FBH-GAME

..■^^miE

California Fish and Game Is a journal devoted to the conserva-
tion of v/ildlife. Its contents may be reproduced elsewhere pro-
vided credit is given the authors and the California Department
of Fish and Game.

The free mailing list is limited by budgetary considerations to
persons who can make professional use of the material and to
libraries, scientific institutions, and conservation agencies. Indi-
viduals must state their affiliation and position when submitting
their applications. Subscriptions must be renewed annually by
returning the postcard enclosed with each October issue. Sub-
scribers are asked to report changes in address without delay.

Please direct correspondence to:

LEO SHAPOVALOV, Ed/for
Department of Fish and Game
926 J Street
Sacramento 14, California

Individuals and organizations who do not qualify for the free
mailing list may subscribe at a rate of $2 per year or obtain indi-
vidual issues for $0.75 per copy by placing their orders with the
Printing Division, Documents Section, Sacramento 14, California.
Money orders or checks should be made out to Printing Division,
Documents Section.

u

1

J

VOLUME 41

OCTOBER, 1955

NUMBER 4

Published Quarterly by the

CALIFORNIA DEPARTMENT OF FISH AND GAME

SACRAMENTO

STATE OF CALIFORNIA

DEPARTMENT OF FISH AND GAME

GOODWIN J. KNIGHT
Governor

FISH AND GAME COMMISSION

WILLIAM J. SILVA, President
Modesto

ANDY KELLY, Commissioner CARL F. WENTE, Commissioner

Los Angeles San Francisco

HARLEY E. KNOX, Commissioner WELDON L. OXLEY, Commissioner

San Diego Redding

SETH GORDON
Director of Fish and Game

CALIFORNIA FISH AND GAME
Editorial Staff

LEO SHAPOVALOV, Editor-in-Chief Sacramento

JOHN FITCH, Editor for Marine Fisheries ^._._ Terminal Island

CAROL M. FERREL, Editor for Game __ Sacramento

J. B. KIMSEY, Editor for Inland Fisheries _._ Sacramento

TABLE OF CONTENTS

Page
Spoond Progress TJcpoft on the Ta;^'<i'iiiji- of White ('jitfisli
[Ictalunis c(tti(s) in the Saei-aniento-.San Joaquin Delia

David E. Pelgen and George W. McCammon 2(il

h'liii't iiiit ions in A linnihim-c of ( 'ci-tnin l^"'ishes in Sont h San h'ran-
eisco Hay as Indicated l)y SaiiiplinL;- at a Trash Screen

Knrl S. Ilti-iihl mill Dniinhl A. Siinpsnn 271

Tolerance of tlie San Franeiseo Topsmelt, Atherinops affinis
(iffi)iis, to Conditions in Sall-l'r<)duein<>' Ponds Borderin<;- San
Franeiseo Bay Lars II. Carpdan 279

Age and Lenfrth Composition of the Sardine Catch OfiF the Paeifie
Coast of the United Stales and ^lexico in VJ^A-^)')

Frances E. Felin, John MacGregor,

Anita E. Bauglierty, and Daniel J. MilJ< r 2.'^5

Nestino; Stndies of Ducks and Coots in Honev Lake Valley

E. G. Uuii'f atul A. E. Naxjhn- 205

'o

Nestinir and Peprodnction of the Band-Tailed Pi<ieon in Califor-
nia Wallace G. Macgvfgov mid Wnlfni} .V. S^niifh 315

Skeletal Differences Between Deer, Sheep, and (ioats

Milton Ilildfhrand 827
Notes

AVhitc Sturiicon With St-vcn Rows of Scutes

Alexander D. Bajkov 347

A Record of an Attack liy a Leopard Shark (Triakis seniifas-
ciaia (lirard) ' Jnlni W. DrWitt 348

Second Known Sliai'lc Attacl; on a Swimjncr in .Moutei'ey Bay

Thomas X. Fast 348

]\ ('views 353

Index t(. Volume 41 359

( 2.-,9 )

SECOND PROGRESS REPORT ON THE TAGGING OF

WHITE CATFISH {ICTALURUS CATUS) IN THE

SACRAMENTO-SAN JOAQUIN DELTA'

DAVID E. PELGEN and GEORGE W. McCAMMON

Inland Fisheries Branch

California Department of Fish and Game

This report i-uiist itutes the second of a series on the results of tag-
ging studies begun in 1952 on the white catfish {Ictalurus catns) of
the Saeraniento-San Joaquin Delta. The ultimate objective is to derive
a valid index of the annual rate of harvest of white catfish in the Delta
fishery. Such information is fundamental to the conservation of this
valuable resource.

These investigations were secondarily aimed at revealing informa-
tion about the movements of white catfish in the Delta. Knowledge of
any regular, seasonal migrations was particularly desired.

The initial report (Pelgen, 1954) described preliminary holding ex-
periments and presented the results of one year of tag returns from
tagged white catfisli released in the Delta. In summary, opercular strap
tags proved unsatisfactory. The disk-dangler tag appeared promising
and worthy of further tests. Tag returns from fishermen indicated only
sporadic, directionless movement of fish.

The present report describes the second year's results of the initial
study, and presents the i)reliminary results of a second experiment
that was designed to compare two types of tags, to further explore
movements, and to estimate the annual rate of harvest.

FIRST STUDY

The inadequacy of the opercular strap tag was confirmed by obser-
vation of experimentally tagged fish, and by tag returns during the
second year of the first study.

Pelgen (1954) reported that both opei-ciiiar straji tags and disk-
dangler tags remained attached to white catfish held in hatchery ponds
for one year, althougli the strap tags became hidden under a layer of
opaque mucus. After the experimentally tagged fish had been held for
22 months, a check revealed that a uumber of strap tags had become
detached. Ten were lost and most of the 'M tags that remained were
very loose.

None of tlie 20 disk-dangler tags were missing, although several had
irritated the flesh surrounding the points of attachment. Excessive
netting and handling are Ijelicved primarily responsible for these irri-
tations. The autlioi's liave I'aii'ly iibscf\ed siii-1i ii'i'itati<»ns 0)1 tagged
catfish in tiie Delta.

1 Submitted for publication .June, 195."). Tliis work wa.s performed as part of Dingell-
Johnson Project California F-2-R. "A Study of the Catfish Fishery of California'",
supported by Federal Aid to Fi.sh Restoration funds.

( 2G1 )

262

CALIFORNIA P'ISH AND GAME

Ta<r returns from catfisli reloased in tlie Delta in 1952 diminished
greatly during' the second year. Table 1 summarizes returns i'roin the
first study for both the first and second years. The superiority of the
disk-danjrler tag over the opercular straj) tag remains evident despite
the few returns duriuL'' the secoiul year.

TABLE 1
Data on Tags at Large and Tags Returned from First Study

Tags at large at beginning of
eacli period

Tags returned during each period

Study jieriod

Strap

Disk-
dangler

Total

Strap

Disk-
dangler

Total

First year

Second year

2,865
*2,790

601
*503

3,466
*3,293

75

(2.6%)

9

(0.03%)

98

(16.3%)

14

(0.3%)

173

(5.0%)

23

(0.07%)

♦ These figures were actually lower than Indicated since natural mortality, loss of tags, and nonreturn of tags were
not considered.

SECOND STUDY

Methods

Two types of tags were compared in this study : the disk-dangler tag
and the staple tag. These tags, with their position on the fish, are illus-

FIGURE 1. Leff: staple tag shown in place. Righf: disk-dangler tag shown in place.

WHITE CATFISH TAGGING 263

tratod in Fip'nrc 1. iJotli tyjK^s Avcro ()i-i<iiiially (l('V('l()])c(l in coiinoction
witli tagyiiiji' studies on striped bass, lioccus sd.ratilis (Calhoun, lO.l.'i).
Tliere is actiiall.\- litili' (lirfei-ence Ix'tween these two tags. The only
distiiu'tion lies in the |)osition of tlie disk in relation to the body of
the tish. Tt was lioped tliat th(> staph* taj^-'s snii<i' position ajjainst the
tish wouhl reduce the ])ossil)ility of entanglement with gill nets, weeds,
and (it lief ol)st I'uet ions.

All tags Avere constructed of the same materials: cellulose nitrate
disks, one-lialf indi in diameter and ().040-ineh tliiek, and n.()20-inch
tantalum wire. The disks were supplied by liastian Brothers Company
of San Francisco.

The method of attachment of both tags to the body of the catfish
was identical to that used by Pelgen (1954).

Thi'ee sepai'ate tagging stations (Figure 2) were selected to obtain
an adequate distribution of tagged fish throughout the population.
The results of the first study, in which six tagging stations were used,
indicated that the dispersion from the point of tagging was sufficiently
thorough to eliminate the necessity for more than three stations.

Fyke nets - similar to those used by commercial catfish fishermen
(Warner, 1949) were used to capture the fisli for tagging. Each net
possessed seven circular iron hoops, grading from 45 inches to 28 inches
in diameter from front to rear. The webbing consisted of number 15
cotton twine with 2 i -inch stretched mesh. Funnels were hung from the
second, third, and fifth hoops. The last funnel had an opening approx-
imately four inches in diameter.

All nets were dipped in heated coal tar at frequent intervals to
prevent rapid detei-ioration of the webbing. It has been observed in
the past b}- commercial fishermen, and more recently by project per-
sonnel, that nets treated in this manner catch significantly more white
catfish in the Delta than do nets treated with other preservatives.

It was originally planned to use 250 tags of each type at each of
the three stations, for a total of 1,500. However, since one disk-dangler
tag was not used at the Antioch Bridge station, the total was actuallv
1,499.

Tagging was carried out between December 16, 1952, and January
27, ]!)5.'l Winter tagging is believed to be distinctly superior to sunnuer
tagging. The reduced metabolic rate of the fish in cold weather maj^
moderate the initial effects of the tagging operation. In addition, some
disease-producing and parasitic orga,nisms are less active and thus less
likely to secure a foothold in the tag wonnd Avhen water teni|)ei-atures
are low.

At two stations. Disappointment Slough and Cache Slough, both
netting and tagging were performed at the release point. This woi-k
was done with the .aid of the Department of Fish and Game research
boat "Stkii'EK." Every effort was made to handle the fish carefully.
Only a few fish were removed from the nets at one time and they were
quickly tagged and returned to the water. Experienced workers usually
require slightly less than 80 seconds to apjily a tag. The procedure at
the Antioch Bridge station was slight Iv dilferent. White catfish Avere

" Nets with funnels but without wings are commonly called "fyke nets" in California.
Such nets are frequently termed "hoop nets" in many other areas.

264

CALIFORNIA FISH AND GAME

® An

tioch Bridge

(2) Disappointment Slough
C^ Cache Slough

Rio Vista

w^^^

:f:<:->=^

Antioch

o

10

=1=

15 Miles

=1

Scale

iacramento

"%

p

•S",

c

o
o

CO

^>

f^^

• Tracy

FIGURE 2. The three localities at which tagged catfish were released.

trapped in Mayberry Cut, two iiiiU's northwest of tlio Antioch Bridge,
and transported to a field hase installation at the south end of the
brid^T', where the fish wci-e i-etained in a liA'e box and tafi'<>'ed from
a doeic.

The foi'k length of eacli eatlisli was measured lo the nearest tenth of
an ineli.

After the catfish were tagged, the success of the study was entirely
dependent upon the return of tags by sport and commercial fishermen.

NVIIITK ('A'l'l'lSll TAGGING

2 (•..")

Tlcnco, tho importance of rctuniiiiji- ta^-s was adNCftiscd t liroii^hoiil the
Delta afca. Posters, press releases, prize (lra\vin<;s, i)ers()iial contact with
resort dw ii(>rs and spoi'tsineii 's clnhs, and einiiiiiendation cards all played
a |)art in the publicity canipai^ni. The prize di-a\vin<,' sponsored by the
Foothill Sportsmen's C'liib of Oakland was f)robabIy the most effective
sin',de i)iiblicity method. An annual i)rize of ^M)(), tog'ether with numer-
ons awards of tishing equipment, was generously donated by that group.

Tag Returns

Table 2 summarizes tag return data during the first 12 months of the
stnd\-. The total return from both sport and commercial fishermen was
I.").! percent. The largest number of tags was recovered from catfish
i'eleas(Ml at the Antioch Bridge, where the return was 21.0 percent. The
l)isai)i)ointment Slough and Cache Slough stations yielded returns of
1 1.2 percent iiud 13.2 percent, respectively.

TABLE 2
Data on Tags Attached and Tags Returned During First Year

Tags attached

Tags returned

TasKing station

Disk-
dangler

Staple

Total

Disk-
dangler

Staple

Total

Antioch BridKe --

249
250
250

250
250
250

499
500
500

54
(21.7%)

33
(13.2%)

35
(14.0%)

51
(20.4%)

23
(9.2%)

31
(12.4%)

105

I )isappointnient Slough

Cache Slough

(21.0%)

50
(11.2%)

66
(13.2%)

Totals -

749

750

1,499

122
(16.3%)

105
(14.0%)

227

(15.1%)

The range in size of the fish tagged was 6,1 to 12,4 inches, fork
length. The mean length was 8,2 inches. The size range at the time
of tagging of the catfisli from which tags were returned was 6.3 to
12.0 inches and the mean length was 8.3 inches. A chi-square test
revealed that a statistically significant difference exists between the
.size of catfi.sh tagged and the size of catfish from which tags were
returned (P<0.003). The actual number of returns from catfish under
8.5 inches fork length was less than would be expected were the return
of tags independent of the size of the fish tagged, while returns from
catfish over 8.5 inches were gi-eater than would be expected. No explana-
tion for this difference is available at the present time.

Suitability of Tags

As shown in Table 2, lt).3 percent of the disk-dangler tags and 14.0
percent of the staple tags were returned. The difference is not statis-
tically significant at the 5-percent level. At no time during the second
study did any evidence appear that definitely demonstrated the supe-
riority of either tag. Comparative tests of the suitability of both tags
on other species of fish are under way in other projects, and they may

266

CALIFORNIA FISH AM) GAME

(liscloso a(lvan(aj,'('s of one ta<r over tlio other that ai-e not now ai)pareiit.
One sueh test has shown tliat the sta])le ta<2: beneath the dorsal fin is
unsatisfaetory for use on lar<re steelhead rainbow trout {Salmo gaird-
ncri (jainlneri) .

The sheddin<r of unsatisfaetory tajOfs ean often be detected by observ-
iuL' the time of recovery of ta^jrs. Tal)le '^ j)i-e.sents the number of returns
ill tliis stuily by monthly intervals following time of tagging.

TABLE 3
Tag Returns by Monthly Intervals During First Year

Tags returned

Elapsed time
in months

Staple tags

Disk-dangler tags

Totab

Number

Percentage

Numlicr

Percentage

Nvimber

Percentage

0- 1 __

1- 2

11

8

14

12

14

10

8

8

9

5

5

1

10.5

7.0

13.3

11.4

13.3

9.5

7.0

7.0

8.0

4.8

4.8

1.0

6

3

9

18

13

10

13

21

0

10

1

6

4.9

2.5

7.4

14.8

10.7

13.1

10.7

17.2

4.9

8.2

0.8

4.9

17
11
23
30
27
26
21
29
15
15
6
7

7.5
4 8

2- 3

10 1

3- 4

4- 5

5- 6

6- 7

7- 8

8- 9.

9-10 .

13.2

11.9

11.5

9.7

12.8

0.0

0 6

10-11 .

■' 0

11-12

3.1

Totals

105

—

122

—

227

—

From these data it is not evident that either type of tag was becoming
detached in significant numbers. It is believed that the variation in
number of returns is largely governed by changes in angling effort.
The periods when the most tags were returned coincide with the periods
of greatest angling intensity in the Delta.

Harvest

T"'ntil F!eptember, Vd'Vi, the Delta Avhite catfish ])opulati()ii was tlie
object of botli a commercial and a sport fishery.

The commercial fishery was relatively small in terms of tlic nnmber
of fishei-men involved and gross income. For instance, in 1!)52, approxi-
mately 20 licensed commercial fishermen reported landing 17-1,452"^
])ouii(ls of catfish with a total value to tlie fishermen of $."ir),7()."} (Pelgen,
lf)52). The actual commercial landings were niKloiihtcdly larger than
the record indicates, since there was good evidence that some of the
marketed catch was not reported as recpiired by law.

