—occlusal and e 


has been ground flat on two sides). 


Miocene brachydont desmostylian 197 


side of the tusk is preserved. In anterior view, it is evident that the tusk 
was not perfectly round. In Figure 1g, the dark shading to the right 
of the tusk fragment denotes an area from which most of the tusk is 
broken away, but where a thin veneer of dentine still adheres to the 
alveolar wall. The curvature indicates a wider tusk with a sulcus or 
groove on one side (Fig. 1f). The walls of the tusk itself are thick 
three centimeters anterior to the basal end, but are reduced in thick- 
ness posteriorly to a sharp-edged margin at the end. The pulp cavity 
is funnel-shaped, tapering rapidly anteriorly (Fig. 1h). 


Bone fragment. On the most convex border of the indeterminate bone 
fragment (LACM 4371d) there are numerous interconnecting shal- 
low sulci which run along the axis of the bone. The broken surfaces 
of this bone were cut and polished on a lap wheel to better observe the 
microstructure (Fig. 11). The bone is quite dense. The sections show 
the characteristic pachyostoseal condition (sensu McLean and Urist, 
1961: 226) that VanderHoof found in Desmostylus and which I be- 
heve to be characteristic of the order Desmostylia (Mitchell, Jn press). 
The Haversian canals are distributed regularly throughout the region 
between the cancellous portion and the distinct circumferential lamel- 
lae. In the center, the cancellous portion is characterized by lacunae 
of small diameter separated by thick-walled trabeculae. 


TABLE 2 


Measurements in millimeters of teeth of Paleoparadoxia sp. (LACM 
4.371) from Miocene of San Clemente Island. See ‘Table 1 for meas- 
urements of cheektooth (LACM 437 1a). 


LACM 4371b, premolar: 


Greatest diameter of root 16.5 
Least diameter of root 12.0 
Height of crown 21.0 
Maximum width of crown 16.0 
Estimated least diameter of adjacent root 8.0 
LACM 437 1c, tusk fragment: 
Greatest diameter as preserved 23.0 
Greatest diameter of pulp cavity as preserved 15.0 
Greatest thickness of dentine wall 9.0 


Remarks. These San Clemente Island specimens are herein referred 
to Paleoparadoxia and not to Cornwallius because the cheektooth 
(LACM 4371a): 1. although fittmg the original generic diagnosis of 
Cornwallius (“low-crowned teeth furnished with a strong tuberculated 


198 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


cingulum” of Hay, 1923: 107), has a “strong tuberculated cingulum” 
that is not in fact diagnostic, or even present, in teeth of Cornwallius 
but is characteristic of Paleoparadoxia (Mitchell and Repenning, 
1963); 2. has vertical striae around the base of the crown, similar to 
more pronounced striae in a specimen referred to Paleoparadoxia 
tabatai by Reinhart (1959, specimen UCMP 40862); and 3. unlike 
teeth referred to Cornwallius has a prominent swollen crown base. 

The Paleoparadoxia sp. specimens (LACM 4371; the “brachyo- 
dont desmostylid” of Mitchell and Lipps, /n press) represent some of 
the earliest collected (1941) identifiable vertebrate fossils from the 
Tertiary of San Clemente Island other than the ‘““mammalian bones” 
of Olmsted (1958: 64) and undetermined fish scales (Smith, 1898: 
490). Mr. J. H. Lipps and I recently collected many fossils of marine 
vertebrates which are still under study (Mitchell and Lipps, Jn press). 

Paleoparadoxia is circum-North Pacific in distribution, and ranges 
from early to late Miocene in age (Mitchell and Repenning, 1963). 
In the eastern North Pacific, Paleoparadoxia is known from west- 
central California (Reinhart, 1959: 95, specimens UCMP 40862 and 
32076; and Mitchell and Repenning, 1963: , specimens UCMP 
45274 and 63981), possibly from southern California (Reimhart, 
1959: 99, specimen LACM 1352), and from San Clemente Island 
(this report) which is the southernmost known occurrence in this area. 
Teeth referred to Cornwallius, a genus which is restricted to the early 
Miocene of the eastern North Pacific, have been found in Alaska 
(Byers, 1959: 289; Drewes, et al., 1961: 606), in British Columbia 
(Cornwall, 1922; Clark and Arnold, 1923), and the eastern side of 
Baja California (VanderHoof, 1942). Fossils of Paleoparadoxia may 
ultimately be found over as great a latitudinal range as fossils of Corn- 
wallius in the eastern North Pacific. 

Hiri and Kamei (1961: 27) stated that “Paleoparadozia, like Des- 
mostylus, is aquatic (marine) herbivorous . . ” and may be “omniv- 
orous grazing’? Yabe (1959) reported that both “Cornwallius”’ 
(= Paleoparadoxia, sensu Mitchell and Repenning, 1963) and Des- 
mostylus have been found in the same beds in Japan. Paleoparadoxia 
and Desmostylus have been found together in the Santa Cruz area of 
California also (Mitchell and Repenning, 1963). 


SUMMARY AND CONCLUSIONS 


Three teeth and a bone fragment referred to Paleoparadozia sp. are 
described from early or middle Miocene deposits on San Clemente 


Miocene brachydont desmostylian 199 


Island, Los Angeles County, California—this is one of the few North 
American locality records of this genus, and one of the first identifiable 
vertebrate fossils reported from the Tertiary of San Clemente Island. 
One of the tusks of Paleoparadoxia sp. was not round in cross sec- 
tion but had an obvious sulcus on one side. 
Pachyostosis occurs in bones referred to Paleoparadoxia sp. 


CoMPARATIVE MATERIAL 


The following specimens were examined: Cornwallius sookensis- 
UCMP 36078, 36079, and 32682, PMBC 491 (cast), PMBC 486 
(cast); Paleoparadoxia- UCMP 40862, 32076, 45274, 63981; Cf. 
Paleoparadoxid- LACM 1352. This last specimen (LACM 1352) from 
LACM locality 1084 was incorrectly referred to as “CIT 857” by 
Reinhart (1959: 99-101). 


ACKNOWLEDGMENTS 


Mr. Mabry Van Reed, Pasadena Range Director, U.S. Naval Ordnance 
Test Station, Pasadena, California, granted permission and arranged 
for field work on San Clemente Island, and Mr. Albert C. Specht, 
NOTS Administrative Officer, San Clemente Island Engineering Divi- 
sion, was very helpful in logistics on the island. Dr. Gideon T: James 
of the Museum of Paleontology, University of California, Berkeley 
kindly loaned comparative specimens. Mr. Jere H. Lipps of the Uni- 
versity of California, Los Angeles, assisted in the field, and Mr. Lipps, 
Dr. Theodore Downs of the Los Angeles County Museum, Dr. Joseph 
T. Gregory of the University of California, Berkeley, Mr. Charles A. 
Repenning of the U.S. Geological Survey, Menlo Park, and Dr. Peter 
P. Vaughn of the University of California, Los Angeles, criticized the 
manuscript. The illustrations are by Miss Mary Butler, staff artist at 
the Los Angeles County Museum, and by the author. Some of the work 
on the San Clemente Island specimens was supported in part by an 
AAAS Research Grant, administered by the Southern California 
Academy of Sciences. 

This paper is part of the results of a study by Mr. J. H. Lipps and 
myself on the paleontology and geology of San Clemente Island (see 
Mitchell and Lipps, Jn press). It is published as a Los Angeles Mu- 
seum-Channel Islands Biological Survey contribution because the spe- 
cimens were collected in 1941 by that Survey. 


200 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


LITERATURE CITED 


BYERS, E M., Jr. 
1959. Geology of Umnak and Bogoslof Islands, Aleutian Islands, Alaska. U.S. 
Geol. Surv. Bull., 1028-L: 267-369. 


CLARK, B. L., and R. ARNOLD 
1923. Fauna of the Sooke formation, Vancouver Island. Univ. Calif. Publ. Dept. 
Geol. Sci. 14: 123-179. 


COMSTOCH, J. A. 

1946. Contributions from the Los Angeles Museum-Channel Islands Biological 
Survey. 33. Brief notes on the expeditions conducted between March 16, 1940 
and December 14, 1941. Bull. So. Calif. Acad. Sci., 45: 94-107. 


CORNWALL, I. E. 
1922. Some notes on the Sooke formation, Vancouver Island, B.C. Canadian Field- 
Nat., 36: 121-123. 


DREWES, H., et ai. 
1961. Geology of Unalaska Island and Adjacent Insular Shelf, Aleutian Islands, 
Alaska. U.S. Geol. Surv. Bull., 1028-S: 583-676. 


HAY, O. P 

1923. Characteristics of sundry fossil vertebrates. Pan-American Geol., 39: 101- 
120. 

1924. Notes on the osteology and dentition of the genera Desmostylus and Corn- 
wallius, Proc. U.S. Natl. Mus., 65(8): 1-8. 


IJIRI, S., and T. KAMEI 

1961. On the skulls of Desmostylus mirabilis Nagao from South Sakhalin and of 
Paleoparadoxia tabatai (Tokunaga) from Gifu Prefecture, Japan. Earth 
Science (Journal of the Association for the Geological Collaboration in 
Japan), 53: 1-27. 


McLEAN, FE C., and M. R. URIST 
1961. Bone, an introduction to the physiology of skeletal tissue. Chicago: Univ. 
Chicago Press. 261 pp. 


MITCHELL, Ee De jx: 
In press. Pachyostosis in desmostylids. Spec. Pap. Geol. Soc. Amer., (Abstract). 


MITCHELL, E. D., Jr., and J. H. LIPPS 
In press. Miocene marine vertebrates from San Clemente Island, California. 
Spec. Pap. Geol. Soc. Amer., (Abstract). 


MITCHELL, E. D., Jr., and C. A. REPENNING 
1963. The chronologic and geographic range of desmostylans. Los Angeles 
County Mus., Contrib. in Sci., 78: 1-20. 


OLMSTED, F H. 
1958. Geologic Reconnaissance of San Clemente Island, California. U.S. Geol. 
Surv. Bull., 1071-B: 55-68. 


Miocene brachydont desmostylian 201 


REINHART, R. H. 
1953. Diagnosis of the new mammalian order, Desmostylia. J. Geol., 61: 187. 


1959. A review of the Sirenia and Desmostylia. Univ. Calif. Publ. Geol. Sci., 
36: 1-146. 


SMITH, WS. T: 

1898. A geological sketch of San Clemente Island. U.S. Geol. Surv., 18th Ann. 
Rept., 2: 459-496. 

VANDERHOOPF, V. L. 


1937. A study of the Miocene sirenian Desmostylus. Univ. Calif. Publ. Bull. 
Dept. Geol. Sci., 24: 169-262. 


1942. An occurrence of the Tertiary marine mammal Cornwallius in Lower 
California. Amer. J. Sci., 240: 298-301. 
YABE, H. 


1959. A problem on the geological range and geographical distribution of des- 
mostylids. Trans. Proc. Palaeont. Soc. Japan, n. s., 33: 44-51. 


NOTES ON THE OVA OF SIX CALIFORNIA MOTHS 


Joun ApAms Comstock 


Del Mar, California 


In the spring and early summer of 1962 there was an unusual number 
and variety of moths that came to light in Del Mar, California. Many 
of these were noted for the first time in this locality. Ova were obtained 
from several species, some of which were reared to maturity and notes 
have been published on their life histories. A few could not be reared 
farther than the first instar because their food plants were unknown. 
The majority of ova were infertile, but drawings were made as a pre- 
liminary step to further study. 

It seems advisable to record six of these ova since little or no mention 
of them occurs in the literature. 


Autographa (Pseudoplusia) californica Speyer 
Figure 1A. 


This common moth, known as the “Alfalfa Looper:’ was present in its 
characteristic abundance, at black light. The reason ova were secured 
was that no illustration of the egg has been found in easily available 
literature. 


ovum. (Laid May 4 and 5, 1962). Eggs were deposited singly in the 
rearing jar. Color, yellow. Form, hemispherical, the base flattened, 
the top well rounded, and the center depressed by a minute micropyle. 

There are approximately 32 prominent ridges running from base 
to micropyle, but many of these fuse with others or become obsolescent 
in the upper third of the egg. The ridges are topped with rows of large 
round “pearls: There are no apparent cross striations running hori- 
zontally between the ridges. The floor of the micropyle is pitted. 

The larva is an “omnivorous feeder on herbs, except grasses?’ The 
moth ranges along the west coast from Canada to California, and east- 
ward to Colorado. 

Owing to the large number of published records of the life history 
of this common pest, only a few selected bibliographical references are 
given at the end of this paper, namely: Comstock (1930: 25, pl. 9); 
Crumb (1956: 258 f.); Dyar (1890: 14); Essig (1913: 161); Hamp- 
son (1913: 540 f.; pl. 238, fig. 5); Lembert (1894: 46); and Oko- 
mura (1962: fig. 45). 


Ova of six California moths 203 


x 


Fae sath 


Figure 1. Ova of six moths., figures greatly enlarged. Reproduced from water 
color drawing by the author. Ova: A, Autographa californica; B, Aseptis genetrix; 
C, Zale lunata; (top aspect); D, Pherne subpunctata; E, Sericosema simularia; F. 
Pterotaea agrestaria. 


Aseptis genetrix (Grote) 
Figure 1B. 


This noctuid moth was described by Grote (1878: 237) as Hadena 
genetrix from material obtained by Dr. James S. Bailey in “Nebraska, 
Colorado and Nevada?’ Dr. McDunnough, in 1937, made it the geno- 
type of “Aseptis” It ranges from western Canada through Washington 
to Arizona, and eastward through Utah to Nebraska. 

Its larva has been described by Dyar (1898: 320) and Hampson 
(1908: 139 f.). 

Specimens taken in Del Mar on April 10, 1962, enable me to de- 
scribe and illustrate the egg. Captive females deposited their eggs 
singly and indiscriminately in the rearing jar. 


ovuM. Hemispherical, the base flattened. Diameter at base, 0.7 mm. 
Height, 0.5 mm. Color, glistening white. The majority have a circlet of 
yellow blotches around the upper one-third, and similar color in the 


204 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


micropylar depression, as shown in the illustration (Fig. 1B.). 
The surface bears about 30 elevated ridges running vertically from 
base toward micropyle, each topped with a line of pearly nodules. 
The eggs proved to be infertile. The larval food plant is reported to 
be Adenostoma fasciculatum H. & A. 


Zale lunata (Drury ) 
Figure 1C. 


Synonymy male edusa Drury, putrescens Guer. 

This member of the Catocalinae ranges through all states east of the 
Rocky Mountains except the Dakotas and Montana. In the west it has 
been reported from California through Oregon to Washington. Sey- 
eral forms or varieties have been named. 

The larva has been described many times by various writers, but I 
cannot locate any illustrations oN the egg. 

Forbes (1954) reports it as “a general feeder on trees and shrubs:’ 
specifically listed are cherry, maple, plum, willow, oak, blackberry, 
raspberry, salmonberry, wisteria, rose and pyrocantha. 

The species is not common in Del Mar, but a single female was taken 
May 7, 1962. This specimen laid a few eggs the following day, one of 
which served for illustrating. Subsequently (May 11) the eggs 
hatched. 


ovum. Spherical, with a flattened base and well rounded top. Width, 
0.75 mm. Height, 0.5 mm. Color, bright green when first laid, chang- 
ing later to cream, with a suggestion of dull green on the upper por- 
tion and with brownish spots sprinkled over the surface, as shown in 
the illustration. There are approximately 45 ridges running from the 
base toward the micropyle. Many of these fuse or become obsolete im 
the upper portion of the egg. These ridges are topped with minute 
nodules. The micropyle is relatively small; its white floor is deeply 
depressed and is covered with small hexagonal cell walls. 

Notes were made of the first instar larva, largely for comparison 
with French’s (1882) description of the same stage. 


FIRST INSTAR LARVA. Length, average about 5 mm. Body cylindrical 
and elongate, the head slightly wider than the body segments. 

Body, yellow, with a slight greenish tinge; legs and prolegs of the 
same color. There are two pairs of functional prolegs in addition to 
the anal pair. No distinguishing lines or marks occur on the body sur- 


Ova of six California moths 205 


face, but small black papillae are distinguishable, scattered over the 
body and head. These bear minute black setae. 

This is a very active larva, but a sluggish feeder. It eats small patches 
from the surface of leaves of both willow and oak. 

In view of the thorough coverage of remaining stages by many 
authors, no further notes were made. 

Nine references in the literature are helpful in the discussion of this 
species: Behr (1870: 28); Beutenmuller (1901: 192); Crumb (1956: 
293); Forbes (1954: 350 f.); French (1882: 130 f.); Guenee (1852: 
13 f., pl. 2); Hampson (1913: 223 ff., fig. 56); Lintner (1888: 58) ; 
and Packard (1870: 229). 


Pherne subpunctata (Hulst) 
Figure 1D. 


This species was named by Hulst (1898) as Metanema subpunctata. 
His brief description recorded it from “California” with no date, col- 
lector, or type locality designated. 

I published a brief note on the egg and first larval instar in 1955, 
but no illustration was included. We have no information of its range, 
but presume it may turn up in Arizona and Baja California, as it is ap- 
parently a lower sonoran zone species. 

Nothing is known of its food plant. I offered young larvae Rhus, 
Eriogonum and Ceanothus in 1955, and again in 1962 tried several 
local plants without success. 