S])ort fishing for catfish is fairly important in California. In lf)58,
an estimated 225,000 anglers (19 percent of all license holders) caught
ai)proximately T.-IOO.OOO catfisli (Rkiniier, 1955).

Fividcnce of a dc^cline in the Delta catfi.sh fishery (Pelgen, 1!)52) led
to the elimination of the commercial fishery by the California Legisla-

' Poundage corrected to cover cleaning loss of 50 percent.

WillTK t'ATFlSll TAGGING 2G7

luiv ill SepteiiilxT. 1!)");}. For all pi-actical piifposos, commercial flsliin<r
ill tlic Delta duiiiiu' l!*-").'} was limited to the inoiiths of January, Febru-
ary, ^larch, and Api'il, since tiie eommorcial closed season extended
from May 1st to Anjjust 31st.

Thus, both the commercial and sport fisheries were active during the
first four months of the tafi-jz-infi: study. In this period, a total of 107
ta«rs was recovered. Si)ort fisli(>rmeii turned in 75 tags, commercial
fishermen i-etui-ned 28 tags, and project personnel accounted for four
tags.

It is believed that very few sport fishermen failed to retui-n tags.
Undoubtedly there Avere instances in which anglers captured tagged
catfish but (lid not return the tags for various reasons. No basis exists
for estimating the percentage of nonreturn by sport fishermen; how-
ever, the absence of any indication of tag nonreturn of significant
propoi-tions leads the authors to presume that nearly all tags were
returned.

In contrast, it is believed that a number of commercial fishermen
failed to return a substantial inimber of tags. During the four-month
l)eriod when the connnei'cial fishery was active, only seven of approxi-
mately 20 licensed commercial men returned any tags at all. These
seven fishermen reported a total catch of 12, .544 pounds. The total catch
by all nshei-men during the same four months was 2S,314 pounds. Thus,
the seven cooperative fishermen, who were responsible for all of the 28
commercial returns, caught al)<)ut 44 percent of the total 1953 landings.
It is surmised that the remaining 56 percent of the 1953 commercial
catch was taken by individuals who were either disinterested or unsym-'
pathetic toward the tagging program. The evidence that some commer-
cial fishermen were not inclined to return tags was verified by personal
contact in several instances.

The etfect of nonreturn of tags by the conunercial fishery upon
the estimate of annual harvest is dit!icult to assess. If it is assumed
that the catch of the seven fishermen who returned tags was repre-
sentative of the total commercial catch, then a correction can be
made to compensate for tags not returned. On this basis, it is esti-
mated by simple proportion that the commercial fishery recovered 63
instead of 28 tags during the first year.

Addition of the corrected commercial tag returns to tlie 195 returns
from anglers increases the total number of tag returns from 223 to
258. The four tags recovered by project personnel are not included
in this total. The revised sum represents a total return of 17.2 percent.

With the correction for commercial nonreturn of tags, a minimal
harvest of api)roximately 17.2 ])ercent of the available white cattish
in the Delta is indicated for 1953. The luirvest by sport fishermen
alone is estimated to be at least 13.0 percent. These estimates are
tentative, pending the results of further studies in ]')rogr(>ss.

Following the elimination of the counuercial fishery in 1115.3. a
third experiment was begun, using only disk-dangler tags. Tlie results
of this study should provide a valid index of the annual harvest of
white catfish by the sjiort fishei-y. A subsequent rejxtrt will describe
the findings.

268

CALIFORNIA FISH AND GAME

Movement of Tagged Catfish

It WHS possible to determine accurately the movements of 194 of
the 227 ^vhite catfish from -which taprs were returned. Sixty-eight fish
traveled upstream, averaging- 10.4 miles; 76 fish traveled downstream,
averag-iii'T 8.7 miles. Fifty individuals were reeai)tured in the imme-
diate vicinity of llicii- point of release.

Tlie movements of the taoffred catfish in both the first and second
studies were snrprisin<j'ly similar. Table 4 summarizes movement data
from both studies. The only noticeable difference lies in the large
number of fish captured near tlie ])<)int of release in the second study,
(^nly 20 catfish were recovered near their release point in tlit^ fii-st
study.

TABLE 4

Movement of Tagged White Catfish in the First and Second Sacramento-San Joaquin Delta Tagging Studies

Number
of fisli

for
which
movement
was de-
termined

No
movement

Movement

Tagging study

Upstream

Downstream

Number
fish

Average
distance

Number
fish

Average
distance

I'irst Delta study

(First year's returns)

'Second Delta study
(First year's returns)

158
194
352

20
50
70

85

68

153

9.8

10.4

*10.1

53

76
129

8.6

8.7

Totals-

*8.7

Mean distance.

Tlie earlier evidence that the movement of white catfish in the Delta
is sporadic and without apparent direction (Pelg'cn, 1954) is cor-
roborated by these results.

ACKNOWLEDGMENTS

Acknowledgment is due Alex Calhoun, under whose guidance the
tagging studies were conducted. We are also grateful to "William E.
Rowley, Jr., who assisted in the tagging of the fish, and to Vincent
Catania, who assisted in their capture. Special thanks are due the
Foothill Sportsmen's Club of Oakland, particularly Mr. Ed Busalak,
who generously sponsored the prize drawings, which stimulated so
much public interest in the project.

SUMMARY

Tagging studies of the white catfish (Ictalurus catus) in the Sacra-
mento-San Joaquin Delta area of California were begun in 1952, in
order to obtain data on annual harvest and migration. This is the
second progress report on the findings of these studies.

All disk-dangler tags on white catfish held in hatchery ponds
remained in place for 22 months, wliile a significant number of oper-
cular strap tags were shed.

W 1 1 ITi; CATFISH TAGGING -^>!^>

Very few taps -wore returned (hii-iii<i' the second year i'roiu ;J,4GI)
tagged white eatiisli reh'ased in the Delta in the sprin<-' of 1952. How-
ever, tlie pereentas'e of retnrn foi- disk-(hiiiu-l(T taj^s was 10 times
that for strap tags.

Ill the winter of ^97^2-^^'^, 1,4!)!) white eat fish were tagged witli
disk-dangler tags and staple tags in ecpial inimbcrs and released at
three localities.

Tlie total return of both types of tags from all sources was 15.1
percent dni'ing the first 12 months. The Antioch Bridge tagging station
produced a high return of 21.0 percent.

A significant dift'erence was noted between size of fish tagged and
size of fish from which tags were returned. Fewer tags were returned
from fish under 8.5 inches than would theoretically be expected.

Fourteen percent of the staple tags and 16.3 percent of the disk-
dangler tags were returned. The difference was not statistically sig-
nificant.

It is estimated that a minimum of 17 percent of the available
white catfish was harvested by sport and commercial fishermen in
the Delta in 1953. The true figure is undoubtedly somewhat larger,
because of failure to return tags.

No evidence of any regular, seasonal migration of white catfish in
the Sacramento-San Joaquin Delta was found.

LITERATURE CITED

Culhoiin, A. J.

1953. Aquarium tests of tags on striped bass. Calif. Fish and Game, vol. 39, no. 2,
p. 2()!)-21S.

Pelgen, David E.

1952. A preliminary analysis of the catfish fishery of the Sacraniento-San
.Toaquin Delta. Calif. Dept. Fish and Game, Inland Fisheries Branch,
Admin. Rept. No. 52-23. 10 i). ( .MIiik-o. )

1954. Progress report on the tagging of white catfish {Ictuhnus catus) in the
Sacramento-San .Joaquin Delta. Calif. Fish and Game, vol. 40, no. 3,
p. 313-321.

Skinner, John E.

1955. California state-wide angling estimates for 1953. Calif. Fish and Game,
vol. 41, no. 1, p. 19-32.

\A'arner, George H.

1949. Catfish. In: The commercial fish catch of California for the year 1947
with an historical review 1".I|(;1947. Calif. Div. Fisli ;ui<l (iaine. Fish
Bull. 74, p. 53-57.

FLUCTUATIONS IN ABUNDANCE OF CERTAIN FISHES

IN SOUTH SAN FRANCISCO BAY AS INDICATED

BY SAMPLING AT A TRASH SCREEN'

EARL S. HERALD and DONALD A. SIMPSON

Steinhart Aquarium, California Academy of Sciences,

San Francisco

The presence of a fixed fish-eollecting device on the west shore of
south San Francisco I>ay has been a considerable asset to Steiiihart
Aquarium over tlie past years. In addition to providin<i' fishes for dis-
play and as food for aquarinm fishes, the records of the catches have
provided valnable information about some of the population dynamics
within tlie hnv.

0, CHINESE

Shrimp nets

Francis co

STATION A

FIGURE 1. Map of a section of Son Francisco Boy, showing location of the Station A

pumping plant of the Pacific Gas and Electric Company and its relation to the fishing area

of the Chinese shrimp nets.

1 Submitted for publication May, 1955. Data for tliis report have been taken bv the
variou.s collectors at Steinhart Aquarium, but chieflv bv the junior author and
Mr. Walter Schneebeli. We wish to thanlc Mr. W. I. Follett and Mrs. Lillian
Demiistei- of the academv s depart mem of ichthyology for their assistance in
identification and in checking records. Also, our thanks are extended to the Pacific
Gas and Electric Company for having made the trash screens available as a
collecting device over the past years.

(271)

272

(AI.II'OKMA I'lSll AND (iA.ME

Tliis fish-collectinf? device is the revolvinp' trash screens of the Station
A power plant of flic I'acifif f!as and Electric, Company, located at
Third and Twciity-l liiid Streets in San Francisco (Figure 1).

The station di-aws c()()lin<i- water from the bay throus'h two concrete
tuhi's 10 feet in diameter, extendinji' outward for about 30 yards from
shore. At high tide the top of each is about seven feet under water.
Thus, at low tide the ojk'u ends of the tubes extend from 2 to 12 feet
beneath the surface and al high tide from 7 to 17 feet. It should be
noted that all of our samples were taken at high tide. The bayside open-
ings of the tubes are protected by guards of perpendicular fiat bars
(I inches x 2\ inches) -which are placed so that there is a 5|-inch
0])ciiing lictwccii adjact'iit bars. These guards prohibit the entrance of
flit" largci- iishcs. When the plant is pumping at its normal rate of
100, 000 gallons per minute, the water travels through the tubes at
about 1?T mpli. This sucks many small fishes into the openings, and they
pass subsequently through to the well in front of the revolving trash
screens (Figure 2). The plant pumps continuously through both tubes
and does not alternate.

FIGURE 2. The junior autlior hauling up the collecting net from the sump in front of the
revolving trash screen. Phofograph by Toshio Asaeda, November, 1954.

SOUTH SAN FRANCISCO BAY FISHES

273

The inoiu'i wire serooiis are I).', feet wide and 33 feet lii^li, witli i-'wch
square mesh. They rotate vertieally, passing over sprockets 3.^ feet in
diametci- ;it the 1(i|) and l)()ttom. At intervals of two feet the screens
are lun<i('d and ("(itiipix'd witli a metal strip or baffle four inclies wide,
which extends outwardly from the face of the screen in tlie form of a
flat scoop. As the screen moves upward, these baffles catch debris and
fish which liave been pinioned against the mesh by the force of the
incominir water. Tliis nuiterial is then lifted over the top of the re-
voIvin<i- screen and deposited in a trough of running water, whence
it is washed back into the bay.

During the suinnicf months fish yield at the screens is small. A haul
of 10 pounds during any day of this period is a good catch, and as
little as one or two pounds is not unusual. By contrast, in the winter
we have often taken as much as 150 to 200 pounds of fish from the
screen sump. The largest amount, 270 pounds, was taken on February
8, in.")!. However, these amounts seem insignificent when compared Avith
the two to four tons that have been taken from the Southern California
p]dison Company's power plant intakes at Hermosa Beach, California
(verbal communication, Mr. Ross McBride).^

32 (31 36 24 27 20 12 10 8 7 16 25

X

80

if)

Ll_

70

Ll.
O

60

if)

50

Q

3

40

O

CL

60

LJ
iD

20

<

CC

10

LU

^

0

65 9

689

68.7

73.1

56.4

liiijiiiiii

,„„„,

;;;!■; -ti!!!!;

VMM

30.8

30.9

1

36.7

iiiiiiiiiiij

;

20.9

11 0

78

18.0

::;::::::;;;

::::::::::::

iiiiiUiii;

\ ...

Jan. Feb. March Apr. May June July Aug. Sept. Oct. Nov. Dec.

MONTHS OF THE YEAR

FIGURE 3. Average number of pounds of fish taken per month from the revolving trash
screens at the Station A pumping plant of the Pacific Gas and Electric Company during the
six years 1945-1950. Figures in circles above each histogram indicate the number of trips
during the six-year period upon which each average month is based. A total of 248 collections

and 14,935 pounds of fish is represented.

2 This aiiKiiiiit of n.sh was u.suall.v taken <>nl.v after a buiUUip period of one to three
weeks. The most abundant fish was the queenfish, .Serip/(».s polilus. followed by
the white croaker, Genyoncnuis lineatus. :Many other species were present in
lesser numbers, but tabulations are not available.

'74

CALIFORNIA FISH AND GAME

rnfortmuitcly l(ir piihlic a(|iiaii;i. this ready soiin-o . t' fisli food and
live specimen apjx'ars to he (hn.iiu'd hy the chaii-iiij,' (l,.si;rn of .seitriis
at these iiistalhitioiis. Studies su<-h as that l.y Kerr (1953) on fisli
srn'.'nin<r d.-vic-s for p(,\v.T plants have inii.l.-nicnted the redesign ol
ohI.T iiistallatiniis to jnoviih' for more cCficifnt tish diversion.

Fi;:ure ;{ presents the avera«;e eateh in |)onnds jxm- month for the six
years HM:, thron-rh liir.O. This represents a total of 248 eoHeetin.' trips,
from w Inch 14.41:) livin- fish.-s were returned to Steinliart Acpiariiuii
tor display, with an additional 14. s:}.") pounds of dead lisli for tood The
live fish have not heen ineluded in the data for Fi«:ure ;j.

Identieal months in diffeivnt years are sometimes quite variahle For
•'xampie. in Fehruary. 1!)4!). the lar-est eateh was 10 pc.unds of fish
whereas in Fehruary. 1 !».->(). it was ]7A) pounds, and in Fehruarv. i:):)l!
L'/O p(.iiiids. However, on an aiiinial basis the monthly variation's tend
to Miiooth out and a fairly reju-esentative histo<rram results (Fij,Mire 3).

The (|nesti<.ii immediately arises re-'ardin<r the reason for hhdi pro-
duetK.ii diiriii- wint.T months and low produetion durinjr s'ummer
months. Temperature does not a|.|.(iii' to piovide the answer hut
st(.rms. ])revailin«r Minds, and tii.' phenomenal feeunditv of the vivip-
arous surfperehes do furnish a part of the solution. Duriu}; and imme-
diately after a storm the yield at the screens is increased sevei-al f<.ld,

TABLE 1

Comparison of Representative Winter and Summer Catcti Samples From the Trasli Screens of flie

Soutti San Francisco Bay Station A Power Plant of tlie Pacific Gas and Electric Company

Species

February 19, 1951

Shiner seapercli, CymatJijanter aggregata

Pilepercli, Hliocuch Hun vacca

Wullcje surfperch, H yperprosopon

arynitenm

Striped seaperch, Embiotica lateralis
Bay blackpercli, Jimbiotiica jackmmi.

L)\sarfperch, Micrnrnetrux minimus

White seai)ereh, I'liaiiirudon /urcatus._

Rainbow seaperch, J/ypsurus caryi

Rubberlip seaperch, Uliacuchilus tuxoles

Boccacio, Sebastodes paucispinis

Blacli rockfish, Sibastodts melanops

LioKCod, Ophiodun elonualua

Striped ba.ss, Hoccus saxatilig

Kntilisli sole, Parophrys vetulus .

San Francisco topsnielt, Atherinops

affinis affinis

Sacramento smelt, Spirinchus Ihalei-

ehthya

Tomcod, Microgadus proximus

Northern anchovy, Kimraulis murdax ^
Stauhorn sciilpin, Li ptucullus armatus

ar mains ..^

Leopard shark, Triakis semi/asciata. . .

California skate. Raja inornata

Detritus and iiiO|jods

Number
speci-

2,217
28

11
1(1
10
12
4
1

Weight

in
pounds

1
2
1
21
8

81.80
2.50

2.20
O.7.".
0.7.-)
0.50
0.25
0.03

Weight
percent-
age of
total
catch

August 13, 1951

12
1

Totals.

2,352

0.22
0.37
0.03
0.40
0.50

0.18

0.25
0.09

l.,50
0.50

87 . 53
2. 08

2.35
0.80
0.80
0..54
0.27
0.03

Number
speci-
mens

Weight

in
pounds

0.23
0.40
0.03
0.43
0..54

0.19

0.27
0.10

1.60
0.54

0.63

93.45

242
93

88
2

11
1
4

7

10

7

1

Weight
percent-
age of
total
catch

2.66
2.81

1.31
0.07
0.75
0.07
0.56

0.67

100.00

3.19
0.44
0.50
0.09

19.00
20.07

9.. 30
0 . 'M
5 . 36
0..W
4.00

0.12
0.56

0.31
0.56

481

22.79
3.14
3 . 57
0.04

0.86
4.00

2.21
4.00

14.00 100.00

SOUTH SAN FRANCISCO BAY FISHES

275

perhaps in l;ii"^-e part a result of the stoi'iii winds hlowini:- fisli lowjii-d
the intake tubes. Durinnj the sprin<i' and suiinnei- months, when the
female siirlperch drop their yonn^', there are a <ireat many small .jnve-
luh's in the catch. The snrt'j)erch i)opiilation during the winter is made
lip mostly of lar^c juveniles and adults.

In order to determine accurately the composition of the catches, 14
of the 24<s eollections. covering 9 of the 12 months (.Iul\- tlirougli ^larch,
inclusive), were broken down and detailed analyses made. Since the
basic pattern seems to be approximately the same in all of the months
during which samples were taken, we present in Table 1 the two
extremes — a wintei- sample (February 19, 1951), contrasted with a
summer sample taken six months later (August 13, 1951). As shown by
the total weight of each collection (88 pounds vs. 14 pounds), the
winter and spring is the period of maximum abundance and the
sinnmer and early fall is the period of minimum abundance fsee also
Figure 2). The shiner seaperch is usually the most abundant species.
In the fall the number of juveniles drops considerably, so that the total
poundage of this species is much less than in the spring. By contrast,
the numbers of some of the other species of surf perches, most of which
HVv adult fish, e.g., pileperch, rubberlip, and walleye, increase in the late
fall. From Table 2 it will be noted that the number of shiner seaperch

TABLE 2

Shiner Seaperch, Cymatogaster aggregata, Contained in 14 Catches From the Trash Screens of the
South San Francisco Bay Station A Power Plant of the Pacific Gas and Electric Company

Date

Weight of

catch
(pounds)

Number

of
shiners

Weight of

shiners

(pounds)

Shiner

weight as

percentage

p{ total

catch weight

March 11, 1949

January 10, 1951

February 19, 1951

July 31, 1951

August 13, 1951

September 11, 1951

October 4, 1951

November 30, 1951

December 26. 1951

February 8, 1952

March 2.5, 1952

July 22, 1952

August 29, 1952

Januarj' 30, 1953

140.00
82.00
93.45
14.00
14.00
1.87

1..50

145.16

111.44

.50.49

1.50.. 50

16.82

14.44

11.00

2,130
338

2,217

670

242

1

(juvenile)

2

189

553

722

289

459

219

138

97.00

12.00

81.80

4.81

2.66

0.002

0.02

7.62

23.43

24.19

17.25

4.62

5.56

4.80

69.28
14.63
87 . 53
.34.. 36
19.00
0.01

1.33
5.25
21.02
47.91
11.46
27.47
38.50
43.64

in the 14 collections critically analyzed ranged from 1 (total catch
weight of 1.87 pounds; September 11, 1951) to 2,217 (total catch weight
of 93.45 pounds; February 19, 1951). The weight of the shiners ex-
pressed as a percentage of the total catch weight for these same two
collections ranged fr'om 0.01 to 87.53.

The minimum number of s|)ecies of fishes in the collections was
usually 7 and the maximum 18, with the usual number 10-14. The
total of all species is 61, as indicated by Table 3. Since the two-foot
diameter net (21-foot handle) used in collecting at the trash screens
is made of %-ineh stretched mesh, many of the smaller fishes are not
taken in their true proportion. For example, the slender bay goby,

I'TU

CAL.1KOKMA FISH AM) IJAME

TABLE 3

Species List and Approiimate Number of Fishes Taken in the 248 Collections Made at the Trash

Screens of the South San Francisco Bay Station A Power Plaht of the Pacific Gas and

Electric ComiMny During the Su Yedrs 1945- 1950

8|>eciai

Num-
ber
of
spoci-
nicna

Species

Num-
ber
of
speci-
mens

Pacific Imtiprvy. Rntotptirnut tridentatui. .

7
1

nu

27
'2

i:»

*j

3

23

701

11

100

17

1

1

2

354

1

34

22

22

168

1

386

5

2

18

7

510
399
344
9
194

383

SrtcIiKill •oWhlinrW. \>iti>ryufhu» maculatum
I.«'<>I'nri| xlinrk, Tnakit trmifatciata
Hr..«ii ••iiiiMillihiiiiml. Hliinutrtaeia heiiUi
iJut^ti^h, S4jualuii fueklryi
Califurnia hkiitf. Haja inurnata

UulilH-rlip »eai«Tch. Hhacochilua tojcotea

Sinner (teapercli. Cymaioyaatrr ayyregata . -

Dwarfperch. Micrometrue minimua

Bocacrio. Srbaattidea pauciapinia .

192

Many

1,000

558

689

LoiiKi»>M- nkulf. Haja rhma rhma
Bat rev, Myliobatu califtirnteua

Vellowtail rockfish, Sebaatodra flatidxta

Black n»ckfi.sh. Sebaaiodta mclanopa

Brown rockfish, Sebaatodea nuriculatua

Gru.sa rockfish. Sebaatodea raatrdliyer

Black-and-yellow rockfish, Sebaatodea chry-
aomelaa

2
252

Kleciric mv. TurfKtIu caiifurnica

1

I'acilir licrrinK, Clui>ra polltui

3

Snriiiiu-, Sarditit>i)ii carrulea

NortliiTti Biifliow. Eniiraulin murdax

4

Wliileliait Miu-ll. Alloamrrus el'jnyatut

SucTHiiii-nto hiuflt, Sfttrmchua thaleichthya

Kelp RreenlinK, Hexaorammos decagrammua
LinKcod, Opftiod-jn eluirjalus _.

16
322

Surf Miiflt, llypomrtuii prttiuaua

Cabezon. Scurpaeniclithya marmoratua

Reef iri^iilord. Hemitepidotua apinoaua

Buffalo sculpin, Enophrya biaon ^..

Northern staghorn sculpin, Leptocottua
armatua armatua

12

Kinu liuliiiun, Oneurhynchus tnhawytscha

Tomc-ml. M tcri>ijadus prjximux.

1
38

I'acilii- ^un(lliltb. Citltarichtliya surdidus

Curlliii turbut. I'Uuruniclithya decurrena

<

Enitlinh Bole, Paru/thrys tetulus

Slipskin snailfish, Liparis fucenais

Show-y snailfish. Liparia pulchellua

Northern threespine stickleback, Gaatero-

ateua aculeatus aculeatua

Tube-nose, Aulorhynchus flavidum

Northern bay pipefish, Syngnathua griae-

olineata griseolineata

1,000
3
1

1
1

Starry Houiider, I'latichthyn ateUatut

Stri|«-<i liasM, Kiiccus aajratilia

Jackxiiielt, Atlitrriiiupeis cali/urnienaia

San Francisco topsmelt, Atherinopsia
affinia affinia

Jackiiiackerel, Trachurua aymmelricua

I'onipano, Paiumeta aimillima

88

White croaker, Genyunimua lineatua .

Bay goby, Lepidogobiua lepidua

1

HeiJtail fturf perch, Amphiatichua rhodoterua

Mudsucker, Gilliclithys mirabilis

1

Walleye surfperch, Hyperproaopon argen-
irum , .

Northern inidshipiuan, Purichthya notatua .

280
4

Bay hlackperdi, Embiotoca jackaoni

Rock-eel, Xiphister mucosus

1

•Stri|jed hcapcrch, Embiotoca lateralia

Saddleback gunnel, Pholis ornata

1

HiiMjow pereU, J/ypaurua caryi. .

Spotted cusk-eel, Otophidium taylori

2

White t>ea|jerch, Phanerodon /urcatua

which is quite common in many parts of the bay, has a maximum
h'n<:th of about 2Vo int-lies; thus, it generally .slips throuoh the dip net.

The occurrence of some of the less common fishes has been surprising.
Of nine specimens of slipskin snailfish (Liparis fucensis) known from
San Francisco Bay, three were taken at Station A. In contrast with
this, Station A has yielded only one specimen of the fairly common
showy snailfish (Liparis ijulchellus). In the south bay the brown
smoothhound is usually twice as abundant as the leopard shark; the
(hitj-fish is rather erratic in appearance; and the sevengill and soupfin
usually form ;"> to 16 jM'n-ent of the elasmobranch population (Herald
and Hipl.-y, l!).-,], p. :{17; Herald, IK.Vi, p. 241). At Station A we find
a icvei-.sal of this condition, f(U- during the six years there occurred
more leopards (140) tlian brown smoothhounds (27), while the dogfish
(2), and .sevengill (1) were both poorly i-cpresented, and the soupfin
was entirely absent.

Table 4 presents a lasting by months of certain fishes whose total
numbers for the six-year period i-aii'jrd between 50 and 1,000. In most
cases these fishes were returned to Sleinhai-t Aquarium alive and so do

SOUTH SAN FRANCISCO BAT FISHES

( I

B
O

— »^

.2 =
^^

« >-

— c

O 'O

<— > a.

I — *->

-o «

« O

E

01

a>

3 00

2 -ra

— Ol'N-fOOC-lOOOQOO
000<NiOOOOO>!7;£tOOO

U5 —

I w o

I ■* « -H C^ t»

I C^ -H T)<

o 00

I CO -r 'O a: 50

M r^ — — ' ;o

aoo

1-5

1 (M
1 (N

O IC

to 00

c o

3 (N

■-5

00 Tt N 1 ^ o o

-^ Ol CC ' CI CO

; ^

fc- CD

C3 CO

00 00 CO o

IN >0 CO M

.-i-^COtD'- — C^ rt

■^.-1 i^00tO00t~O5 lO
lO 1 t>. C^ CO 1 •-■

O -H

IN

I 05 to ■* CO 02 ■* I O

(M02 lOO it-00'* itO
Nin lOOQ IN — -^ i-H

a.

e

1= -

^ !o « 3 n -*; r^

3 -i

2

Q as

3 «

C a

» O

s«

si a

X .s£ -6

o s ■

•e T3

n

2.

'r'

o

o

fa

:2

-'<:

ti

5

=

u

2h

-/i<5

P5

« J-

. I-
cn S3

■=■ o

2 "~ 2

•2 S
■a e

s j:

Ol

<o

o a

S Ea

ty o :-

o o ~

o

278 CALIFORNIA FISH AND CAME

not appear in the data (tf Fi^rni-c ."{. Of tlu' spceios listed in Table 4,
only two, the Idaek nickfisli and the hoeeacio. uccnrred tlnrin^ every
month of the year. These were invariably yonn'j: horn a few months
earlier. Three sjjecies, the hrown ntcktish, lin<rt'()d, and hci-rin^-, oc-
cnrred dnrin^: 11 months of the year. The fact that San Franei.sco Baj'
is a nnrsery <.'roniul for two of these speeies. boecaeio and linjjreod, has
{generally been overlookecl. .Most of the species in Table 4 folloAV the
al)nndance liistnirram of Fij/ure 1. However, some exce]>tinns are (a)
tin- al)si'ni-e of midsliipmen foi- six months, Aii<:ust tlirou<;li January,
whieh is nndonbtedly correlated with the spawninjr cycle, and (b) the
absence of the northmi aiiclio\y (liii-ju'j- the tlwce months January
throiij.'h March.

In total numbeis the (il fish species from Station A compare favor-
ably with the ()() species wiiicli oiir as yet-unpublished records show for
the Chinese shrimp nets fisliin^^ in deeper water (4r)-()() feet) about two
anil oiu'Jialf miles nortlieast. From the north bay shrimp trawlers wlio
fish in water of 8 to 35 feet we luive records of 47 species. Detailed
comparative analyses of the catches made in these tliree areas will be
inrluiled ill a subse(|uent ]ia]>er on the sjiriiii]) fislierics and associated
or<:anisms.

SiitlHce to sa>- foi- tin- pi-esent that the list of 61 species from Station
A includes 11 that have not been taken in the Chinese shrimp nets.
These are: jackmackerel, '.'■rass rockfish, yellowtail rockfish, black-and-
yellow rocktish, lonjiiiose skate, Calil'oi-uia skate, tubenose, kinp' salmon,
rock-eel. noithern stickleback, and redtail snrfpereh. From the Chinese
nets, howevei-, there are 11 species that are not known from Station A.
These are: bonehead sculiiin, Arfrdius //(^/r>.s7;//o/«.s; padded sculpin, Ar-
tedins fenestra] is,- arrow <ioby, Clevelandia ion; river lamprey, Lam-
petra flitrlnl His ; lln-esher, AJopias nil piints .- hake, Merluccius product-
ns: (lover sole, M icrosfoniKs pucificKs: saudlance, A}iiii)odjifes fohiiDtufi
personal us ; anjifelshark, S(jii(itiii(i ctdifornica ; i)it-head jioacher, Odon-
topyxis trispinosa, and the rubber seulpin, Ascelichthys rhodorus. Inas-
nnich as these records of jiresence and absence deal Avith species that
are not abuiulant in the catches, their sijiuificance is not very preat,
excej)t as an indication of ditl'ei-euce between the two samplinu' sites.
The Chinese nets fish oidy on iiicomjip,;- tides and IVoiii a faii-ly central
position in tlie south bay. Thus, these nets should l)e expected to catch
some lisiies fi-om offshoi-e and deejier watei-. I'y contrast. Station A
operates continually, and we believe tli.it many of llie fishes appearin<>'
here are brought by shore curi'ents alon<>' the south side rather than
thi-ou-i-h the center of the Coldeu Cafe. AVhether or not this tentative
exjjlanation for some of the diU'erenecs in the catches is valid must
await further studies.

REFERENCES

ll.Tiil.l. K.irl S.

V.iTi'A. 'I'll.- M)~)2 shail< dcrliies at Elkhorn Sloiifili. .Mcpiiterey Bay, and at Coyote
rc.iiit. San rrancisco Bay. r.-ilif. Fisli .ind C.inic, vol. ?,U. u>k 2. p. 2:-.7-243.
Il.iMld. Karl S., and Wni. Kllis Riiilcy

r.i.'l. 'riic r<'lati\c ahiindanco of sliarks niid liat stingrays in San I'"rancisc(> Bay.
("alif. Fi.sli and (lame. vol. 1^7, nn. ;;., |). .'U ■")-.■ >2!).
Ki'iT. .lanics E.

I'.i.'i:;. Stndics on fish prcsci-v.-ition at the Contra Costa steam jilant of the Pacific
(las and Klt'clric Company. Calif. Dept. Fish ami Game, Fish lUill.
92, GO p.

TOLERANCE OF THE SAN FRANCISCO TOPSMELT,

ATHERINOPS AFFINIS AFFINIS, TO CONDITIONS

IN SALT-PRODUCING PONDS BORDERING

SAN FRANCISCO BAY'

LARS H. CARPELAN
University of California, Los Angeles

INTRODUCTION

The topsmelt, Atherinops affinis, is tolerant to a wide range of salin-
ity: it lives in fresli and brackish water, in the sea, and in salt concen-
trations greater tlian in the ocean. Dnring a year-ronnd ecological study
of five salt-producing ponds bordering San Francisco Bay, the San
Francisco topsmelt, Afhcrinops affinis affinis (Ayres), was observed
to si)awn in water with twice the salinity of the ocean. Since it does not
live the year around in j)()nds of such high salinity, concurrent records
were made of 1lie development of the young fisli. Jiiid of the seasonal
changes that took place in certain factors of the environment. These
factors Avere : pH, concentration of dissolved oxygen, temperature, and
salinity.