I include herein an illustration of the egg, and an amplified descrip- 
tion made from specimens taken in late June of 1962. The very brief 
note on the first instar larva previously published will have to suffice 
for the present. 


ovum. Oval, the base flattened and the top evenly rounded. Height, 
0.95 mm. Horizontal diameter through the center, 0.7 mm. (the meas- 
urements given in 1955 were in error). When first laid the color is 
deep cream with the white lines of the vertical ribs showing in strong 
contrast. 

At the base a circlet of raised points marks the origin of the vertical 
ridges. There are from 16 to 20 of these and the edges thereof are 
topped with a line of minute rounded tubercles. The troughs between 
these ridges are crossed by striations corresponding in position and 
numbers to tubercles of the vertical ridges. 

The flattened base of the egg is irregularly pitted. The micropyle is 


206 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


not clearly defined, and is restricted by the terminations of the verti- 
cal ridges. This gives a roughened and crowded effect. 

The majority of the eggs were regularly oval, but a few tended 
toward elongation. 

‘Two references are helpful in the study of this species: Hulst (1898: 
218); and Comstock (1955: 105). 


Sericosema simularia (‘laylor) 


Figure 1E. 


This is a rare moth in the San Diego-Del Mar area, but is probably 
more common in the environs of Pasadena, California, the type lo- 
cality. 

It was described as Enemera simularia by ‘Taylor (1906: 190 f.) 
from specimens collected by Fordyce Grinnell. Nothing has been pub- 
lished on its life history, range, or food plant. 

We confined a female taken at Del Mar May 4, 1962, which laid 
17 eggs, none of which hatched. In late June and early July, addi- 
tional specimens were captured and generously added to our “‘clutch? 
As with the others, none hatched, though apparently fertile. All were 
retired for the winter but, unfortunately, without the moisture and 
temperature of their natural environment. This year (1963) they are 
apparently dead, but show dark bodies internally. This suggests that, 
normally, the embryo overwinters in the egg. 


ovum. Elongate-oval, very flat at the base and evenly rounded at the 
top. All were laid singly on their sides. Length, from base to rounded 
top, 1.0 mm. Width through center, 0.5 mm. Color, bright green. 

The surface is covered by 12 or 13 longitudinal ridges, running 
from base to micropyle, with no fusion or discontinuance at their ends. 
The spaces between the ridges have low striae, placed close together, 
running at right angles to the main ridges. Each of the latter is topped 
by a row of very minute brownish nodules. The micropyle is small and 
irregular, faintly pitted, and not deeply depressed. 


Pterotaea agrestaria (Grossbeck ) 
Figure 1F. 


This geometrid moth was first described in 1909, as Cleora agrestaria, 
by John A. Grossbeck, from material originating in Monterey and San 
Diego Counties. Apparently most of his specimens were collected by 
George H. Field, pioneer lepidopterist of San Diego. 


Ova of six California moths 207 


A group of eggs laid May 8, 1962, showed, typically, the following 
characteristics: 

ovum. Width, 0.4 mm. Height, 0.8 mm. Form, ovoid, with a flat- 
tened base, an evenly rounded top, and no formed micropyle. Color, 
at first salmon-red, fading later to a lighter shade. The eggs were laid 
singly on their sides, not on their flattened bases. 

There are from 12 to 14 longitudinal ridges running from base to 
rounded top. Horizontal ridges of equal height, from 7 to 9 in number, 
run at right angles to the longitudinal ridges. This gives a pattern of 
irregular squares and hexagons with deeply depressed and pitted 
floors. All of the ridges appear to have hyaline edges which show 
white in cross lighting. 

The base appears to possess a ragged membrane suggesting that, in 
a state of nature, they may be laid with their flattened bases on a sur- 
face to which they may be adherent. These features are brought out 
in the illustration (Fig. 1F.). 

It should be mentioned that a second batch of eggs obtained May 25, 
1962, showed a considerable number that were more elongate and sub- 
cylindrical than the typical oval examples of the first lot. 

None of the eggs hatched. We were prepared to feed the larvae on 
Adenostoma fasciculatum H. & A., as this was reported (in litt.) by 
Noel McFarland as the food plant. 


LITERATURE CITED 
(Annotated ) 
BEHR, HERMAN 
1870. Synopsis Noctuidarum Hocusque in California Repertarum. Trans. Amer. 
Ent. Soc., 3: 23-28. (Larva, as H. salicis and H. rosae) 


BEUTENMULLER, WILLIAM 
1901. The larva of Homoptera edusa. J. New York Ent. Soc., 9: 192. (Larva) 


COMSTOCR, JOHN A. 

1930. Studies in Pacific coast Lepidoptera. (Continued). Bull. So. Calif. Acad. 
Sci., 29(1): 22-31. (Pupa) 

1955. The egg and young larva of a geometrid moth from California. Bull. So. 
Calif. Acad. Sci.,54(2): 105. (Egg, young larva) 


CRUMB, S. E. 
1956. The larvae of the Phalaenidae. U.S. Dept. Agriculture, Tech. Bull. no. 
1135, 356 pp. (Larva, food plants) 


DYAR, HARRISON G. 

1890. Preparatory stages of Plusia californica. Entom. Amer., 6: 14-15. (Egg, 
larva, pupa) 

1898. Descriptions of the larvae of fifty North American Noctuidae. Proc. Ent. 
Soc. Wash., 4: 315-332. (Larva) 


208 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


ESSIG, E. D. 
1913. Injurious and beneficial insects of California. Bull. St. Comm. Hortic. 


Calif., 2(1-2): 1-367. (Egg, larva, pupa) 


FORBES, WILLIAM T. M. 
1954. Lepidoptera of New York and neighboring states. Noctuidae. Pt. III. 
Cornell Univ. Agricultural Exp. Sta.. Mem. 329. 433 pp. (Larva, food plants) 


FRENCH, G. H. 
1882. Preparatory stages of Homoptera lunata, Drury. Canadian Ent., 14(7): 
130-134. (Life History) 


GROTE, A. EB: 
1878. New N. American Lepidoptera, with notes on a few little known. Canadian 
Ent., 10: 231-238. (Original description) 


GUENEE, A. 
1852. Species general Noctuelites. (from Abbott mss.). 3: 13-14. Colored figs., 
Pl. 2. (Larva, pupa) 


HAMPSON, GEORGE EF 

1908. Catalogue of the Lepidoptera Phalaenae in the British Museum. 7: 1-709. 

1913. Catalogue of the Lepidoptera Phalaenae in the British Museum. 13: 1-609. 
(Ege, larva, pupa) 


A455? 
HULST, GEORGE D. 
1898. Descriptions of new genera and species of the Geometrina of North Amer- 
ica. Canadian Ent., 30(8): 214-219. (Original description) 


LEMBERT, JOHN B. 
1894. Food plants of some California Lepidoptera. Canadian Ent., 26(2): 45-46. 
(Food plants) 


LINTER; J. A: 

1888. Fourth report on the injurious and other insects of the state of New York. 
4th Rpt. N.Y. St. Entom., In 41st Rpt. N.Y. St. Mus. Nat. Hist. pp.1-347. 
(Larva) 


OKOMUBRA, GEORGE T. 

1962. Identification of lepidopterous larvae attacking cotton, with illustrated key 
(primarily California species). Dept. Agr. Calif.. Spec. Pub. No. 282, 80 pp. 
(Larva, setal maps, food plants) 


PACKARD, A. S. 
1870. A few words about moths. Amer. Nat., 4: 225-229, (Larva, pupa) 


TAYLOR, GEORGE W. 
1906. On some new species of Geometrid moths from Arizona and California. 
Ent, News, 17(6): 188-192. (Original description) 


A STUDY OF THE ZYGOPINAE (COLEOPTERA: 
CURCULIONIDAE) OF AMERICA NORTH OF MEXICO, I.* 


ELBERT L. SLEEPER 


Long Beach State College, Long Beach? 


While engaged in a study of the genus Cylindrocopturus Heller of the 
subfamily Zygopinae, it became increasingly obvious that there was 
a need for a study of the whole of the Zygopinae. This is the first of 
three papers on this group. 

The author is indebted to numerous individuals for aid on the papers, 
especially: Drs. Fred Truxal, Charles Hogue, and Mr. Lloyd Martin 
of the Los Angeles County Museum for making material and library 
facilities available; Miss Rose E. Warner (now Mrs. T: J. Spilman) for 
aid in studying material at the United States National Museum 
(USNM); Dr. Barry D. Valentine for information on the type ma- 
terial at the Museum of Comparative Zoology (MCZ), Harvard; and 
Dr. William D. Stockton for aid with the manuscript. Acknowledg- 
ment is made to National Institute of Health, research grant AI-3407 
(Richard B. Loomis, Principal Investigator), for aid in studying ma- 
terial first hand in northwestern Mexico. In addition to the above ab- 
breviations the following have been used: ELS=E.L. Sleeper Col- 
lector, (ELS)=E.L. Sleeper Collection, FDPI=Entomological Col- 
lection Florida Department of Plant Industry, and OSU=Ohio State 
University. 


KEY TO THE GENERA OF Z,YGOPINAE IN THE UNITED STATES 


ieeeyeacimmexposed; (Nig. 1) S555 082.6. 5 5. 5. Peltophorus Schoenherr 
tea cidmm completely concealed) (Pigi7) Wee este ace eel 2 
2. Abdomen horizontal throughout (Fig. 4); eyes approximate on the front; 
hunrenignotobliquely: truncate). 4... 5.44 salen etc: ae aie aeons 3 

2a. Abdomen ascending rapidly distally (Fig. 6); eyes and humeri variable. 
2 0.6.06 6 Bb. S SLSR E NMS R oR RAE Troe orci ane Och one Gata din. Alas Gee lela te aaae Pi bee eels Ye Tee 4 
3. Femora minutely toothed; antennae stout with well developed club; body 
denseliyascaliy;; ely tral striaevcoars?’ 94)... 445.4. -0-." 3. Acoptus LeConte 
3a. Femora not toothed; antennae very slender, the club small; body subglabrous 
abovermelyitrall: striae fires 301 Wyss ee nee eat oie 4. Psomus Casey 


1Contribution #23 to the knowledge of the Curculionoidea. 
Biological Science Paper #9, Long Beach State College. 


209 


210 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


4. Pectoral channel extending over the mesosternum. occasionally ending in the 


6a. 


N 


metasternum, channel clearly limited laterally by a raised ridge or carina 


(Bigs 10). sei. ie ose ie elas sae ok ieseeee ele canto atc ok 6) 
. Mesosternum not excavated, apex of rostrum free (Fig. 3) .............. 7 
Femora minutely or moderately toothed ......... 1. Lechriops Schoenherr 
- Femiora unarmed 3 ic 22. fer font ee bie as ods heck sa ee eee ee 6 
Mesosternum and anterior margin of metasternum deeply impressed or ex- 
cavated by pectoral channel (Fig. 10) .............. 2. Eulechriops Faust 


Mesosternum with a pectoral channel indicated only by a feeble concavity 
and prominent lateral carinae, metasternum not impressed, the latter in- 
clined: imitromty cee sega tera: 7. Cylindricopturinus, new genus 
. First two segments of funicle little different in length; second never more 
than one and one-half times as long as first .................-5-..:-5--- 8 
Second segment of funicle very long, more than twice as long as the first 
Co a Pa eh A calor A eLah years ea a 8. Copturus Schoenherr 
. Mesosternum flat, slanting, non-carinate laterally 6. Cylindrocopturus Heller 
. Mesosternum very feebly concave, carinate laterally between meso-and 
metacoxal Cavitlesi-c: vas wine carats ae ae 7. Cylindrocopturinus, new genus 


1. Lechriops Schoenherr 


Lechriops Schoenherr 1826, p. 306. Type species Rhynchaenus sciurus 


Fabricius. 


Piazurus LeConte 1876, p. 259. (Not Schoenherr, 1826) 
Gelus Casey 1897, p. 667. Type species Cryptorhynchus oculatus 
(Say ) 


This genus occurs in North and Central America and northern 


South America as well as on some of the islands of the West Indies. 


KEY TO THE SPECIES OF Lechriops oF AMERICA NortTH OF MEXICO 


. Prothorax about twice as wide as long; common sutural spot of elytra not 
conspicuous; elytra much wider than prothorax; total length 1.7 mm. .... 


RA Ua E ES hs tenth Rene Beinn, aPC OR eo Eero BOR ore & subfasciata (LeConte) 


. Prothorax normally slightly wider than long (varying from 5:5 to 5:8) never 


more than one and one-half times wider than long; common sutural light 
spot of elytra very conspicuous .......... 0550.0... 2 eee eee 2) 
Elytra but little wider than prothorax; outline more elongate, slender (Fig. 
5); elytra noticeably longer than wide (measurements of length down sutu- 
ral line); total length 2.0-3.5 mm.; width 1.2-1.6 mm. 7{27Seereeee 
Re fain ee eben ee REE His il rls Blas tad &. 0-0 californica (LeConte) 


. Elytra much wider than prothorax; outline more robust ((Fig. 7); elytra 


but little’ longer than wide .. 2.2.0. 060.0. 0000 os oe ene 3 
Smaller, 3.0-3.2 mm.; elytral suture with a prominent light spot near middle: 
pronotum with a conspicuous pattern of pale brown scales; east of the con- 
tinental divide south into Mexico and Central America ....... oculata (Say) 


American Zygopinae Daa 


3a. Larger, 3.5-4.5 mm.; elytral suture with a prominent light spot and sutural 
line from declivity to apex; pronotum without pattern of lighter scales at 
wanGelles (Mey 18 VNGLAOEY Teme acbooogahoogooobeedoo grisea, New species 


Lechriops subfasciata (LeConte) 


Piazurus subfasciatus LeConte 1876, p. 260. Type locality: “New York, 
Mr. Guex;’ type in MCZ. 

Gelus subfasciatus (LeConte), Casey 1897, p. 668. 

Lechriops subfasciata (LeConte), Blackwelder 1948, p. 47. 


This species is still known only from the unique type, which may 
represent an introduced species from an undetermined Latin Ameri- 
can Country. 


Lechriops californica (LeConte ) 
Figure 5 


Piazurus californicus LeConte 1876, p. 260. Type locality: “Calveras, 
California,’ type in MCZ. 

Gelus californicus (1eConte), Casey 1897, p. 668. 

Lechriops californica (LeConte), Blackwelder 1948, p. 47. 


Distribution. Rather abundant in western United States and northern 
Mexico. Specimens are at hand or have been seen from southern Ore- 
gon, Utah, Nevada, California, Arizona, New Mexico, Baja California 
Norte, Sonora, and Chihuahua. It is limited to areas of pme growth, 
being, however, infrequently encountered on pinyon pine (Pinus cem- 
broides var.). It is very abundant on P ponderosa and P. jeffreyi 
throughout all of the range of this weevil. The larvae and pupae have 
been encountered beneath or in the bark of P jeffreyi in southern Ca- 
lifornia and Baja California Norte. 


Lechriops oculata (Say ) 


Cryptorhynchus Say 1824, p. 308. Type locality: “Inhabits Missouri,’ 
here restricted to St. Louis, Missouri, type lost. 

Lechriops oculatus (Say), Heller 1895, p. 14. 

Gelus oculatus (Say), Casey 1897, p. 668. 

Lechriops oculata (Say ), Blackwelder, 1947, p. 879. 


Distribution. Generally distributed throughout North America east of 
the Rocky Mountains and the Sierra Madre Oriental. Its range in 
Canada is undetermined, but examples are at hand from Quebec, On- 


212 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


tario and Manitoba. Numerous examples have been seen from Mexico 
and Guatemala. 

This species has been taken on Quercus spp., Hicoria sp., Sassafrass 
sassafrass L., Fraxinus spp., Fagus grandifolia Ehrh., Viburnum spp.. 
and Crataegus spp. 


Lechriops grisea, new species 
Figures 6 and 7 


Holotype. USA., Arizona, Cochise Co., Chiricahua Mtns., Rucker Cyn., 
VIII-10-59, 7000’, type no. 74, (ELS). 


Male. Length 3.3 mm., width 2.0 mm.; elongate-oval; black with 
antennae and tarsi reddish brown; head and basal third of rostrum 
sparsely clothed with oval, black and white scales, vertex of head with 
scattered seta-like scales in punctures, prothorax with overlapping oval, 
yellowish-white scales on sides, a few black and brownish seta-like 
scales on disc, elytra moderately clothed with black, brown, white and 
yellowish white scales, the latter in patches at base of elytra and on 
apical half of sutural intervals. 

Rostrum nearly three times as long as wide at base (11.5:4), nearly 
six times as long as width at antennal insertion (11.5:2), as long as 
the prothorax (11.5:11), sides convergent from base to antennal in- 
sertion, parallel sided thence to apex, in lateral outline strongly ar- 
cuate; coarsely, rugosely punctured and with a prominent median 
carina in basal third, apical two-thirds smooth, sparsely, very finely 
punctured; mandibles feebly bidentate. Antennal insertion at basal 
third of rostrum, scape shorter than the first two segments combined, 
not attaming base of rostrum; funicle with first two segments longer 
than remainder combined (ratio 1.3:2.2:.8:.6:.5:.5:.6), sparsely 
clothed with fine setae. Club oval-acuminate, basal segment constitut- 
ing nearly half club length (1:.5:.2:.4), sparsely clothed with fine 
setae. Head continuous with rostrum, depressed but not foveate be- 
tween the eyes, dorsum closely punctured with only the vertex with- 
out punctures, vertex strongly alutaceous, front narrow, the eyes 
separated by only one-third width of rostrum at antennal insertion, 
eyes finely granulate. 