From the data obtained, it is not possible to determine the exact
iippei' limits of tolerance to the factors measured. However, since there
seem to be few records of the actual environmental conditions that
prove limiting to fish, this is presented as an account of the extreme
conditions under which a school of Afhcrinops developed, and existed
until cniiditions exceeded the upper limits of its tolerance.

The ponds under study are located near Alviso, at the soutliei-n
end of San Francisco Bay. These Alviso ponds are early in the evap-
orative series. Afhcrinops lives the year aroiuid in the first three ponds;
ill these the mean salinity ranges from 27.5 parts per thousand in the
tii-st to 45 parts per thousand in the tliii'(|. In the fifth |)<)n(l. which has
an annual range of salinity from (il ])ai'ts per tlionsand in tlie winter
to as high as 150 parts per thousand in the fall, Alhcrinops does not
live throughout the year. It may, however, spawn there in the spring.

Coneiirreiit data were recorded on the growth of fish in a school
spawned ;it the inlet of the fifth pond, and on seasonal changes in
certain environmental factors. The young fish lived from April until
August. The school then disajipeared, presumably because of intoler-
ance to the seasonal change in some factor or factors of the environment.

1 Submittert for publication February, 19.55. The work was done at the Hopkins
Marine Station of Stanford University and was aided by a fellowship from the
Leslie Salt Company.

( -*T'." I

280 CALIFOKN I A I ISH AND OAME

METHOD OF FISH SAMPLING

Duriiij;- the I'oiu'sc oi' the ('L-ul();^ical sludy, plankton t-oUectiuns were
made at weekly intervals, nsinp- a net of nine inch diameter, No. 12
silk boltin*;: cloth, 120 meshes ])er iiu-h. Fisli taken by this means were
incidental to the main stndy. Tlie method was not well adapted to the
collection of flsli, and did not take the faster-inoYin<i', larger fish.

The fish of tlie salt ponds were not intensively studied. It also seems
necessary to point out that the samples enumerated below were taken
from a school of fish, so that tlie number i)er volume are not repre-
sentative of the fish popnlation throughout the pond.

BIOLOGY OF ATHERINOPS

From March to August, plankton eollections from all stations in the
ponds contained j^oung topsmelt. Eggs were eolleeletl as early as March.
Spawning thus seemed to begin in Mareh and to reach a peak in April
and Maj', when many larval fish were taken. However, the presence
of fish 5 nun. long as late as August indicates a long spawning season.

During the long spawning season, there was considerable range in
size of fish at most collection stations in the Alviso ponds. For example,
the following is a tabulation of collections taken in the third pond :

Atherinops Collected in Midpond No. 3, 1951

Average number
Month Size range in mm. per 4 AP

February — 0

March 8 1

April 5-12 9

May 5-18 26

June 10-30 39

July 15-30 37

August 5-25 2

September — . 0

However, topsmelt apparently spawned over a shorter time interval
at the inlet of the fifth pond:

Atherinops Colleded at Pond No. 5 Inlet 1951
Month Size range in mm.

April 5-10

May 10-30

Juue 25^0

July 35-55

August 30 (onefisii)

According to Schultz (1933) the various subspecies of Atherhiups
afflnis on the Pacific Coast are from 65 to 100 mm. long by the end of
their first year, the greatest size being attained by the most northern
subspecies. The growth of the Atherinops in the salt ponds under study
seems more rapid than that reported by Sehultz for A. affinis affinis,
presumably the subspecies in the ponds. Schultz reported 70 mm. as
the average length reached by the end of the first year. In the salt
ponds that length was attained by August. Fish in August collections
in the first of the Alviso ponds ranged from 60 to 80 mm. By the end
of the year the average length was from 90 to ]00 nun.

TOPSMELT IX SALT-ri;iil)l ( IN(; I'ciMDS '2.X\

Topsnu'lt of the s;ilt jjoiuIs i-c.-icIkmI scxnnl mat iirit_\- when in thoir
second year; that is. fish tialdicd in April spawned tlie t'ollowiiii;- April,
wiien one year old. Tln'ir Iota! lent:! 1 1 when mat n re was 10(1 to lit) mm.
Fisji larger than this wei-c not ol)sei'\'e(l in the ])onds.

MORTALITY OF ATHERINOPS IN POND NO. 5

Although topsmelt are present tlirouj^lnnit the year in the first three
of the Alviso ponds, they are not present in later ponds dnring winter.
Beginning in April or INFay. when the water levels of the ponds fall
due to evaporation, watei- is |)niii|)ed fi-om jiond No. ^ into the later
ponds.

When pumping operations began, in eail.v April of l!*;")!, mature
fish, about 100-110 mm. in lenutli, which had wintered in the third
pond, found their way iido tln' foni'th and into the irdet of the fifth
pond.

At the inlet of pond No. 5, gravid females spawned in water which
then had a salinity of 72 pai-ts per thousand. The eggs hatched, giving
rise to a large school of fish which jiersisted at the inlet of the pond
from April until July, with the increase in size shown previously.

Tabulated below are data showing the size of the population, and
some of the changing environmental conditions to which these fish were
subjected at the inlet of the fifth jiond.

Atherinoiis Mean Mcait initer

(average no. salinity Mean pTf temperature

Month pcr]M') (p.p. 1000) (P.M.) (P.M.)

Mnnli 0 fi.S.l' S.S I'.C C

.\pril 36 72.3 !).0 21° C

-May 05 73.1 S..! 23° C

June 112 SO.O S.3 24° C

July 48 OO.O 8.5 27° C

August 1 II3.1 8.3 26° C

September 0 02.2 8.3 2.">° C

The data above and the size range of th(> fish are illustrated in
Figure 1.

The tabulation above is of monthly means, based on weekly collections
and samples. A great change, not apjiarent in the table, was the number
of fish collected during the first two weeks in -Inly. The number de-
creased from 227 per (piai'tei- cubic meter on .luly 2d to five per
quarter cubic meter on .luly Dth. The large school of fish virtually dis-
appeared.

DISCUSSION OF FACTORS CONCERNED

Various en\iiM)nniental factors (Figure 1 ) in the ponds change sea-
sonally; one of the following, oi- some combination of them, may be
implicated in the disapjx'arance of the fish.

Salinih/. The eggs hatched in water with a total salt content of 72
parts per thousand. The salinity increasiMl L;i'adiiall.\ until it was 85.5
parts |)ei- thousand on -Inly '2t\. DuriuL;' the week I'l-om .Inly I'd to Dth.
it increased greatly to IM).2 j)arts pci- thousand. This could be in-
terpreted to mean that somewhere in the range between 80 and iM)
parts per thousand conditions hecame intolerable for Athcritiops. How-
ever, it is impossible to estimate the exact upper linnt of salt tolerance

282

CALIKOKMA FISH AND GAME

1201

ri2o

FIGURE 1. Histogram: Average number and size of Afherinops in collections from

Alviso pond No. 5 (inlet), 1951.
Graphs: Seasonal changes in S / '", pH, and temperature.

from the data. Furthonnorc, increase in osmotic pressure due to salt
concentration is probably not the only factor involved.

Hydrogen Ian Concoitratiox. The table and the figure sliow the
decrease in pll that took place during the period concerned. The peak
in April was associated with uptake of carbon dioxide during a spring
bloom of the alga Hiichococcus. After this bloom, tlie pll decreased to
a value more nearly like that of the oceans, but in this case, the decrease
indicates a loss of carbonate. In the salt ponds, the increasing salt

TOPSMELT IN SAI/r-l>K()DUCING PONDS 283

concentrations event ii;ill\' lead to the pi'ecipitation of salts (the {)nri)ose
for wliieh tlie ponds are maintained). Caleinm earbonate is the first salt
to ])reeipitate ; it hej^ins to come out of solnlion at salinities above
about 80 ]iarts per thousand. Loss of (•ai-bonat<' not only increases the
hydrogen ion concentration, but creates an upset in the ratio of ions
present in sea water. Cakdmn as well as earbonate is lost from solution.
This change in ionic ])i'opoi-t ions may, in some way, ])c i-esponsible for
the iiuibility of Athcrinops to tolei-ate salinities beyond 80 parts per
thousand.

Temperature. The temperature of the water, though rising during
this period, was no higher than in the other ponds, in which Atherinops
continued to live. Athcrinops is remarkable in its tolerance to tempera-
ture changes. The use of means in the table covers up great diurnal
temperature ranges. For example, the extreme diui-nal range observed
in the ponds was 12° C, from a morning low of 21° (69.8° F.) to an
afternoon high of 33° (91.4° F.) ; tliis occurred in August. Ailierinops
must also withstand an annual range of 25° C. : f i-om a low of 8°
(46.4° F.) in January to as high as 33° (91.4° F.) in -Inly and August.

Dissolved Oxygen. Not tabulated is another fa(*tor that should be
considered. The concentration of dissolved oxygen may become a limit-
ing factor because of the inei-eased salt concentration. Water with a
salt content two and one-half times that of ocean water can hold only
about one-half as much oxygen, at saturation in equilibrium with air,
as can be held by sea water of ocean salinity. Because of the oxygen
evolved by algae during photosynthesis, evening values for dissolved
oxygen in pond No. 5 are above saturation. (There may be as much as
six or seven milliliters of oxygen per liter of water.) However, morning
values are below saturation, due to the respiration of all the organisms
])resent, without compensating oxygen production by the algae at night.
Very low concentrations of dissolved oxygen at dawni may limit the
continued existence of Atherinops in pond No. 5.

Comhination of Factors. No single factor can be isolated as the
factor limiting Athcrinops to salinities below 80 parts per thousand.
But the combination of factors (high temperatures, high osmotic pres-
sure, an upset in the ionic ratio, variable oxygen concentrations) finally
becomes too miicli for e\iMi so liai-dy an organism as the topsmelt.

Differences in Tolerance With Age. There is at least one other factor
that might be considered. The school of fish was of uniform age. It
may be that these ycnuig topsmelt are more tolerant than older ones,
and that the disappearance of the school might be due, not to changes
in the environmental factors described, but rather to attainment of a
certain age or size at which the fish may become less able to withstand
the environment. However, the fish have not been studied physio-
h)gically for such diiferences, noi- have they been studied for possible
morphological changes which might have been induced by tlie extremes
of tlie environment to wl)ieh they wer(> (>x]ios(>d during their d(>v(do]i-
me]it.

SUMMARY

Athcrinops, the topsmelt, is the most salt tolerant fish observed living
in ponds used for the prodm-tion of salt by cvapoi'al ion of sea water.

284 CALIFORNIA FISH AND GAME

It lives throufiflinut tlie year in {)()iids with a salt content as great as
one and one-hall' times tliat of ocean water. It has been observed to
spawn in salinities double thai of the ocean (72 parts per thousand)
and to exist in still lii^'her salt concent rations.

In ad(liti(jn to hij^li salinities, Atlurinops in the salt ponds must also
witlistand other environmental extremes, iueludin<): high summer tem-
peratures (with a great diurnal range), higli pll values, and variable
concentrations of dissolved oxygen.

LITERATURE CITED
Schultz, Leonard P.

1933. Till' aj,'e aii<I jjrowth of Atherinops affinis oregonia Jordan and Snyder and
of other subspecies of l)aysnie]t alonj; the Pacific coast of the United
States. Wash. Univ.. Puhl. I'.iol., vol. L'. no. 3, p. 45-102.

AGE AND LENGTH COMPOSITION OF THE SARDINE

CATCH OFF THE PACIFIC COAST OF THE

UNITED STATES AND MEXICO IN

1954-55'

FRANCES E. FELIN and JOHN MacGREGOR

United States Fish and Wildlife Service

and

ANITA E. DAUGHERTY and DANIEL J. MILLER

Marine Fisheries Branch

California Department of Fish and Game

This i-cport on age and length composition of the catch of sardines
(Sardinops caendea) off the Pacific Coast of North America is the
ninth of a series which gives similar data from 1941-42 to the present
season (Felin et al., 1954).

During the 1954-55 season, 65,918 tons were landed in the San
i'edro region; this included fish landed at Santa Barhara, Port
llueneme, Santa Monica, Los Angeles-Long Beach Harbor, and New-
])<)rt. Fish from some of these points were trucked to Monterey, Moss
Landing, and San Francisco for processing during October, November,
and December. At San Diego 510 tons were landed and trucked to
the Los Angeles area for processing. In August and September, 830
tons were landed at Avila and trucked to Monterey and Moss Landing
for processing. No fisli were landed at Monterej^ Moss Landing, or
San Francisco.

The fishing season at San Pedro and San Diego, as for the several
preceding years, extended from October 1st to February 1st ; that in the
Monterey region extended from August 1st to January 15th. For Ense-
nada, Baja California, where fishing is carried on throughout the year,
data for a period approximating the California canning season only
were used — in this case, July 30th to January 7th.

Sampling was carried on, as described for recent preceding seasons,
at Monterey, San Pedro, San Diego, and Ensenada. Sampling and
age determinations were continuecl by personnel of the California
Department of Fish and Game and the U. S. Fish and AVildlife
Service.

Tables 1-5 show, by sex and area of catch, the length-frequency
distributions of sardines of each year class as taken in the 1954-55
random scale samples.

Table 6 gives calendar dates for the lunar months included in
Table 7.

Table 7 gives total tonnages and numbers of fish of each age caught
in eacli region, estimated in tlic same inaiiiicr as described for preceding
seasons.

' Submitted for publication April, 1955.

(285)

286 CALIFORNIA FISH AND GAME

Taljlc !^ pivcs. hy sex and repion of oatoh, tlio moan lonp-fli and
istandard error of the mean for eaeli year class sampled in the I'Jo-l-oo
season. These are based on the random scale samples.

REFERENCE

Feliii, Frances E., .lolui .M;icC;ri'j,'ur, Anita K. ] )augliert.v, and Daniel .7. Miller.
1954. Age and length composition of the sardine catch off the Pacific Coast
of the I'nited Stat.-s and Mexico in l!»r):{-.")4. Calif. Fish and Game,
vol. 40, no. 4, p. 4Li;!-4:il. (See this report for preceding references.)

SARDINE CATCH, 1954-55

287

>-

CO to

e
o

Eh

— -hoOiO'MM— ''><0: — 000-f'Mc»3— '

-t<

« 01 M M ti M CO -M -- —

'O

■0 _

CI

So

&<

cc 00 -r 0 •M -f -M 1^ -c CO cc — •ri — '

— CI — CI —

CD
CI

O*'

""^

ie-(

— — 00 3-. -rt^ — p'OOicc-rci — — '-H

00

<;

CI

rt

.

rt(N-<TjiOM^»M — —

^

?,

■a

01

0

t<

^ CI CO CI ?l 71 ^ ^

-r

s

w

S

— CI CI CO CI -H

CI

1— 1

H

.-c cc-t<oot^0500a5 — 0500>-«'cieo-^

00

2

^rt^CICOClCO^^

CI

'S _.

CI

0 S

r.

10 '-^ CO 00 0 Cl Cl CD CO t^ CO -1 01 .-

CI

C-1

^2 0

►— <

IN — CI rt

»— 4

3)

r

>a

<-> croiClTlirt00^o;c0-1<CI— 1— irt

CD

(^

f-H f-i CI i-H ^

"u

C3

0

H

rt rt rt CI C) -f — 1 -H

CO

0)

a;

IM

Q

tn

^ - CI ^

CD

ira

c

05

e4

OQ

s

rt ,-1 — . CI CI

t~

2

H

■-H iraeot^>or^^>'5-HO!050>coc>co-H

Tf

i-(i-Hi-(ClcO<NCOt-i

^H

T3

CI

<1)

fa

Tfi CD ci 00 00 -J ci cc c r^ CI — ' CI — 1

rt — ■ CI — ■

0

c

eg

1-1 ino5-<coojcD^ajeococi^rt.-i

2

>.

H

1— 1

^

CJ

u.

il

fa

0

^

s

s

rt

fH

•3

H

rt -H CI

'I'

0

■•J

-a

fa

.- -H CI

'?'

c

?

I— 1
0

L.
0

S

bti

es

n

03

H

f-H

•~

«

T3

c3

C3

■3

fa

1— 1

—

^

in

IS

B

>>

w

§

CO

05

0

H

— CI

CO

fa

— CI

CO

c

eg

t.o 0

a) aj _h

« w, "^

>> a„"

h

'-iCOiCCOO-t'CO-H

•If

0

f vT"

CO

!2 " =

-■oOJ

•B

1

■itE

§gi

0

OS

'fccoooci-rcDoo cDoooci-fcDx^ci-rcrooociTcroooci
eococo-r-r-f-r-r 00003-. 3-. c:C5^. c^oco — — — — ■ — cici

rt rt —. — ,-. rt « _H .-.-« — -. -H — — CI CI CI CI CI CI CI CI CI C) CI CI

M CI

N (N

H

288

CALIFORNIA FISH AND GAME

TABLE 2
Length Composition of tlie 195t Year Class in 1 954:55

Ml

">1 year <

•lass, atic 3

Standard
length Mini.

Monterey

San Pedro

San

I )iego

California
total

Ensenada

Grand
total

M F

T

M F T

M

1' T

M F T

M

F

T

M ]■■ T

186

1

1 2

1 1

2

1

1

1

1

3

2
2
1

1

1

1 1

2 2

3 3

4 2 6
11 3 14
14 2 16

10 4 14
17 11 28
It; 13 29

8 8 16

11 17 28
7 12 19

6 10 16

7 8 15

5 3 8
2 5 7

1 7 8

2 6 8
1 2 3

1 1
1 1

1
1
1

1
1

1
1
1

1 3

1 1

1 1
1 1
1 1

1 1

2 2

3 3

4 2 0

12 3 15

15 2 17
11 4 15
17 11 28

16 14 30

8 8 16

13 18 31

9 14 23
6 10 16
8 10 18
6 5 11
2 6 8

1 9 10

2 7 9
1 4 5

1 1
1 1

1
3
1
1

3

1
1

1

4

3

1
1
1
1

1
1

1
1
7
1
4

4
1
2
2

1
1

1 1

188

2 2

190

3 3

192

4 2 0

194

12 4 16

196

16 2 18

198

14 8 22

200

18 11 29

202. . --

17 17 34

204

8 8 16

206

16 19 35

208 - .

9 15 24

210-

7 11 18

212 .-

9 11 20

214

216

6 5 11
2 7 9

218

1 9 10

220 ---

2 7 9

222

1 4 5

224.

1 1

226

2 2

Totals

2 9

11

128 115 243

7

5 12

137 129 266

11

14

25

148 143 291

SARDINE CATCH, I954-55

289

TABLE 3
Length Composition of tlie 1950 Year Class in 1954-55

l!l."iO year

■lass, aco 4

Standard
li'iintli mm.

Monterey

San I'c

Iro

San 1 )iogo

California
total

Ensenada

Oranfl
total

M F T

M F

T

M F T

M F T

M F T

M I- T

190

2 1 3

3 3

1 1

1

2
1
2

2 1

3 1

3 1

4 1
6 1
2 4

5 3

2 2

3 5

4 5
1 2

4

4

1

1 1

1

1

2
1
2
3

4
4
o

7
f'l
8
4
8
9
3
4
4
1
2

1

1 1

1 1

2 2

1 1

2 1 3

2 1 3

3 1 4

3 I 4

4 1 5
6 I 7

4 5 9

5 3 8
5 2 7

3 5 8

4 5 9
1 3 4

4 4

4 4

1 1

1 1 2

I 1

1 1
1 1
1 1
1 1 2

1 1

2 2
1 1
1 1

1 1

1 1

1 1

192...

194

19H _

198

2 2

1 1
'' 1 3

200

202

2 1 3

204 .-

3 '> 5

206

4 1 5

208

210.

.■) 1 f,
7 •) 9

212 .. .

5 5 10

214

7 3 10

216

(Jig

218

220

222-.
224

22t;

228

230

232

234

236

4 5 9

4 5 9

1 3 4

5 5

4 4

1 1

1 1 2

1 1

238

1 I

240

Totals

5 2 7

42 39

81

1 1

47 42 89

8 4 12

55 46 101

2—20860

290

CALirORMA FI.SII -\XI) GAME

TABLE 4
Length Composition of tlie 1949 Year Class in 1954:55

1949 year class, age 5

Standanl
length mm.

Monte

rey

San Pedro

San Diego

California
total

Ensenada

Grand
total

M F

T

M F T

M F T

M F T

M F T

M F T

190

1

1 1

1

1

1

o

1

1

1 1

1 1
1 1

1 1

2 2

2 2
4 4

3 2 5

4 1 5

1 2 3

2 2

1 2 3

2 1 3
1 1
1 1
1 1

1 1 2

1 1
1 1

1 1
1 1

1 1

1 1
1 1

1 1

2 2

3 3

4 4
4 3 7
4 2 6

1 2 3

2 2

1 3 4

2 2 4

1 1

2 2
1 1

I 1 2

1 1
1 1

1 1

1 1 2
1 1
1 2 3

1 1
I 1 2

1 1
1 1

192.

194 .

196

1 1

198.-
200..
202..
204..
206..

208

210

212

1 1 2
1 1

1 1

2 2

3 3

4 4

5 4 9

214

216

5 2 7
2 4 6

218

2 1 3

220

222

224

226

228

2 4 6

2 3 5

1 1 2

2 2

1 1

230

244

258.

1 1 2

1 1

1 1

Totals

2 3

5

27 13 40

2 2

29 18 47

5 7 12

34 25 59

SARDINE CATCH, I954-55

2m

10

'a-

ru

CO

■el 00
I — -"a-

o
o.
E
o

en

c:

1

"3

H 1

— '

0

-^ •— ' .-1

M

T)

b

C

2

S

^ « -^ — . — C4 ^

00

0

l^

1

&

0]

1— 1 —

71

w

_trl

h

00

.2

a

"o

fe

Ih

CO

C3

c

«

U

>>

^

05

I— 1

0

H

^_^„„ -l^^^

01

«"

, ■— t l^-l

CO

Ph

fc.

C

CO

s

■"* — —

0

^

cc ^ ic ^ ^ 0: t^ rc c; CO — cc >o — ■ --• 'M f-H ^

r^

-^

H

— ,— ,—.—.

0

0

■*^

^ ^ ^ Cl CC OC >C lO GO Oi ■M -r ^ C-l rH —.

05

"^

tu

'I'

0

s

(M^-t ^t-'ro-roooO'*co<N

00

,-r ^ (M C^l (N (N ^H

^M

d

El

*~*

^M t— ' •— < ^1 1— 1 >— t

t^

s

fe

•*

co^-t-— ..-^air^^it-- — -rccrc'j:— ^ ^^

0

2

H

05

CO

"2 „

01

fa

- -iccro-f-t-^cr. --r-- ^-^

71

d

_s *"

s"

0

s

c^i ^ -t — t^ -r c; cc 1^ 00 -^ 01 CI

.5

^

w

71

a

H

00

.2

■^

X

<N

71

en

■-^

fa

S

_2

E-i

cOrtTt4^.-<c:w — t^05'-*cococo^^ •-•'-^

01

00

)r,

0
Pl,

fa

^ — M?cco-t-rcoc. ---r^^ ^^

00

c

tl

S

CI ^ -r — t^ CO X CO 0 X ^ ci c^i

0

1

1

• - -H (M -

1/3

>>

Eh

a>

k.

0
1 c

fa

-"

— '

1 0

1 <5
i

S

^^ ^- C'l

tT

<

.3 =
It

0

1 1 1 1 1 r 1 r 1 1 > ■ > 1 1 ) 1 I • •
1 t 1 1 1 1 1 1 1 1 1 1 1 1 I I 1 1 • •

1 00 0 ?1 'I' 0 00 0 Ol Tf CC 00 0 <M ■* 0 00 0 (M ■* to

C-. COOOO — — — '-^-^c^iiMCincMcommco

JO

"3
0

— 71 :^l 71 71 n 71 7) 71 71 Cl 01 IN <M C-l 04 IN C^ <N (74

292

CALIlOliNIA FISH AM) (iAME

TABLE 6
Calendar Dates of Lunar Months tor the 1954-55 Season

"July" July 15-August 13

"August" August 14-Septeniber 11

"September" September 12-October 11

"October" October 12-November 9

"November" November 10- December 9

"December" December 10-January 7

"January" January 8-February 7

TABLE 7

Age (Year Class) Composition of the Sardine Catch in the 1954-55 Season
(Numbers of fish are given in thousands, i.e., 000 omitted)

Catch

Number of fish by age (year class)

Tons

Nos.

0

1

2

3

4

5

6

7

1954

1953

1952

1951

1950

1949

1948

1947

Monterey

"July"i -.- -

29

579

220

1

1

181

3,642

1,375

5

4

--

--

6

121

40

71

1,433

541

2

1

42
850
321

1
1

33

655

247

1

1

29
583
220

1
1

"August"

Totals Monterey- ^_

San Pedro

"September"'

"October"

830

4,110
35,026
22,296

4,309
177

5,207

30,473
270,390
179,184
34,065
3,766

3,766

2,294

173

8,229
91,067
61,209
11,947

2,048

10,208
101,235
08,663
11,173

1,215

4, '723

30,716

17,202

3,134

937

3,961
16,050
11,754

1,4.55

834

2.895
29,202
15,625

5,859

457
1,514

"November"

2,437
497

Totals San Pedro ..

San Diego^

"October"

65,918

42
267
201

517,878

326
2,047
1,610

3,766

2,294
"26

172,452

110
700
565

191,279

122
785
528

55,775

37

196
148

33,826

20

134

69

53,581

35
178
277

4,905
2

"No\-ember"

28

"December" _

23

Totals San Diego . -

510

3,983

--

26

1,375

1,435

381

223

490

53

Totals f'iilifiiriiia- -

Knsenada'

"July"'

"August"

"September"

67,2.58

865
1,394

993
1,112
3,664

679

527,068

6,.539
10,541

7,510

8,407
27,699

5,133

3,766

2,320
484

174,000

2,747
3,560
2,907
2,018
6,833
1,266

194,762

2,223
2^328

2,090

13,110

'2,430

.57,371

547
2,666
2,354
4,240

787

34,986

1,569

1,916

969

336

1,108

205

54,905

2,190

484

1,009

1,848

342

4,958

"October". .

"November" _

554

"December"

103

Tot-ils Baja Cali-
fornia _

8,707

65,829

484

19,331

22,781

10,600

6,103

5,873

657

Grand totals

75,965

592,897

3,766

2,804

193,331

217,543

67,971

41,089

60,778

5,615

' Includes deliveries August 1-13 only.

'^ Ineludes deliveries Octiiher 1-11 uiily.

=* Numbers of fish in the San Uiego catch were prorated at the same ratio as the San Tedio catch for the comparable

months.
* Numbers of fish were prorated according to the average weight of fish taken November 3-23.

SARDINE CATCH, 1954-55

293

TABLE 8

Number of Fish, Mean Length, and Standard Error ot the Mean for Each Year Class in the
1954-55 Season, by Region of Catch

Monterey

San Pedro

San Diego

Ensenada

\ ear Clas8

1

No.

M.

S.E.

No.

M.

S.E.

No.

M.

S.IO.

No.

M.

S.E.

1954

Malo

--

--

--

Female

._

_.

..

_^

--

Total

--

--

--

30

141

4.84

--

--

--

--

--

--

1953

Male

..

..

..

0

0

Female

..

..

3

196

2.00

1

Total ...

--

--

--

3

196

2.00

--

--

--

1

--

--

19512

Male

1

108

201

0.52

7

199

1.36

12

200

2.70

P'einale

0

100

205

0.51

6

202

2.76

14

201

1.32

Total

1

--

--

214

203

0.39

13

200

1..50

26

200

1.41

1951

Male

2

128

202

0.65

7

203

3.02

11

203

1.63

Female

9

213

2.00

115

208

0.69

5

216

2.92

14

205

2.34

Total

11

212

1.67

243

205

0.50

12

208

2.68

25

204

1.48

1950

Male

5

214

0.98

42

210

1 . 33

0

_.

8

212

1.44

Female

2

39

218

1.33

1

_.

..

4

218

6.40

Total

7

215

1.37

81

213

1.04

1

--

--

12

214

2.26

1949

Male

2

27

213

1.84

0

..

__

0

217

2.16

Female

3

217

4.38

13

223

3.34

2

..

._

7

215

2.76

Total

5

214

3.06

40

216

1.79

2

--

--

12

216

1.80

1948

Malo

4

214

1.50

50

213

0.95

0

..

4

216

2.16

Female

1-

..

39

217

1.19

2

._

7

217

3.02

Total

5

214

1.47

89

215

0.78

2

--

--

11

217

2.00

1947

Male

..

_.

6

217

2.04

_.

..

..

2

Female

3

220

4.1C>

__

..

..

0

__

Total

--

--

--

9

218

1.84

--

--

--

2

--

--

NESTING STUDIES OF DUCKS AND COOTS
IN HONEY LAKE VALLEY'

E. G. HUNT and A. E. NAYLOR

Game Management Branch

California Department of Fish at.d Game

INTRODUCTION

Durin<i- the spring ot' 1951 and 1!)"):] studies were made to determine
the status of nesting ducks and coots in Honey Lake Valley, Lassen
County, California.

Tlu' iii;iiii (il)jeetives of the studies were to obtain information on
nesting ])(i|)\ilations, nesting density, preferred nesting habitat, nesting
success, and the production derived from successful nesting. The 1951
study was the first survey made on nesting ducks and coots in the
valh'v. The 195.'3 study determined what changes had occurred in nest-
ing actixitics and also supplied data additional to those obtained in
1951.

Sample areas were established and utilized during each study, and
nest histories were completed on all nests found in these areas.

The results obtained in both years were similar in most eases. The
most apparent difference was a shift in the bulk of nesting from dry
upland areas in 1951 to marshy areas in 1953. The average nesting suc-
cess for l)()th years was approximately 50 percent for ducks and 96
percent for coots.

Brood data showed a slight reduction in the brood size of ducks and
an extensive reduction in the brood size of coots during the first week
of life.

ACKNOWLEDGMENTS

Tlie authors wisli 1(» thank several individuals for tlicir lielp. John K.
LeDonne assisted with the collection of data in 1951 and Robert Kirk-
patrick and Charles Victor Oglesby assisted in 1958. Tlianks are ex-
tended to the following memboi's of the Department of Fish and Game:
M. E. Foster, Refuge Manager of Honey Lake Waterfowl Management
area; Cliff a E. Corson, who prepared tlie map and graph; and A. "\V.
^rillci-. wlio assisted Avith tlie preparation of tli<^ maiinscri]it.

LOCALE OF THE STUDIES

The study areas were located in Honey Lake Valley in southeastern
Lassen 'County (Figure 1). Most of the preferred waterfowl nesting
habitat in this valley was in an area along the lower reaches of the
Susan River, from the mouth of Willow Creek to the river's entrance

> Submitted for publication INIarch, 1955. Federal Aid in W'ildlife Restoration Act,
California Project 30-R, "A Study of AVaterfowl in California."

( 205 )

296 CALIFORNIA FISH AND GAME

into lldiicy I.iiki'. Diiriiiiz- the wcl cycles, when Honey Lake is lilUnl to
cajiacity m- ncai- capacity, siiiialile n('stiii<r habitat is available from the
inoiitli (if the Siisaii Ki\ei- west ahuiL;' the lakeshore luf approximately
five mih's. A lurtiier descfipt ion ol' waterfowl nestin<r areas in the
vallev has been i)ublislied previously (Xaylor, 1LI33 ; Xaylor and Hunt,
1954).

Apj)roximately 80 percent of the nesting population of ducks and
coots in the Honey Tjake A'alley was found alon;^: the Susan Kiver and
its diversions frt)ni Litchfield to Honey Lake. It was in this area that
the studies were located.

The ]!'.") 1 study Avas conducted by Naylor on the Fleming unit of the
state-owned Honey Lake Waterfowl Management area. The second
study was conducted by Hunt in 1953 on the Fleming unit and on
j)i-i\'ately owned land.

HISTORY

The water level of Honey Lake throughout the years has been char-
acterized by fluctuation, determined mainly by wet and dry climatic
cycles. The lake has contained Avater several years after filling, but at
times i-everfs to a dry alkaline lake bed. When full the lake covers
approximately lUU square miles and has an average depth of 18 inches.