Prothorax wider than long (11:15.3), widest just before base, sides 
convergent from base to apex, apex tubular but not constricted, ocular 
lobes absent, base bisinuate, apex emarginate; disc closely, coarsely 
punctured, each with a subclavate scale, punctures of sides coarse, but 
obscured by vestiture, each with a large oval overlapping scale, disc 


American Zygopinae 213 


Figure 1. Outline of Peltophorus polymitus seminiveus (LeConte). Figure 2. 
Outline of Peltophorus adustus (Fall), male. Figure 3. Outline of pro- and meso- 
sternum of Copturus floridanus (Fall). Figure 4. Lateral outline of elytra and 
sternites of Acoptus suturalis LeConte. Figure 5. Lechriops californica (LeConte). 
Figure 6. Lateral outline of Lechriops grisea, new species. Figure 7. Lechriops 
grisea, new species, holotype. Figure 8. Lateral outline of aedeagus of Peltophorus 
polymitus suffusus (Casey). Figure 9. Lateral outline of aedeagus of Peltophorus 
adustus (Fall). Figure 10. Outline of pro- and mesosternum of Eulechriops 
minutus (J. E. LeConte). Figure 11. Dorsal outline of aedeagus of Peltophorus 
adustus (Fall). Figure 12. Dorsal outline of aedeagus of Peltophorus polymitus 
seminiveus (LeConte) with outline of dorsal part of tegmen. Figure 13. Dorsal 
outline of aedeagus of Peltophorus polymitus suffusus (Casey) with outline of 
dorsal part of tegmen. Line = 1 mm. 


214 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


carinate from base to apical fourth. Scutellum prominent, smail, 
rounded, punctured and alutaceous. 

Elytra only slightly longer than wide (12.5:11.1) ; humeri rounded 
off not prominent, but slightly wider than base of prothorax; sides 
strongly arcuate from base to apex, apices feebly emarginate; disc 
feebly convex, feebly depressed behind the scutellum; striae narrow, 
deep, with rectangular punctures, each separated by a narrow trans- 
verse carina, each puncture with a recurved subclavate seta, striae 
seven and eight not attaining base; intervals flat with close, deep, con- 
fusedly placed punctures each with a broad clavate scale. 

Sternal side densely clothed with oval, white, rarely overlapping 
scales, which are for the most part appressed, each originating in a 
coarse puncture, area between punctures strongly alutaceous on ab- 
dominal sternites 3-5, on remainder of venter feebly alutaceous. Pros- 
ternum and mesosternum as in other species, metasternum concave 
between mesocoxae; intercoxal process very broad and truncate; ab- 
dominal sternites 10.5:4:1.5:2:3, first very long and with an oval con- 
cavity longitudinally at middle, second abruptly bent upward along 
posterior margin, apex of fifth truncate. Legs clothed with white ap- 
pressed oval scales. Front coxae with a backward projecting tooth. 
Femora linear, compressed, all prommently unguiculate externally, 
the posterior pair mucronate internally. ‘Iarsi elongate, as long as the 
tibiae, first two segments slender, clothed with prominent white seta- 
like scales, third broadly bilobed, naked, glabrous above, densely pilose 
beneath, fourth slender with inconspicuous reddish brown setae, ratio 
lengths of segments of hind tarsi 4.5: 1.6:1.6:2.5. Claws slender. 

Allotype. Female, 4.25 mm., width 2.4 mm., differing from the 
male only in the less concave first abdominal sternite and having the 
fifth sternite rounded apically. 

Other localities. USA, Arizona, Cochise Co., Huachuca Mtns., Park- 
er Cyn., 6500’, VITI-12-59, ELS, (ELS). 

Type material. Holotype, allotype, 12 paratype all with the same 
data, 2° paratypes, from Parker Cyn. The @ paratype from the 
type locality deposited in the Entomological Collections, Los Angeles 
County Museum, remaining type material in (ELS). 

Biology. All of the examples from Parker Cyn. were taken while 
beating oak, where they were observed feeding on leaf petioles. One 
example from the type locality was also atest ed on oak. 

This species may be separ ated readily from the other species known 
from the United States by its larger size ‘and the prominent sutural spot 
and line, the latter bemg very conspicuous. It does not compare favor- 


American Zygopinae 215 


ably with any of the material described in the Biologia Centrali-Amer- 
icana nor any known species from northern Mexico. 


2. Eulechriops Faust 
Figure 10 


Eulechriops Faust 1896, p. 91. Type species Eulechriops erythroleucus 
Faust. 

Zygomicrus Casey 1897, p. 667. Type species Eccroptus minutus J. E. 
LeConte. 


This genus is distributed from northeastern United States to north- 
ern South America. Three species were previously listed in the litera- 
ture from America north of Mexico. One of these species is here refer- 
red to a new genus, leaving E. minutus (J. E. LeConte), from the 
United States and FE. sobrinus (Horn) from the Cape Region of Baja 
California Sur. The undescribed forms will be the subject of a future 
paper now in preparation. 

All of the United States forms are associated with various species 
of Quercus. 


3. Acoptus LeConte 
Figure 7 


Acoptus LeConte 1876, p. 264. Type species Acoptus suturalis Le- 
Conte, the type in MCZ. 

Homogaster Provancher 1877, p. 530. Type species Homogaster que- 
becensis Provancher, location of type unknown, a synonym of A. 
suturalis, according to Blackwelder and Blackwelder, 1948: 47, 


A monotypic genus with the single species, described from “New 
York;’ distributed in the United States and Canada east of the Rocky 
Mountains, and in extreme northeastern Mexico. 

In eastern United States this has been taken from Quercus spp., 
Cercis canadensis L.., Hicoria sp., and Platanus occidentalis L. 


4. Psomus Casey 


Psomus Casey 1892, p. 458. Type species Orchestes armatus Dietz, 
type im MCZ; Psomus politus Casey, type in USNM, a synonym of 
O. armatus; Fall 1913, p. 64. 

A single species occurs in the United States and Canada east of the 


216 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


Rocky Mountains, and three species are known from Central America. 

P. armatus (Dietz) is frequently associated with green ash (Fraxi- 
nus lanceolata Borck. and F. americanus L.) It is uncommon in col- 
lections. It is most frequently collected in June. 


5. Peltophorus Schoenherr 
Peltophorus Schoenherr 1845, p. 451. Type species Peltophorus poly- 


mitus Boheman. 


This genus is restricted to arid areas of the United States and Mexico. 
The type locality for P polymitus polymitus Boheman is “Mexico, 
Villa Alto in Oaxaca?’ In the United States three kinds are known from 
western Texas, New Mexico and Arizona. 


Key TO THE KINDS OF Peltophorus oF AMERICA NortH oF MEXICO 


1. Prothorax with sides parallel in basal two-thirds to three-fourths (Fig. 2); 
abruptly subrectangular constricted in apical third; pronotum coarsely, 
cribrately punctured, their edges forming longitudinal ridges in some exam- 
Plesot yeh wint cele canta sales Paes a eee adustus (Fall) 

1a. Prothorax with sides convergent from base to apex with a feeble apical con- 
striction (Fig. 1); pronotum coarsely punctured, with a few punctures coal- 
escent, their edges never forming prominent longitudinal ridges ......... 2 

2. Range, southwestern Texas; dorsum of body with black, white and brown 
scales; punctures of elytral striae large, encroaching upon intervals; fifth 
ventral abdominal sternite without a prominent spot of scales each side of 
middle; aedeagus as in Figs. 8 and 13 .......... polymitus suffusus (Casey) 

2a. Range, southern Arizona and southwestern New Mexico; dorsum of body 
with black and white scales; punctures of elytral striae no wider than striae; 
fifth abdominal sternite with a prominent spot of black scales each side of 
middle; aedeagus as in Fig. 12 ......... polymitus seminiveus (LeConte) 


Peltophorus adustus (Fall) 
Figures 2, 9 and 11 


Zygops adustus Fall 1906, p. 61. Type locality “Arizona; here re- 
stricted to Arizona, Pima Co., Santa Rita Mtns., Lower Madera Cyn.., 
type in MCZ. 


Distribution. Moderately abundant in the Santa Rita, Huachuca and 
Chiricahua Mtns., in southern Arizona, and at Rodeo, New Mexico. 
Nearly all examples examined were taken from Agave palmeri Englm. 


American Zygopinae 217 


Peltophorus polymitus suffusus (Casey ) 
Figures 8 and 13 


Zygops suffusus Casey 1892, p. 459. Type locality “Texas (southwest- 
ern)” here restricted to Texas, Jeff Davis Co., Ft. Davis, type in the 
USNM. 