The main source of water for Honey Lake is the Susan River drain-
age. The water in this drainage is either stored for domestic use and
irrigation or allowed to flow into the lake. The amount of Avater that
eventually enters the lake is determined by the amount of spring runoff
and the demand on the Avater supply. Only during the years of ex-
tremely large spring runoff does an appreciable amount of AA^ater enter
Honey Lake. Some Avater enters the lake from Long Valley Creek and
several other small streams, but the volume is small and is not con-
sidered an important source for Honey Lake.

Honey Lake filled during 1937 and then gradually receded until it
became dry in the late 1940 's. AboA^e normal Avinter rains and snoAvfall
combined to fill the lake partially during the Avinter of 1950-51. Water
was abundant in the vicinity of the lake until late spring (May, 1951),
at Avhich time the Avater level of the lake receded rapidly until fall. Avhen
little or no water renmined. The abundant spring Avater supply enabled
dornumt emergent vegetation to attain some groAvth before the Avater
le\('l diopped in the fall. However, Avith the continuation of heavy
winter j)recij)itat ion during the Avinters of 1951-52 and 1952-53, the
emergent vegetation became abuntlant. in 1!)53 growth was considered
to have reached a maximum. Vegetative growth along the lakeshore,
excludiiiu' the large area at the mouth of the Susan River, Avas not
appreciably increased during 1952 and 1953, AA'heu Avater Avas abundant.

in general, drier conditions prevailed during the nesting season of
1951 than was the case in 1953. In 1953 the vegetation had become rank
and ideal for nesting in the area at the mouth of the Susan River.

Tjand use practices in Honey Tjake Valley haA'e remained relatiA'ely
stable dui-ing the last fcAv years. The chief agricultural activities are
concerned with past nriiig livestock and i-aising cereal crops.

XKSTIXG S'lTDIKS OF DL'CKS AND COOTS

297

>s

>»

TD

T3

a

3

«_

•♦—

(J)

CO

—

ro

lO

ID

(Ti

0^

II

11

CD

C

(6

1

<

—

OT

in

•^-

^_

O

o

Q.

CL

FIGURE 1. Map of study areas in Honey Lake Valley.

3—20860

298

CALIFORNIA FISH AND GAME

METHODS

Because of the large size of the study area, \>U)ls were usrd to sample
the nesting activities. The methods used in locating and marking the
nests differed slightly in the two studies. None of the differences was
of groat significance, and methods utilized in both studies obtained
satisfactory results.

FIGURE 2. Field recording of nest history data on a unisort analysis cord.
Pbofograph by C. V. Oglesby.

NESTIXO STT'DTKS OK DUCKS AND COOTS

299

Each i)lot was visited at least every 10 days. A rope was diji-r^rd
several times on plots to flush the nesting- birds wIkmi the ii;iliiic of
the vo<:'(>ta1inii made this practice possible. As each nest was toiiiid it
was assiyned a number and markeil by placing a willow marker several
feet from the nest to facilitate location on return visits. The marker
was aligned with the nest and a fixed object, a mountain peak easily
seen from any spot on all study plots. The distance between the marker
and nest varied from o feet in very dense cover to 15 feet in sjjarse
cover. The markers were p]a('<'d away from the nests to reduce the
chance of predators being attracted to the nests. The top of each willow
marker Avas cut on an angle, and the number assianed to the nest was
written on the cut surface. In the 19.").'^ study a white specimen tag was
tied to the top of each willow marker that was placed in dense cover
and to some markers that were placed in sparse cover. The flashing of
tliis white tag facilitated the finding of nests in all types and colors of
vegetation. It was found that little or no increase in the amount of
predation occurred in areas where the white tags were used.

At each visit to a nest all necessary information was recorded on a
nest card. A nest card was assigned to each nest, and all data gathered
during subse([uent visits to the nest were recorded on the same card
(Figure 2).

Table 1 gives the species composition of the nests found during both
studies.

TABLE 1
Species Composition of Nests Found

Species

Total nests

Percentage of total

1951

1953

1951

1953

Mallard

63
45
22
18
17

3

2
15
17

143
53

125
13

7

1

6

11

31.2
22.3
10.9
8.9
8.4
0.0
0.0
l..i
1.0
7.4
8.4

39 8

Pintail

Cinnamon teal —

Gadwall

Shoveller.

Baldpate

Ruddy duck

Redhead _.

14.8
34.8
3.6
2.0
0.3
1.7
3.0

Unclassi fied

Unclassified, destroyed when found

Unclassified, hatched when found

0.0
0.0
0.0

Total ducks

Coot - ,

202
20

359
143

100.0 100.0
9.0 ""SO

An attempt was made to find as many of th(^ n(^sts as ixissihic, l)ut
on the densely vegetated |)|ois all eonld not be found, in lli.')l it was
estimated that the percentage of nests found in relation to the actual
number on each plot ranged from ap|)roximate]y (iO percent in the
densely covered plots to lOO percent in some of the sjiarsely vegetated
plots. In 1953 an estimated 75 percent of the nests in the densely cov-
ered Idiots were found .iml 100 pei'cent of those in some of the sparsely
vegetated plots.

300

CALIFORNIA FISH AND GAME

The soieiitinr' iiaiiios of all birds, maiiirnals. and plants rofon-fd to in
this study arc ^'iviMi iu Appendix A.

SELECTION OF STUDY PLOTS

Because most of the nestiii«r in Honey Lake Valley in 1951 was con-
fined to the Fleniinjr Unit of the Honey Lake Waterfowl Manaprenient
Area, the l!l.')l study was made on this unit. In 1953 nesting activity
was nil 111' \\ i(l('s|)i-ead, and the study was conducted on both the Fleming^
T'nit and private land. Data on the number of breeding pairs of ducks
and coots are given in Table 2.

TABLE 2

Breeding Pairs of Ducks and Coots in Honey Lake Valley 1951-53 *

Species

19olt

1952

1953

Mallard

734
364
266
668
173
323

5
40

6

1,214

201

262

714

110

129

1

28

18

1,010

Pintail -

220

283

Gad wall

Slioveller

Rerlliead

Scaup

294

68

212

8

Ruddy duck - - -

10

Other - -

3

Total ducks -. .

2,579
126

2,677 2,108

Coot . -- . . -

398 620

* ropiilation rumbers taken fiMim California bieedini; ground surveys of 1951-1953.

t Part (if tlie 1H51 breedinn groiiiid survey was taken from the ground and is not completely compaiable to 1952-53,
when all figures were made from aerial counts.

Two strip plots were used in 1951. These plots included all cover
types ])re.-ent on the area. The combined area of the two strip plots
was approximately 300 acres, or 15 percent of the total area of the
Fleming L^iiit.

In 1953 eleven study plots were established to sample nesting on ap-
proximately 20,000 acres. They contained 328 acres, or aj^proximately
l.() pci-ceni of the total acreage in the study area.

Descripfion of the 1951 Study

Practically all the o|)eii water and marsh area in northeastern Honey
Lake \'alley existed on the Fleming I'nit of the Honey Lake Waterfowl
Management Ai-ea. Water was impounded in ai'tificial ponds diii-iug the
nesting season and early summer. As a result of the availability of this
water and mai-sh area, most of the waterfowl nesting in the valley Avas
believed to have been confined to the Waterfowl Management Area.

The information gathered during the 1951 study was compiled from
nests fouinl on the two strip i)lots located on the Fleming Unit which
were repi-esentativo of the cover ty])es found on the unit. Plot A con-
tained the following cover types: pasture grasses, volunteer barley,
cultivated Avheat, five-hooked bassia, rye grass, Baltic rush, hardstem
bulrush and other-s. Plot 1> contained five-hooked bassia. rye grass, Bal-
tic i-ush, hardstem bulrush, sagebrush, greasewood, and other cover
types.

NESTING STUDIES OF DUCKS AND COOTS 301

Description of the 1953 Study Plots

Eleven study plots were <'stal)lisli('(l in nine difTerenl. cover types yc[)-
resentin*; the general eovei- types nsed most extensively by nestinpr
waterfowl in the Hnncy L;ikr X'allcy in U»58. Seven plots of 40 acres
each, two plots of 20 acres each, and two plots 1 mile long by :{0 feet
M'ide were used. The two one-mile plots Avere along a ditchbank and
a levee and were both approximately four acres in area. In order to
sample 40 acres of cover growing on ditchbanks and levees it would
have taken 10 miles of ditchbank and levee, a factor not feasible in that
study. The two 20-acre plots were of Baltic rush cover type. The seven
40-acre plots were established to include samples of the following major
cover types: hardstem bulrush, river bulrush, sagebrush and grease-
wood, five-hooked bassia, rye grass, salt grass, cereal crops, and otlier
cover types.

NEST SITES AND COVER TYPES

In compiling data on both studies, two broad headings were used in
describing the locations of waterfowl nests. These headings or classifica-
tions were nest sites and cover types. The nest site classification de-
scribed the physical characteristics of the terrain where the nest was
located ; e.g., in a marsh, on an island, or on a dike. The most abundant
species of vegetation in the immediate vicinity of the nest was used to
designate the cover type found at each site. As an example of cover
type and nest site relationship, most of the duck nests found in 1953
were constructed in mai'sh ne.st sites and the dominnnt cover type
around the nests was Baltic rush.

A description of the different nest sites used during the studies
follows :

Dike or Difchbanl'. Elevated margins of any slough, creek, river,
irrigation ditch, or dam enroankment were classified as dike nest sites.

Marsh. Areas such as lakeshores, artificial ponds, and all semiwet
land were recorded as marsh-type sites.

Island. Any sizable piece of land completely surrounded by water
was considered to be an island nest site.

Agriculfural Laud. All land used for agricultural i)urposes was
listed as agricultural nest sites. During both studies most of the agri-
cultural land was either in irrigated pastui-e or in cereal crops.

Uncultivated Land. Dry upland-type areas not under cultivation
were classified as uncultivated land nest sites.

A differeiu-e was shown in the location of nest sites by ducks in 1951
and 1953. In 1951 nests were located primarily in dry upland areas.
Dikes and uncultivated fields were the most common nest sites used by
ducks that year. Results of the 1953 study showed an over-all change
of location to the marsh type site. The change was attributed to the
increased proportion of marsh nest sites available to the nesting water-
fowl. The marsh nest sites contained 14.7 percent of all duck nests found
in 1951 and 67.4 percent of all duck nests found in 1953. All coot nests
found in 1951 and 98.fi percent of the coot nests found during the 1953
study were located in marsh nest sites. Location of nest sites by .species
is shown in Table 3.

4— 2O8G0

302

CALIFORNIA FISH AM) (iA.ME

TABLE 3
Nest Sites (Percentage in Each Site)

Species

Dike

Island

Marsh

Unculti-
vated

Agricul-
tural

Total

Mallard

1951

39.7
18.9

20.7
22.0

59.0
1.0

44.4
7.7

29.4
0.0

0.0
0.0

0.0
0.0

0.0
0.0

50.0

1.6
0.0

4.4
0.0

9.2
0.0

0.0
0.0

0.0
0.0

0.0
0.0

0.0
0.0

0.0
0.0

0.0

23.8
(i2 . 9

4.4
37.7

9.2
89.0

0.0
23.1

17.7
0.0

0.0
0.0

0.0
100.0

100.0
100.0

0.0

19.0
15.4

22.2
34.0

18.1
8.0

27.8
09 . 2

23.5
85.7

0.0
100.0

0.0
0.0

0.0
0.0

0.0

15.9
2.8

42.3

5.7

4.5
0.8

27.8
0.0

29.4
14.3

0.0
0.0

0.0
0.0

0,0
0.0

50.0

100.0

1953..

100.0

Pintail

1951

100.0

19,53 -

100.0

('iiiii:uiii>u teal

1951 -

100.0

1953

100.0

Gad wall

1951

100.0

19.53 ..-

100.0

Shoveller

1951

100.0

1953.. -

100.0

Baldpate

1951 -. _

0.0

1953

100.0

Kuddy duck

1951

1953

0.0
100.0

Redhead

1951

100.0

1953

100.0

Unclassified

1951

100.0

Average for ducks

.37.7
11.7

0.0
0.7

2.9
0.0

0.0
0.0

14.7
07 . 4

100.0
98.0

20 . 0
18.4

0,0
0,7

24.1
2.5

0,0
0 , 0

100.0

1953

100.0

Coot

1951 -

1953 -.

100.0
100.0

'I'lic change in nest sites between 1951 and lft")-'5 was accompanied by
a clianj^c in cover types. The cover utilized most often by nesting ducks
in li).")! WHS ry(' grass, five-hookod hassia and salt grass. These three
})lant species provided cover for (iO.O percent of all duck nests found
that year. In 1!)53, 59.7 percent of all duck nests found were in Baltic
rush. Emergent plants were the preferred cover tyj)es utilized by coots
during both studies. During 1951, 95 ])erceiit of the coot nests found
were in hardstem bulrush. Baltic rush, river bulrush, and hardstem bul-
rush proxidcd cover for !)8.6 percent of the coot nests found in 1953.
The utilization of the different cover types during both studies is pre-
sented in Table i.

NESTING STUDIES OP DUCKS AND COOTS

303

-S
o

o

o

o o

O 00

O 00

o q

q q

o o

o o

o o

q ^

o o

d oi

d d

d d

d d

d d

d d

d -^

88

o r~

(M so

o —

^» o)

oi r^

•?! d

CI '?!

Cl CO

r~ CO

o o

Ln o

■' -a

O 05
— lO

d -^

•r d

00 -r

CO

— ■ CI

00 d

-r CO

88

<

""

.It

o o

o o

o o

o o

o o

c o

q q

o o

o o

q q

d d

d d

d d

d d

d d

d d

d d

d d

— ■ ;:^

d d

lO

'0

o

t3

•a

'T3

t^ M

cc t^

o -^

q q

o o

o o

o o

o o

o o

o o

d 00

d c-i

O 05

d d

d o

d o

d d

d d

d d

d d

(D

?:; — .

M r~

o o

«

^«

■^^

o o

o -^

o o

q q

o o

o o

o o

o o

o o

o o

■§§

d d

O M

d d

d d

d d

d d

d d

d d

d d

d d

tf^

W

o

4)

c3

a

o o

o o

c o

o o

q q

c o

q q

o q

- =>

o o

2

d d

d d

d d

d d

d d

d d

d d

d d

d d

d d

C3

o

m

u

<u

rj< ^

o o

o o

O CO

O! o

"O o

-f '.O

00 o

q q

q q

>
o

SJ^i

d d

d d

d -^

d d

d d

d 00

— d

d d

d d

c-x to

f-*

C-l

CI Cl

o o

^

I—* f^

OJ

=3

&

O !M

o o

o o

O -H

(N O

00 t»

CD O

c) o

CO o

o o

-0

in d

d d

d d

d m

c-x d

00 (^

lO O

'-' o

d d

d d

t3

o

(N

<N

CO

r^

o o

a

d

o

S •-:

o o

o 00

o o

lO 00

>0 O

o o

O O

o o

C 00

o o

a ?

c 5

d d

d d

d d

•* d

T O

1- o

00 —

d d

d d

d d

C

05

o

c o

O

'3

o o

05 'O

o r:

-r M

00 M

CO b'

-r o

— o

O -l>

o o

e

d ^

00 t-i

d -^'

-r cc

t^ —

d 10

CO o

>c d

d 05

d d

E

"5

CO

o o

"H

03

00 t^

!M Tt«

O -f

00 T)<

>o t^

« t~

CO 00

00 t~

00 CI

o o

03

t^ lO

C-I —'

d -

t>i 00

1^ ts.

CO t^

CO CI

-r o

-r -r

d d

§

w

M e<5

IM

M

o o

0}

a

>>

".•«

u

o

00 ■ ■

>
o

■a 1 1

C3 ' '

O

■£! ! 1

CO 1 1

3 ' '

M I 1
0) 1 1

t-i ( 1

i ; :

i : ;

3 ' '

"m • '

3 ! !

S ' i

2 1 1

.2 Sfo

•^-'d

t, -. CO

.ads

03 1 <
^ ^ CO

C 'H CO

U 1 1

— — CO

— CO

— CO

>- lO lO

X! lO >ra

Mm iO

Mira ira

es m >o

t- in lO

— -0 o

■** 05 Ol

» o: 05

V oi a>

« 3) OS

C 05 2

_„ OJ o>

« 05 Ol

2 Ol 05

J 0-- 05

"a -^ —

a '^ —

r*i

3

"a

O

304

CAIJI'OKNIA IISII ANDCAMK

In order to j^et the over-all trend ol' tlie preferred nestinjj: sites
and cover types, iiironiiaf ion from these two classifications was com-
bined in Table 5.