Peltophorus polymitus suffusus (Casey), Blackwelder 1947, p. 881. 
Distribution. Texas: Davis Mtns., VII-2-40, V-9-41, VII-17-46, 

V1-29-49, DJ & JN Knull, (OSU); Chisos Mtns., VII-17-46, DJ & 

JN Knull, (OSU); Val Verde Co., V-13-46, DJ & JN Knull, (OSU) ; 

Sanderson, VI-15-56, D. G. Genung, (ELS, FDPI). 


_ Peltophorus polymitus seminiveus (LeConte) 
Figures 1 and 12 


Zygops seminiveus LeConte 1884, p. 31. Type locality “Arizona,” here 
restricted to Arizona, Cochise Co., Ft. Huachuca, type in MCZ. 
Peltophorus polymitus seminiveus (LeConte), Blackwelder, 1947, p. 

881. 

Distribution. Arizona: Many records from the Santa Rita, Hua- 
chuca, Chiricahua, Pinal, Santa Catalina, Dragoon and Patagonia 
Mtns., May through September. Common in several species of Agave 
including A. palmeri. 


Cylindrocopturus Heller 


Cylindrocopturus Heller 1895, p. 56. Type species Zygops quercus Say 
1831, p. 20. 

Paratimorus Heller 1895, p. 58. Type species P ganglbaueri Heller. 

Copturodes Casey 1897, p. 667. Type species, none designated, Zygops 
guercus Say, by present designation. 

Gyrotus Casey 1897, p. 668. Type species Gyrotus minutus Casey, by 
monotypy, mew synonmy. 


The genus occurring throughout the United States and Canada (ex- 
cept for northwestern United States and Western Canada), southward 
to Argentina. Thirty species occur in Baja California and America 
north of Mexico. Forty-three species are known from the New World. 
The species of this genus will be treated in subsequent papers. 

The designation of Zygops quercus Say as type species of Copturodes 
Casey was necessary in order to establish its synonomy with Cylindro- 
copturus. Z. quercus seemed the logical choice inasmuch as it is also 


218 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


type species of Cy lindrocopturus and it was Z. quercus and forms re- 
lated to it that were treated in the 1897 paper, for the most part. 

After a study of many examples of G. munitus and comparing this 
species with numerous examples of Cylindrocopturus from Mexico I 
have come to the conclusion that there is no characteristic that will set 
Gyrotus apart from Cy lindrocopturus. The pronounced modification 
of the prothorax 1 in the region of the postocular lobes used by Casey 
fora major characteristic is repeated 1 in varying degrees 1 im Many spe- 
cies of Cylindrocopturus from Mexico. The other characteristics grade 
into or are shared in varying degrees with members of that genus 
found in the United States. As a result of the synonomy G. munitus 
Casey must now be known as Cylindrocopturus munitus (Casey), new 
combination. 


7. Cylindrocopturinus, new genus 


Type species Eulechriops pictus Schaeffer, here designated. 

Rostrum as in Lechriops; first two segments of funicle subequal; 
eyes large, separated by less than one-fourth the width of the rostrum 
at the point of antennal insertion; prothorax moderately bisinuate at 
base. without indication of ocular lobes or modification thereof; scutel- 
lum conspicuous, round; elytra deeply and conspicuously striate. 
completely concealing pygidium from above; mesosternum feebly con- 
cave, metasternum feebly convex, not exacavated, mesosternum cari- 
nate laterally between pro- and mesocoxae, apex of rostrum resting 
free; abdominal sternites ascending rapidly distally in lateral outline. 

This genus is differentiated from the other North American forms 
by the characteristics pointed out in the key. particularly in having 
the non-excavate metasternum and the lateral elevations on the meso- 
sternum which form a pectoral channel. Lechriops has the femora 
armed with a very small to moderately large tooth and the pectoral 
channel extending into the metasternum. Eulechriops has unarmed 
femora as in this genus but differs in the deep excavation of the meta- 
sternum. Cylindrocopturus has no indication of a pectoral channel on 
meso- or metasternum. 


Cylindrocopturinus pictus (Schaeffer). new combination 


Eulechriops pictus Schaeffer 1908, p. 219. Type locality “Huachuca 
Mtns., Arizona;’ type in the USNM;; Sleeper 1954, p. 182. 


A rather rare and without doubt the prettiest of our Zy gopinae from 
the United States. known only from the mountains of south central 


American Zygopinae 219 


Arizona. Specimens have been examined from the Santa Rita, Hua- 
chuca, and Patagonia Mountains. 
A single example was beaten from Quercus arizonica in Madera 


Canyon of the Santa Rita Mtns. in July. 


8. Copturus Schoenherr 


Copturus Schoenherr 1826, p. 302. Type species Rhynchaenus lamella 
Fabricius. 
Coptorus Schoenherr 1826, p. 302. (Error in spelling. ) 


This genus is represented in America north of Mexico by a single 
species found only in southeastern United States. The genus ranges 
generally: southward to Argentina. More than 156 species have been 
described. 


Copturus floridanus (Fall), new combination 


Figure 3 


Piazurus floridanus Fall 1906, p. 61. Type locality “Florida (Key 
Largo and Elliots Key): here restricted to Key Largo, type in MCZ. 


This species is frequently encountered in Florida in Dade and Mon- 
roe Counties particularly on the Keys, and in Cuba. 

In Florida it was taken by beating miscellaneous vegetation on hey 
Largo, and in Matheson Hammock. It has also been taken on Swietenia 
mahogont. 


LITERATURE CITED 
BLACKWELDER, R. E. 
1947. Checklist of the Coleopterous insects of Mexico, Central America, the West 
Indies and South America. Part 5, Bull. U.S. Nat. Mus., 185: 765-925. 


BLACKWELDER, R. E. and R. M. BLACKWELDER 
1948. Fifth supplement 1939-1947 (inclusive) to the Leng cataloguz of the Cole- 
optera of America north of Mexico. Mount Vernon, N.Y., 87 pp. 


CASEY, T. L. 
1892. Coleopterological notices II. Ann. New York Acad. Sci., 6: 359-712. 
1897. Ibidem. VII. Ann. New York Acad. Sci., 9: 285-684. 


FALL, H. C. 

1905. New species of American Coleoptera of the tribe Zygopini. Trans. Amer. 
Ent. Soc., 32: 53-56. (Continued in 1906.) 

1906. Ibidem. Trans. Amer. Ent. Soc., 32: 57-61. (Continued from 1905.) 

1913. A brief review of our species of Magdalis with notes and descriptions of 
other north American Rhynchophora. Trans. Amer. Ent. Soc., 39: 23-72. 


220 ~=©>— Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


FAUST, J. 
1896. Reise von Simon in Venezuela. Curculionidae. Stettiner Ent. Zeitung, 57: 
33-135. (Pars tertia.) 


HELLER, kK. M. 

1895. Zygopiden-Studien II, mit besonderer Beriicksichtigung der Gattung Cop- 
turus. Abhandlungen und Berichte des koeniglichen zoologischen und an- 
thropologisch-ethnographischen Museums zu Dresden, 1894/95, No. 11, pp. 
1-70, illus. 


LeCONTE, J. L. 
1884. Short studies of North American Coleoptera (No. 2). Trans. Amer. Ent. 
Soc., 12: 1-32. 


LECONTE, J. L. anp G. H. HORN. 
1876. The Rhynchophora of America north of Mexico. Proc. Amer. Philos. Soc.. 
15: 1-455. 


PROVANCHER, L. 
1877. Petite faune entomologique du Canada precedee d’un traite ellementaire 
d’entomologie, vol. 1—Les coleopteres, Quebec, 786 pp. 


SCHAEFFER, C. FE A. 
1908. New Rhynchophora. III. J. New York Ent. Soc., 16: 213-222. 


SCHOENHERR, C. J. 

1826. Curculionidum dispositio methodica cum generum characteribus, descrip- 
tionibus atque observationibus variis, seu prodromus ad synonymiae insec- 
torum partem 4, Lipsiae, 339 pp. 

1845. Genera et species curculionidum, cum synonymia hujus familiae. Species 
novae aut hactenus minus cognitae, descriptions a Dom. Leonardo Gyllenhal, 
C. H. Boheman, et entomologis aliis. Illustratae. Vol. 8, pt. 2, pp. 1-504. 


SLEEPER, E. LL. 
1954. New Rhynchophora II (Coleoptera, Curculionidae). Ohio J. Sci., 54: 180- 
186, illus. 


THEODORE PAYNE 
1872 - 1963 


With the passing of Theodore Payne on May 6, 1963, the Southern California 
Academy of Sciences lost one of its most active members and the last surviving 
Charter Member. ; 

Mr. Payne was born in Northamptonshire, England, June 19, 1872. As a small 
boy he was always fond of flowers and had a small garden of his own. At the 
age of 12, he was sent to Ackworth School in Yorkshire where he joined a natural 
history society. His special choice, from the first, was botany, and while at this 
school he became secretary of the botanical section, made a collection of pressed 
plants for which he was awarded the first prize, and became active in conserva- 
tion and horticulture. 