TABLE 5

Nest Site — Cover Type Relationstiips

Nest site *

Cover type

Species

1951

1953

1951

1953

Mallard...

General

General

General
Dike

Dike

Marsh

General

General

Marsh

Unculti-
vated

Unculti-
vated

Unculti-
vated

Marsh

Marsh

Bassia

Rye grass

Hardstem bulrush

Rye grass

Bassia

Rye grass

Rye grass

Bassia

Salt grass

Baltic rush

Baltic rush
Hardstem bulrush

Baltic rush

Pintail

Cinnamon teal

GadvvalL-

Hardstem bulrush

Baltic rush

Baltic rush
Baltic rush

Shoveller ..

Baltic rush

Baldpate . .

Grasses
Sagebrush-grease-

Ruddy duck.

wood
Hardstem bulrush

Redhead

Hardstem bulrush

Average duck.

Dike

Marsh

Rye grass
Hardstem bulrush
Bassia

Baltic rush

Hardstem bulrush

Coot

Marsh

Marsh

Hardstem bulrush —

Baltic rush

River bulrush
Hardstem bulrush

* Where the term "Knioial" is iisprl. there was no definite preference for a nest site, and nesting occurred in the
nest sites with ahuut the same frequency.

NESTING PERIODS

The springs of 1951 and 1953 were considered favorable for nest-
ing waterfowl in Honey Lake Valley. Although a change in loca-
tion of nest sites and cover types was found in the two studies, the
nesting periods and hatching dates were quite similar. The first nest
found in 1951 was on April 19 ; in 1953 the first nest was found on
April 22. The last nest history in 1951 was completed on July 25
and in 1953 on July 17. Nesting continued in the valley after these
dates during both years, but it is believed that the number of nests
hatched after July 25 was nominal and had little or no effect on the
peak of hatch.

Nesting periods similar to those shown above were recorded at
the Tule Lake and Lower Klamath National Wildlife Refuges in
Siskiyou County (Miller and Collins, 1954).

Information concerning peak of hatch for both ducks and coots is
illustrated (Figure 3).

NESTING RTUDIKS OF DUCKS AND COOTS

:!().")

UJ
CD
<

UJ
O

cr

UJ

ql

lO

o

ID

ro

ro

a>

ro

4>

>N

>»

>s

c

■^—

O

3

c

ZJ

3

^

-3

3

-3

-3

2
1

1

-3
1

1

1

<o

..

CD

—

(0

o

c

3

4»

C

>»
3

3

S

-3

-3

->

-3

FIGURE 3. Hatching periods and peak of hatch for duck and coot nests

in Honey Lake Valley.

306

CALIFORNIA FISH AND GAME

FATE OF NESTS

The categories used in ilassifying fates of nests -were the same as
those used in several other nesting studies in California. The fate
of nest classification used was as follows: (1) hatched nests, (2)
deserted nests, (8) flooded nests, (4) destroyed nests and (5) fate
iiiikiiow 11 nests. A (lefinitioii of these categories has been published
by Miller and Collins (liioij). Table (i shows the fate of all nests
found during the studies.

TABLE 6

Fate of Nests *

Species

Mallard

1951

1953

PintaU

1951

1953

Cinnamon teal

1951

1953

Gadwall

1951

1953.. .

Shoveller

1951

19.53

Ruddy duck

1951

1953

Redhead

1951

1953

Total ducks

1951

1953..

Coot

1951

1953

Number
of nests

63
143

45
53

22
125

18
13

17

7

3

11

168
359

20
143

Percent-
age
nests
hatched

GO. 4
47.6

33.3
45.3

54.0
56.0

66.7
38.5

58.8
28.6

0.0
66.7

33.3
54.5

52.5
50.1

95.0
97.2

Percent- Percent-
age age
nests nests

destroyed deserted

28.5
36.4

57.8
45.3

13.6
28.8

22.1
46.1

41.2
42.8

o.n

0.(1

66 . 7
U. 1

35.0
34.3

5.0
2.1

7.9
14.0

4.5
9.4

31.8
13.6

5.0
15.4

0.0

28.6

0.0
33.3

0.0
36.4

9.5
14.2

0.0
0.7

Percent-
age
nests
flooded

3.2
2.0

4.4
0.0

0.0
1.6

5.6
0.0

0.0

0.0

0.0

0.0
0.0

3.0
1.4

0.0
0.0

Total

100.0
100,0

100.0
100.0

100.0
100.0

100.0
100.0

100.0
100.0

0.0
100.0

100.0
100.0

100.0
100.

100.0
100.0

34 nests not shown from the 1951 study were: 2 unclassified, 15 destroyed when found, 17 hatched when found.
One nest not sliown from llie 1953 study wa.s a baldpate nest that was destroyed.

Successful Nests

The average nesting success for the duck nests found on study
])lo1s (lurin<>- both years was similar. In 1!)51, 52.5 percent of the
(luck nests hatched, while in 1953 success rate dropped slightly, with a
hatch of 50.1 percent.

The over-all hatching success of dabbling ducks was lower in 1953
than in lH.")]. The nintfiil \\;is the oiih- dabblinii' duck that showed

NESTING STUDIKS OK Dl'CKS AND COOTS

MOT

any apprcciabb' ^■aiii in ncstin;^- .siiccoss in I!)-"):!. 'I'lir i;itr of .success
foiuul in cinnanion Ir.il nesting- was fclati\cl\ constaiit diiriii}; both
studies. In 3951 gathvall. mallard, and sliovclcr were iIk; most suc-
cessful nesters of tlie dabblini:' ducks, with success rates of ()(i.7, ()').4,
and 58.8 percent, respect i v. 'ly. In ]!).').■{ the three species of dabhl inn-
ducks that Avere the most successful nesters were the cinnamon teal,
mallard, and pintail. The success rates for these s])ecies were .')().(),
47.6, and 45.8 percent, respectively. The success rates of the uests
of diviufT dueks found (luriii<r the studies were 33.8 percent for red-
heads in 1051 and 54.5 ])ercent in 1!I5;5. The hatehiuf? success of ruddy
duck nests was (ili.T percent in 1958. No ruddy duck nests were louiid
in 1951.

All recent duck and coot nesting' surveys in iiortlKjastern Calif(jrniu
have indicated that the coot is the most successful nester with respect
to hatching success and hatchability of epgs. The success rate for coots
at Honey Lake Avas 95.0 percent in 1951 and 97.2 percent in 1953.
The success rate for coots at the Tule Lake and Lower Klamath
National Wildlife Refuges was 94.6 percent in 1952 (Miller and Col-
lins, 1954).

Unsuccessful Nests

DestrHcfiou. Predation on nests of both ducks and coots was the
greatest single cause of nesting failures during both studies. The
amount of destruction attributed to predation was relatively constant
during both studies. Of all the duck nests that were found, 35.0
percent wei-e destroyed in 1951. whili; 84.8 i)ercent were destroyed
in 1958. Tlie amount of predation on coot nests was light, with 5.0
percent of the nests destroved in 1951 and 2.1 percent destroyed
in 1953.

The cause of nest destruction was difficult to determine in many
cases. The lack of sufticient evidence to establish definitely the cause

TABLE 7
Percentage of Destroyed Duck Nests Found in Each Nest Site

Dike

Island

.Marsh

Unculti-
vated
land

•Agri-
cultural
land

Total

Destroyed by mammal

1951 .

64.0
25.7

12.0
20.0

24.0
54.3

0.0
(t.O

100.0
100.0

0.0
0.0

0.0
0.0

0.0
0.0

0.0
(1.0

0.0
0.0

33.3
40.0

.^)0.0
27.5

10.7
32.5

0.0
0.0

100.0
100.0

85.7
45.3

0.0
11.9

14.3
42.8

0.0
0.0

100.0
100.(1

38.9
10.7

5.5
16.7

27.8
6(> . 0

27 . 8
0.0

100.0
100.0

55.4

1953 . ---

36 . 6

Destroyed by bird

1951 . -

12.5

1953 --

19.5

Destroyed by unknown causes
1951 . . -

23.2

19,53

43.9

Destroyed by uiinatviral causes
1951 -

8.9

19.53

0.0

Total

1951

100.0

1953

100.0

One coot nest was destroyed In 1951 and three coot nests were destroyed in 1953 in marsh nest sites and are not
shown in the table.

308

LALUOKMA FISH AM) GAiNIE

of predation was responsible for the larjre iiumbei' of nests attributed
to destrnetioii by unknown causes (Table 7). If there was any doubt
as to the identity of the i)redator a nest was listed as destroyed by
unknown causes. Mammalian predators known to occur in the area
were the striped skunk, coyote, house cat, badger, bobcat, and weasel;
the avian species Avliicli prey on nests were the California and ring-
billed gulls, the black-billed magpie, the crow, and the raven. An
instance of nest destruction by unnatural causes occurred during
1951, when five lu^sts were destroyed by land-leveling o])erations.

Preseason trapping of predators bj^ a State trapper on the Honey
Lake Waterfowl ^Management Area resulted in the capture of 23
striped skiuiks. 4 coyotes, 9 bouse cats, and 2 bobcats in 1951 and
'.U striped skunks, 1 coyote, and 5 house cats in 1953.

Ihscrtiun. The amount of desertion found in duck nests was 9.5
percent in 1951 and 14.2 percent in 1953. This higher rate of deser-
tion was the greatest difference found in comparing tbe results of
the fate of nests in the two studies.

Competition for preferred nest sites among the ducks probably
accounted for some desertion. However, data concerning desertion

FIGURE 4. Mallard duck nest parasitized by a pheasant. The six pheasant eggs show darker
and smaller than the five duck eggs. Phofograph by E. G. Hunt.

NESTING STUDIES OP DUCKS AND COOTS

:!()

due to competition foi- prefei-i-ed nest sites ^vere ineonsistent, and no
definite statement can be made on lliis subjeet.

Pai'asitism occui'i'rd in '2.7 pci-cciit of llic total duck nests found
in 1951 and in (i.'i percent of the lota! duck nests found in 1053. An,v
nest eontaininji' e<i<'s laid by two species of ducks or a duck and a
pheasant was considered parasitized (Figure 4). There were no
instances of a duck nest being parasitized by another species of duck
during the 1951 study. In 1953, 13 (42 percent) of the duck nests
parasitized contained eggs laid by another species of duck. Para-
sitism of duck nests by i)lieasants occurred in all five of the para-
sitized nests found in 1951 and in 18 (58 percent) of the duck nests
parasitized in lf)53. Some dcsiM-tion resulting dii-cctly from jiarasitism
in duck nests probabl.y occurred, but data gathered during both
studies showed that parasitism w^as not an important cause of deser-
tion. No instance of parasitism was found in coot nests during either
study.

Only one coot nest was recorded as deserted during the 1953 study,
and none was deserted during the 1951 study. It was believed that
overcrow'ding in preferred nest sites and parasitism that may exist in
duck ni^sts were not factors that affected coot nesting. The pugnacity

TABLE 8
Clutch Size and Average Hatch Per Clutch

Species

Successful
nests

Total
eggs

Average
clutch

Average hatch
per clutch

Mallard
19.J1
1953-

Pintail
1951 ,
19.53-

Cinnanion teal

1951 .
1953.

Gatlwall

1951

19.53-.-

Shoveller

1951

19.53

Ruddy duck

1951

19.53--

Rpdheai
1951.
1953-

All ducks
1951-..
1953...

Coot
1951.
1953.

38
64

15

12
04

12
5

10

88
ir.8

19
123

339

529

111
170

125
583

120
50

109
20

0
15

14
42

824
1,415

154
913

8.9
8.3

7.4
7.0

10.4
9.1

10.5

1 0 . 0

10.9
10.0

0.0
5.0

1 l.O
8.4

9.4
8.4

8.1
7.4

8.3
0.9

6.8
5.9

9.7
7.8

9.0
7.4

10.1
10.0

0.0
5.0

4.0
5.4

7.9
7.4

310

rAT.TFORXI.\ FISTT .VXD 0.\:\rE

Mitli wliicli the rnni i\c\\']]{\s ;i iii'stiiiii 1 1'l'i-i 1 oi'v iiii'jiit 1)(> a reason
for sucli ;i low ilcscrtioii rate.

FUxidiini. The f1o()(liii<r of nests was of minor si^nificanco in the
success of (luck and coot iiestin<i'. Five duck nests were found flooded
(luiiiii; each of the sliidics. Xo instance ol' a coot nest beinj:' flooded
was recorded. Stable nr I'ecedin^- water levels dui'iiig the nesting
season acconnti'd \{\v \\\v Inw incidence of floochnl nests.

FATE OF EGGS AND CLUTCH SIZE OF SUCCESSFUL NESTS

The inf(!i-iiia1 ii)ii cdllected from successful nests was used to deter-
mine the average clutch and fate of eggs. The average clutcli size of
hotii ducks and coots was found to be sliglitlv lower in 1953 than
in lf»51 (Tabh' 8).

All available data concerning the fate of eggs were recorded in the
following categories: (1) number of eggs hatched, (2) number of eggs
desti'oyed, (3) number of eggs infertile, (4) number of eggs containing
dead embryos, (5) number of eggs missing, and (6) number of dead
in nest. The fate of eggs in successful nests is shown in Table 9. The
number of ilead young in nests were included in the percentage of eggs
hatched.

TABLE 9

Fate of Eggs Expressed in Percentages

Species

Hatched

Dead
embryo

Infertile

Destroyed

Missing

Total

Mallard

1951 - - ...

93.5
83.5

91.9
84.2

92.8
85.5

01.2
71.0

92.7
100.0

0.0
100.0

28.C)
01.9

91 .7
83.9

97.5
99.3

5.0
12.8

7.2
10.7

0.8
11.2

3.2
Ki.O

4.0
0.0

0.0
0.0

71.4
20.2

5.4
12.0

0.0
0.3

0.0
0.2

n.9
1.1

0.8
0.0

1 .11

0.0

0.9
0.0

0.(1
0.0

0.0
11.9

0.0
0.0

O.fi
0.0

0.0
1.3

0.0
0.6

0.0
1.0

0.0
4.0

0.0
0.0

0.0
0.0

0.0
0.0

0.0
1.0

0.0
0.4

1.5

2.2

0.0
3.4

5.6
2.3

4.0
0.0

1.8
0.0

0.0
0.0

0.0
0.0

2.3
2.5

1.9
0.0

100.0

1953 . .

100.0

Pintail

1951 -

100.0

19.53

100.0

Cinnamo teal
1951

1953

100.0
100.0

Gadwall

1951

19.53

Shoveller

1951

19.53

Kmiiiy duck

1951

100.0
100.0

100.0
100.0

0.0

19.'-)3.

Redliead

1951

1953. .

100.0

100.0
100.0

Average of all ducks

1951

1953

100.0
100.0

Coot

1951

100.0

1953

100.0

NESTING STUDIKS OK DfCKS AM) Coo'lS .'il 1

Tiu' siicccssl'iil (luck Jicsfs produced Sli4 ixjns in llic lit.")] study ;iiid
1,41") ('••••i-s in the 1958 study, of winch T.'),') luitclicd in I!).')! and 1,187
hatched in 1!>53. The hatchiuL: success of the duek e<>:<xs in If).")! was
91.7 percent ; in lO'j.'J it was <s;{.!) percent. The hatehiiij;' success of coot
e<iji-s was !)7..') pei-cent in 1951 and 99. :{ |)ereent in 195:5. Oidy four coot
e<>'<i's out of 154 in 1!I51 and seven out of !)!:! in 1!»5:! did not hatch.

The total number of eo-gs atl i-il)ule<i to pai-asit ism in successfully
liatched duck nests was 2(1 in 1!t5;{ and two in 1!)51. Of tiie 2() ey-o's
fouutl ill l!)5:j. Ki wei-e duck e^j^s and 10 wert' piieasaut cirjis ; both
of the eggii found in 1951 were plieasant eggs. The small number of
eggs resulting fi-om parasitism made little difference in the total
nnmber of eggs in the snccessfully hatclu'd nests and was not com|)ute(l
in the fate of eggs or average clutch.