After leaving Ackworth, he was apprenticed for three years to the firm of John 
Cheal & Sons, Lowfield Nurseries, Crawley Sussex, to learn the nursery and seed 
business, a vocation which was to become his life-long career. 

In the spring of 1893 he decided to come to California. On June 3rd, as he 
neared his 21st birthday, he sailed for New York and arrived in Los Angeles, June 
28th. Shortly after arriving here he secured a position as gardener to Madame 
Helena Modjeska, the famous Polish actress, at her country estate, “Forest of 
Arden?’ Santiago Canyon, Orange County, where he remained for two and one 
half years. 

He then entered the employ of the Germain Fruit Company (now Germain 
Seed Co.) in April, 1896, and was placed in charge of their flower, tree and palm 
seed department. While with this firm, he also handled much of the correspond- 
ence pertaining to technical information and prepared all their catalogues. 

Mr. Payne, having decided to go into business for himself, resigned from the 
position with the Germain Fruit Company in 1903 and bought the Hugh Evans 
Nursery then located at 440 South Broadway. Two years later he moved his busi- 
ness to 345 S. Main Street, and subsequently bought a nursery at 33rd and Hoover 


221 


222 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


Streets which he used for growing-grounds. He maintained his nursery at the 
Main Street address until 1922, when he purchased 10 acres of land fronting on 
Los Feliz Blvd.. and moved the nursery to that location. Here, he continued his 
nursery, growing mainly native trees, shrubs, and wild flowers, until he retired 
in June, 1961, at the age of 90. 

As a seed and plantsman, Mr. Payne has contributed a wealth of information 
about the nature of plants. the growing of cuttings, and the germination of seeds: 
particularly of native plants. Many of the plants that dot our orchards, parks, 
highways, and landscaped homes, passed through his hands. 

Early in his career. he made a study of the Eucalyptus and when the Euca- 
lyptus boom occurred about 1907, he soon became the headquarters for these 
seeds in the United States. Between 1911 and 1913 he sent out hundreds of pounds 
of these seeds to Central and South American and European countries. 

When Mr. Payne first came to California he was impressed with its native 
flora. With deep regret, he saw the wild flowers rapidly disappearing from the 
landscape. He decided, then early in his career, to awaken interest in the native 
flora and began growing wild flowers and native plants. To further the interest 
in beautiful native plants for home gardens, he sowed wild flower seeds in a 
vacant lot in Hollywood, and in several such lots in Pasadena. He secured from 
Walter Raymond, of the Raymond Hotel in Pasadena, a plot of ground on the 
hotel site and sowed it with wild flower seeds: the following spring his efforts 
resulted in a profusion of wild flowers. His first wild flower catalogue, of many 
that were to follow, was a modest little booklet published about 1906. 

In 1915, Mr. Payne was commissioned by the City Council of Los Angeles to 
plant wild flower beds in Exposition Park. This “wild flower garden” consisted 
of 5 acres and was planted with native trees, shrubs, perennial plants, bulbous 
plants, and a large area of annual wild flowers. In all, the garden contained 
262 species, the whole being a reproduction of a natural landscape; each specimen 
was labeled with its botanical and common name. 

In 1919, and later, Mr. Payne did landscape work in addition to his nursery 
work. He landscaped a number of estates in the Santa Barbara area, Ojai, Los 
Angeles, Pasadena, and elsewhere; the largest was the 140-acre estate of Mrs. 
Lora J. Knight in Montecito. Among the native plants that he planted there in 
quantity were 121 Monterey Pine and 30 pounds of seed of the “Blue-eyed Grass”’ 
He kept 10 men on this place to maintain and cultivate the plants and made 2 
trips each month to supervise their work. 

During this period, shortly after the commencement of work on the Knight’s 
estate, Mr. Payne formed a landscape architects partnership with Ralph D. Cor- 
nell and for 5 years, during their association, they completed some interesting 
developments including Torrey Pines Park and Occidental College. 

In 1926, he was requested by Mrs. Susanna Bixby Bryant to help her select a 
site on her ranch in Santa Ana Canyon for a native botanic garden. The Rancho 
Santa Ana Botanic Garden was founded the following year. Mr. Payne served on 
its advisory council for over 20 years, until the time the Garden was moved to 
Claremont. 

During his career in horticulture, Mr. Payne introduced into cultivation in 
California between 400 and 500 species of wild flowers and native plants and made 
them available for general use. He wrote many articles on native plants, acquaint- 
ing readers with the desirability of these plants for home gardens, and gave talks 


Theodore Payne 223 


in nearly all the towns in southern California on “Preserving the Wild Flowers 
and Native Landscape of California?’ 

Mr. Payne received a number of honors and awards for his outstanding achieve- 
ments in horticulture and in recognition of his work in conserving the native 
flora of California. He held a number of memberships in botanical societies, 
nature conservation groups, and horticultural societies. He was a charter member 
of several organizations. He held one honorary fe membership, one honorary 
membership, four life memberships, and was Fellow of the Royal Horticultural 
Society of England. He served as president of several horticultural, arboricultural, 
wild flower and nurseryman’s associations. 

The County of Los Angeles and the County Department of Parks and Recrea- 
tion honored Mr. Payne by setting aside a plot of ground, consisting of 320 acres, 
near Llano in the Mohave Desert as a wild flower sanctuary. This was dedicated 
as the Theodore Payne Wildlife Sanctuary on January 28, 1961. In 1958, Mr. 
Payne began the development of a 5-acre site in the Descanso Gardens as a wild 
flower garden, planting 23 species of conifers, 14 groups of other trees, 92 kinds 
of shrubs, 8 perennial plants, and seeds of 39 species of wild flowers. The dedica- 
tion of this garden was held in the spring of 1959, at which time Mr. Payne was 
presented a scroll from the California State Legislature and another from the Los 
Angeles County Board of Supervisors in recognition of his work in conserving 
the native flora of California. 

In addition to this activity in the many organizations to which he belonged, 
he was an active member of the Southern California Academy of Sciences for 
the greater part of his life; only in the past several years, as age and declining 
ill health came upon him, did he become less active. He first became a member in 
1898, and in May 1907, at the time the Academy was incorporated, he became 
a charter member. In May, 1918, he was elected secretary of the Botanical Sec- 
tion and for many years thereafter served in this capacity. He was elected to the 
Board of Directors on June 20, 1920 where he served for about 30 years, later 
serving on the Advisory Board. He was President of the Academy from May 
1932 to May 1933. Already a Fellow, he was made an Honorary Life Member 
of the Academy in May, 1957. 

Mr. Payne will always be revered by all who knew him for his many kind- 
nesses and for his wise and patient nature, for his love of flowers and the Cali- 
fornia native plants. Many who came to him for information about plants or to 
chat with him, whether amateur or professional horticulturalist or botanist, left 
richer in knowledge and with great respect for a wonderful man. He was never 
too busy to share his time, his knowledge of botany, and his experiences in grow- 
ing native plants. 

As his life was nearing to a close, he saw one of his hopes being realized—that 
of a foundation devoted to the purpose of perpetuating the knowledge and con- 
servation of California native plants and the growing of these plants for the home 
garden and roadside planting. This organization, ““The Theodore Payne Founda- 
tion for Wild Flowers and Native Plants?’ was named in his honor. 

A final honor given Mr. Payne, posthumously, was the “Man of the Year” 
award for 1963 by the California Garden Clubs, Inc. Although Mr. Payne re- 
ceived many similar honors, he was a modest man and never allowed pride in 
these things to overshadow his devotion to love for wild flowers and his service 
to mankind. 


224 Bulletin So. Calif. Academy Sciences / Vol. 62, Pt. 4, 1963 


A Memorial Fund in the Southern California Academy of Sciences and in the 
Theodore Payne Foundation has been established in his honor, Many friends 
have contributed and are still contributing to these funds. 


| Bonnie C. TEMPLETON 
Los Angeles County Museum 


SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 


VOLUME 62, 1963 


INDEX OF SUBJECTS 


A contribution to the biology of the 


mreeny einen Gli S66 encuecesee 83 
A fossil bird, Caracara, from Santa 
Rosa Island. Erratum. ......... 18 


A study of the Zygopinae (Coleop- 
tera: Curculionidae) of America 


north of Mexico .............. 209 
VA\GTMACONTMUTG jos leas. 105 
VACTIIGCOMPCILE Nein eisai sei. ee as 104 
A\GODOUS oom 85.6 AHO COR SO Ee 215 


An experimental study of the echo- 
location ability of a California 
sea lion, Zalophus californianus 


GEessorl) Meee vse cieie De ss aiej oes 165 
IATISCRILOGIMES: Gy. shoe est eee een 182 
ASG IIS [FIRGIUE Meo Stones ee oon. 202 
ANSHO O1BUS. oo 0-8 Seo Sito 180, 183 


Attraction of insects to exudates of 
Verbesina encelioicles and Iva 


ambrosiaefolia ............... 109 
VAUOST AP WOANDUIODA . 2... 2. 2. 36 
Autographia (Pseudoplusia) cali- 

JOTTUNOL. i 61 alo eae ROE Bee 202 
BOUGTIUSKCOTIOSUSE inc. fame ess 2 - 106 


Bess Reed Peacock—Obituary ... 22 
Birds and indians in the West ....178 


Boccardia columbiana ..........:132 
Boccardia proboscidea ......... Ley, 
BOMOSAMUTMOCILUSH sos eyee 183 
PSULCOMANLACETISIS 22)... = see 179 
calico, Protochlorotettix ......... 78 
Centrocercus urophasianus ...... 183 
(COUDHIGS CLP Sood sab SRO Oe 180 


Comparison of Boccardia colum- 
biana Berkeley and Boccardia 
proboscidea Hartman (Annelida, 
Polychaeta 6... ec, cee ooo 132 

Contributions from the Los Angeles 
Museum—Channel Islands _bio- 
logical survey. 37. Brachydont 
desmostylian from Miocene of 
San Clemente Island, California . 192 


CODERS. Bis min Cc ec ie P sane hes 219 
Copturus floridanus ............. 219 
CORO MG) (IZIUTR 5 on On ened oon 106 


Corvus brachyrhynchus ....181, 184 


(CORDYS COREE sob0ccnc00cb0000060 181 
Cylindrocopturinus .............. 218 
Cylindrocopturinus pictus ........ 218 
QAM OCH OTS coscsavcessso0 217 
Dendragapus obscurus ...... 179, 182 
Deroceras reticulatum .......... 83 
Desmosty Way yee. ww) sok iy ee ce 193 
dietrichi, Hyperodes ............ 145 
Hmertiayanalogaie nr pe 45 
ESCRTICRtIUS SRLQUCUS Ta arin ere 99 
Ele cRntopsny sa ney ee eee ine oe 215 
Euphydryas eurytion ........... 19 
Euscelis palmeri ................ 2 
Halcoicolumbaritus) ee ree 182 
IICNCO) GeiPABUS sooshoe0c00000+ 179 
Halconiformiesieeea son re oe 182 
floridanus, Copturus ............ 219 


Fossil arthropods of California. 
No. 25. Silicified leafhoppers 
from California mountains nod- 
WS Sis eave Ae sen tn Wiara sees carey ase ea 69 

Functional morphology of the ex- 
ternal appendages of Emerita 
GNGIOS AYE eve ME ee 45 

Further studies of the benthic fauna 
in a recently constructed boat 


harbor in Southern California .. 23 
Gallitormes (09 e sides eters 182 
gibroni, Miochlorotettix ......... 73 
gracilis, Orchestoidea ............ 3 
grisea, ’cchriopses nena sere ee 212 
Haplotrema vancouverense ...... 106 
hoodi, Hyperodes ............... 144: 
Hyperodes dietrichi ............. 145 
Hyperodes hoodi ............... 144 
Hyperodes texana .............. 142 
Hyperodes wallacei ............. 140 
Iva ambrosiaefolia .............. 116 
juliae, Miomesamia ............. 81 


225 


kirkbyi, Miochlorotettix ......... 77 
Lacinipolia quadrilineata ........ 33 
Lechriops LE ae ee ene tele PN 210 
Eechriops CAUjOrmiCG =) <5. cei. 911 
LECCRTIONS: GriSeG = :42 2. 2 ache) sea 912 
Gechriops OCulaia 20. oss se: 211 
Eechriops subjasciata 2... -. a). 211 
UE CDOS: | ASCLCLLICT IS. eae ena 130 
Lepospondyl remains ........... 151 
Miochlorofettix ................. 73 
Miochlorotettix gibroni .......... Wi 
Miochlorotettix kirkbyi .......... 77 
Miomesamia ................... 81 
Miomesamia juliae .............. 81 
Molluscs from Pacific Northwest 


archaeological sites, 2. Washing- 

ton: 45-CA-30, a coastal shell- 

midden in the Ozette area ..... 101 
Mytilus californicus 


Neptunea tabulata 2 o.4..4.824)- 104 
New information on the structure 
of Permian lepospondylous ver- 
tebrae—from an unusual source. 150 
New species of Hyperodes Jekel and 
a key to the Nearctic species of 
the genus. (Coleoptera: Curcu- 
honidae ) 
Notes on the barnacle Lepas fasct- 
cularis found attached to the jel- 
lyfish Velella 


Notes on the larva and pupa of Eu- 
phydryas eurytion (Lepidoptera, 
Nymphalidae) 

Notes on the life histories of two 
Southwestern phalaenid moths .. 33 

Notes on the ova of six California 


INVOUTS saree chee ramus rR re 
Olvella ibiplicatas enna fast 104 
Orchestoidea gracilis ............ 3 


Orchestoidea gracilis, a new beach 
hopper (Amphipoda: Talitridae) 
from Lower California, Mexico, 
with remarks on its luminescence 1 


ORCOFEY 2, PIClOnas woke rac yeer toa: 183 
OLS A510 Cl os fcr NO estan: 180 
Paleoparadoria 2) act. arson 193 
Paleoparadoxadaees 4-1-0. ciekciccre 193 


palmeri, Huscelis ............... 72 
Passeriformes 2... secre 184. 
Pedioecetes phas: ‘anellus RRS 180 
Peltophorus. .- > 2 eee 216 
Peltophorus adusiusss--eeeeere 216 
Peltophorus polymitus seminiveus .217 
Peltophorus polymitus suffusus ...217 
Pero macdunnouchi == epee 44 
Pherne subpunctata =p ee 205 
Phlepsius weissmanae ........... 80 
Pica pica... <2. eee 181, 184 
pictus, Cylindrocopturinus ........ 218 
Proceedings of the Academy ..... 159 
Protochlorotettix ............... 78 
Protochlorotettix calico .......... 78 
Protothaca lacinaia =. Eee 103 
Psomus . o53 0h) ote oe 215 
Pterotaza agrestaria ............ 206 
Schizotherts nuttalliy oe See 104 
Sericosema simularia ........... 206 
Strigiformes .:. 32/2656 sees 183 
Siurnella neglecta. sn ee 182 


Surf-riding by the California gray 


whale... 0.0.) 2.2) eeeeeer 99 
legula ‘funebralis.: 2) see eee: 104 
texana, Hyperodes 3.5450 peace 142 
Thais lima . i... a= oe eee 104: 
The early stages of Pero macdun- 

noughi Cassino and Swett (Lepi- 

doptera, Geometridae) ........ 41 
The late Pleistocene 150 foot fresh 

water beach line of the Salton 

Sea area ...0. 50 ccc g 
The number of conenose bugs, Tria- 

toma, infected by one engorge- 

ment on a mouse with Trypano- 

SOM CTUZI |... soe ee 176 
Theodore Payne—Obituary ......221 
Triatoma p. protracia sa... 176 
Trypanosoma cruzi” 5. - eee 176 
Turdus migratorius .2...s-oee as: 181 
Velella. . .24.3+. 25 eo eeeeOee 130 
Verbesina encelioides ........... 110 
wallacei, Hyperodes ............ 140 
weissmanae, Phlepsius .......... 80 
Zale lunata ... 3.54 eee 204 
Zalophus californianus .......... 165 
Zygopinae ..... .: «mse eeeeey: 209 


226 


INDEX OF AUTHORS 


ANTRIAG, IR, (O)e., oacsene ete, cele aie ene 83 
Boolootian, Richard A............ 45 
IXoQGMEG), 1, Ibs coogupenoueoege. 1 
Caldwell, David Kk. ............. 99 
Caldwell, Melba C.............. 99 
(Gaziersa VA eis a elo coe sone es 109 
Comstock, John Adams .. .33, 41, 202 
Crowe lly Ws cass ak Gees 83 
WrakewRoberntie o- 6: ss 52202. 5: 101 
Emmel, Thomas C.............. 19 
Evans, William E. .............. 165 
Flaugenwhuthy Mis 2... eas 165 
ilawieemv apes eyeich sii chseicle see te ihe 1 
knox Gameromyon ss s4 2.5). sa) a 68 45 


to 


Knudsen, Jens W................ 130 
Teimsleyee he Gag. te orn ates: 109 
Malllersmltovesisiy easign ita se cee 178 
IMGT AEIDL, ICh, ID, Wie; socb00cece 192 
Bierce. Nie Owaglitieeere nn eee: 69 
ReishysDonal di|aeaeeeneeeece 23 
Sleepers lb exten r ence 209 
Stockton, William D. ........... 140 
Templeton, Bonnie C......... DODO 
Thomas, Robert G. ............. 9 
Waushn Peter sei ee aneeer rie 150 
Wood, Sherwin E .............. 176 
Woodwick, Keith H.............. 132 


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