Infertile eggs and eggs containing dead end)i-yos that wei-c found
in successfully luitched nests were classified by the method described
by Kossack (1950). An (>gg was considered infertile if the yolk was
su.spended in the albumen and no indication of development was pres-
ent. Any egg that contained a dead embryo in any stage of develop-
ment or coutaiued yellow custarddike material was classified as a dead
embryo.

In all, 45 eggs containing dead embryos and ti\-e infertile eggs were
found in successfully hatched duck nests in 1951, and 172 eggs con-
taining dead embryos and eight infertile eggs were found in 1953. In
all the successfully hatched coot nests oidy thi-ee eggs containing dead
embryos and no infertile eggs were found in '[i)'r.\, and no dead embryos
and one infertile egg were found in 1951.

A total of 50 eggs was either destroyed or missing from the success-
fully hatched duck nests in 1953, while in li>51 there were 19 eggs
missing from successful duck nests, but no eggs -were destroyed. Suc-
cessful coot nests contained four destroyed eggs aud had no eggs miss-
ing in 1953; there were no destro\-ed eggs and foui- e<:us were missiu"-
in 1951.

BROOD DATA

The number of duck brood counts taken during the two studies was
132 in 1951 and 151 in 1953. Most of the duck broods counted were
either one or two Aveeks old. Only 30.1 ])erceid of the dui-k broods
counted diii-ing 1951 and 44.3 pcfceiit of the duck l)i-oo(ls counted dur-
ing 1953 were ovei* two weeks old. Losses in broods din'ing the first week
of life averaged 0.1 bii-d in 1!)51 and 0.7 bii-d in 1953.

Xo coot broods wei'e counted durinji- 1951. and only 24 coot broods
were counted durini;' 1!I5;). Since a paii' of coots will often split the
brood between them, bi'ood counts may not accurately reflect the actual
brood size. Therefore, obsei'vers tallied only those broods that could be
considered eom|)lete. An average loss of 3.0 coots ]^e\' brood during
the first wi'ck of life oeeuiTcd in tlie t'l'w coot broods taken. This loss
was undoubtedly tlue to the .ueneral helplessness of youuL; coots dui'ing
their first few days of life (Gulliou, 1954).

DISCUSSION

The amount of water in lIone\- Lake has a definite bearins: on the
number of waterfowl utilizing the valle\ throu'jliouf the vear. The lake

312 CAT.TrnrtxiA ftsh and game

ill wet years provides ;iii ;i(liM|iiate resting place for tlie spring migrants
and attracts breeding pairs that i-emain to nest in the area. Fall mi-
gi-atiiig Avatert'owi feed and rest in the vicinity of tlie lake and, together
with tlie waterfowl pnidneed in the valley, provide Inmting during the
waterfowl season.

In rj.")l most of the duck nesting oeeui-red in dry, upland-type hab-
itat that was adjacent to artificial ponds. These nesting areas provided
a combination of good nesting cover and sufficient water for rearing
liroods. Tile iti-eFei'i-ed nesting (•()\'ei' under If).")! conditions was rye
grass, five-hooked liassia, and salt gi-ass. These plant species grew in
clumps and ])i-ovi(|ed eoxcr that was relatively low and dense. In 1!)");]
there was iioi only an ahniidant growth of upland plant species, in-
clmling rye grass, five hooked bassia, and salt grass, but also an abund-
ant growth of emergent plant s])ecies, such as Baltic rush. The Baltic
rush offered llic same coiieealnient factors as the rye grass, five-hooked
bassia, and salt grass and was usually growing in or near water. A\^-
proximately the same percentage of duck nests was found in the Baltic
rush in ]95;5 as was found in rye grass, five-hooked bassia, and salt
grass in 1951. Apparently the conditions that prevailed at the nest
location, such as concealment and proximity to water, were more im-
portant to the nesting ducks than the selection of a certain plant species
in which to build a nest. The coot nesting during both studies was con-
fined to areas that grew emergent plant species.

From observations made of coots it appears that both parents par-
tici])ate in incubation of the eggs. A further observation is that the
coot often commences incubation at some interval after the first egg
is laid and before the final egg of the clutch is laid. This would enable
the coot to hatch several of the young and allow one parent to take
the young from the nest and the other parent to continue incubating
until all eggs in the clutch were hatched. These observations followed
the coot nesting behavior described by Gullion (1954) and others. As
an example of the frequency of this behavior, 83 of the 139 coot nests
hatched during the 1953 study were hatched in this manner. The early
start in incubation would also give the coot eggs more protection than
that received by duck eggs.

The most frequent cause of nest failure was predation. Approxi-
mately one-third of all nests found during both studies were destroyed
by predators. The species of ducks that nested in the dry upland
locations sustained the majority of the nest destruction in each study.
Many of the ducks nesting in the upland areas preferred ditch banks
and dikes for a nest location. Mammalian predators, principally the
striped skunk, seemed to hunt these areas extensively in search of food.
Nest destruction in the marsh area was very limited, apparently be-
cause of protection afforded by standing water. Undoubtedly the ab-
sence of such aquatic predators as the mink was also a factor in the
low incidence of predation in the marsh area. The rate of nest de-
struction by avian predators was low in marshy areas and moderate
in the upland areas during both studies.

Since coots habitually built their nests over water they were pro-
tected from most mammalian predators, and thus were the most success-
ful nesters studied. Another factor which may have contributed to the

XKSTIXCi STl'I)li:S OK DUCKS AND COOTS -11;!

high rate of nesting success of coots was the participjition of both
parents in guarding the nest.

An insufficient nunilxT of Iji-oods was counted to (h'tei-iniiic accurately
brood regression during citlier study. Tlic utilization of dense escape
cover by the duck and coot broods made brood counting difficult. The
one- and two-week-old broods were the only age classes that were
counted frequently. Tnfornuition taken from the brood cards regarding
week-old duck broods indicated that there was a slight reduction in
brood size during the first week of life. The coot broods that were
counted showed a loss of approxinuitcly 40 percent of the luuiibcT- of
hatched young during the first week of life.

SUMMARY

1. Studies on nesting ducks and coots were conducted during the
spring of 1951 and 1953 in Honey Lake Valley, Lassen Comity,
California.

2. Two sample strips with a total area of 300 acres were studied
during 1951 ; 11 study plots with a total area of 328 acres were studied
during 1953.

3. Xest histories were completed on 202 duck nests and 20 coot nests
during 1951. In 1953 nest histories were completed on 359 duck nests
and 143 coot nests.

4. The peak of hatch for coot nests was between June 1 and June 15
during 1951 ; for duck nests, between June 1 and June 30. The peak of
hatch for both ducks and coots during 1953 was between June 16 and
June 30.

5. The nesting success for all nests found in 1951 was 52.5 percent
for ducks and 95.0 percent for coots; in 1953 the nesting success for all
nests found was 50.1 percent for ducks and 97.2 percent for coots.

6. Predation was the most important cause of unsuccessful nesting
of ducks and coots during both studies.

7. The hatching success of eggs in the successful nests in l!i51 was
91.7 percent for ducks and 97.5 percent for coots ; in 1953 the hatching
success was 83.9 percent for ducks and 99.3 percent for coots.

8. In 1951, 132 duck broods were counted, while in 1953, 151 duck
broods were tallied. The brood count data showed that on the average
less than one duck per brood was lost during the first week of life. The
24 coot broods counted in 1953 revealed an average reduction of 3 coots
per brood during the first week of life.

REFERENCES
Gull ion, Clordon W.

1954. The rcprodiu-tivc cycle of American coots in California. Auk, vol. 71, no. 4,
p. 3G6-412.

Kossack, Charles W.

1950. Bieediuj,' habits of Canada geese under refuge conditions. Anier. Midi. Nat.,
vol. 43, no. 3, p. 627-049.

Miller, A. W., and B. D. Collins

1953. A nesting study of Canada geese on Tulo Lake and Lower Klamath
National Wildlife Refuges. Siskiyou County, California. Calif. Fish and
Game, vol. 39, no. 3, p. 385-396.

1954. A nesting study of ducks and coots on Tule Lake and Lower Klamath
National Wildlife Refuges. Calif. Fish and Game, vol. 40, no. 1, p. 17-37.

314

CALIFORNIA FISH AND GAME

Na.\l..r. AllMTt E.

1!>5;J. I'rodiiciioii (if the ("aniida >;"'os<' on Honey Lake Refuge, Lassen County,
("alifornia. Calif. Fish and (lame, vol. 31), no. 1, p. S3-94.

Naylor, A. K., and E. (i. Hunt

l!)ij4. A nesting stud.\ and |)oipulation survey of Canada geese on the Husan
Kix'er, Lassen County, ("alifornia. Calif. Fish and Game, vol. 40, no. 1,

APPENDIX
Scientific Names of Animals and Plants Listed in the Text

JUnls

-Mallard — .1 iiny i)hil i/rlujiiclios
Pintail — Anas (lentil tzitzifioa
Cinnamon Teal — Anas ci/diioiitciii
( ;;i(l\v:ill — .1 nnx strcpern
S 1 1 o \ ( ■ 1 1 e r — Spat Ilia chjpeata
I'.aidpate — M ari'ca uniriicn iia
Kuddy Duck — Oxyura janinirenxis

r u h ida
K I 'ill lead — .1 th i/d luiiciiriitni
( 'ooi — h'liUra (uiicrivana
California (inll — Larux valijornicus
King-hilled (lull — Lams delawarensis
J>lack-hilled Magpie — Pica pica

hudsonia
Western Crow — Corvus liraihi/-

rliynchos hesperis
Raven — ('arr us coraj-

M annuals

(Ireat J'asin Striped Skunk —

Mephitis iiirphitis major
-Mountain Coyote — Caiiis latians
lestes

I'allid Bohcat — Lynx rufiis pallcscens
California Badger — Ta.iidea taxiis

neglecta
"Weasel — Musiela sp.
Houseeat — Felis domesticus

Plants

Grasses — Gramineae

Cultivated Barley — Jlordeum vulgare

Cultivated Wheat — Triticuni

aesiiruni
Black Greasewood — Sarcobatus

vermiculatus
Sagebrush — Artemisia trident a ta
Five-hooked Bassia — Bassia

hyssopifolia
Rye Grass — Elymus sp.
Baltic Rush — Juncus haltjcus
Ilardstem Bulrush — Scirpus acutus
River Bulrush — Scirpus fluriatilis
Alkali Bulrush — Scirpus paliidosus
Salt grass — Distichlis spicata
Alfalfa — Medicayo sativa

NESTING AND REPRODUCTION OF THE BAND-TAILED

PIGEON IN CALIFORNIA'

WALLACE G. MACGREGOR and WALTON M. SMITH

Game Management Branch

California Department of Fish and Gome

INTRODUCTION

The State of California is the iiriiicipal wiiiteriiij'' area of the band-
taihul |)i;^-eon {CoJiniihii fasciotd iiioitilis), which is one of the important
nii<iratory {jame birds of tlie J'acifie Coast. Hunting- pressure is exerted
on this speeies not only in California, but also in Oregon and Wash-
ington. Past fluctuations in population due to one eause or another
have been reflected in crop (h'pi-edation complaints and hunting
closures.

In order to determine the status of this species and secure adequate
data on which to base recommendations for management, a Pittman-
Rol)ertson I'esearcli project was set up in 1950. This paper reports on
the results of a part of this study.

PRODUCTIVITY

The band-tailed pigeon has been credited with having the lowest
breeding potential of any game bird in the United States. Ever since
Grinnell (VJVl) wrote, "it seems impossible that, at least as far as
our knowledge of the species on the Pacific Coast is concerned, more
than one brood can be raised each year by a single pair of birds * * *,"
authorities have assumed that pigeons rear only one brood per year,
(ilover (li».');}) states that '"there is no reliable evidence that the band-
tail nests more than once each year in California, except when its
first nesting is disturbed. Within the coiitriK^s of tliis study, no data
was obtained to the contrary.''

Tlie authors first began to siis|)ect that the reproductive potential
was higher than one squab i)er year while conducting checks of pigeon
hunters in ^*^'y2. Age ratios were obtained at that time which would
be possible oidy if there was a higluM- i-eproductive rate, or a dif-
ferential migi-ation of young and old hii'ds. These age ratios were
obtained by examining the hni-sa of Fabrieius in bii-ds killed by
hunters. Although imature birds can l)i' distinguished from adult
birds by idumage this characteristic is not completely valid during the
fall hunting ]ieriod. as some of the earlier hatched l)irds have com-
pleted the moult and ha\c full adult pluma-v. The presence of tlu^
bursa of Fabrieius as deserihed l.y Kiddle il!iL'Si is the ])est method
of telling immature from adult birds.

1 Submitted for puMication .Tune, T.*."..'. Thi.s .'<tudy was made possiMo with funds of
Federal Aid in Wildlife llestoration, Califurnia Project W-42U, "Tlie Life History
and Management of the Kand-tailed Pigeon in California" and I'roject W-4TR.
"Upland Game Investigations."

(315)

316

CALIFORNIA FISH AND GAME

Table 1 izives a<;e ratios of birds in Oregon and California from
1950 through l^'A. All of these ratios were based on examination of
the bursa in ])ii-(ls killed durinii' the liuiitinf? season.

TABLE 1
Band-tailed Pigeon Age Ratios Obtained Froni Hunter Killed Birds

Locality

Oregon*

Oregon*

Sliasta County

Monterey County

Kern County

Monterey County
Monterey County

Date

September, 1950
September, 19.51
September, 1952
December, 1952.
December, 1952.
December, 1953.
December, 1954.

Adult:

Immature

1:0.25
1:0.41
1:0.26
1:0.79
1:2.00
1:0.97
1:0.81

Number in
sample

4.39
350
77
171
161
146
352

•Morse (1951).

It Avill 1)(^ noted that the age ratios obtained in September, both
from Oreg-on and California, are below 0.5 immature bird per adnlt.
In contrast, the ratios obtained durinp' December range from 0.7!)
immature per adnlt up to 2.00 immature per adult. As a ratio of 0.5
would be the maximum expected ratio if the production were one squab
per season, there must have been higher production or differential
migration of vonng and old birds.

NESTING

In 1952 a nesting study was initiated in the Carnlel area of Monterey
County. This area was chosen because of its accessibility and high
density of nesting birds. During the period from July, 1952, to ^Fay,
1955, a total of 26 pigeon nests and 33 nestings was observed. Although
this is a fairly small number of nests, it equals the number of all the
other pigeon nests found in California and i-ecorded in the literature
to date.

Grinnell, Bryant. ;ind Rtorer (1918) gave data on 17 nests recorded
in the literature or known to iIkmii from 1877 to 1918. More recently
Glover (1953) re])orted on five nests from northwestern California.

Table 2 summarizes the nesting data obtained by this study. The
locations of the nests are shown in Figures 1 and 2.

Nest Location

All the nests were located in trees. The nests were found at heights
from 12 to 95 feet, Avith an average height of 36 feet. Sixty-two per-
cent (16) of the nests were found in ^Monterey pine (Pinus rarliata) ;
19 percent (5) in coast live oak {Q"crc\'S agrifolia) ; 15 percent (4)
in acacia (Acacia sp.) ; and 4 percent (1) in hawthorn (Crataegus
sp.). The pine and oak are native species and the acacia and hawthorn
are cultivated exotics. Most nests were const meted of a flimsy aggre-
gation of twigs placed in the fork of a limb. In the case of a nest lo-
cated next to the Carmel Police Station, Avith each new nesting addi-
tional material was added on top of the i)i-eceding nest, until it became
quite a substantial structure.

BAND-TAILED PIGEON NESTING

:>>]

KEY

D Nests Active in 1953
O Nests Active in 1954
A Nests Active in 1955
• Nest Active in 1952-1955

.C7"

r

03

4^G
5

8G
O

10

o,

Ocean

-O

O

FIGURE 1. Map of band-tailed pigeon nests in Carmel, 1953-1955.
Drawn by C/iffa Corson.

Number of Eggs

Due U) the liei^ilil of sunn,' of the nc>ts ami othor factors it was
possible to observe the contents of only 26 nests, 'rwi-nty-five (96 per-
cent) of tlie nests (•(nitaiin'il (nic ("j:v: of s(|uab and one nest contained

two eggs.

:nH

CAI.IIOKXIA I'lSlI AXDHAATE

FIGURE 2. Map of band-tailed pigeon nests in Carmel Highlands, 1952-1954.

Drown by Cliffa Corson.

Nesting Period

Tlio cai'licst (late of i>^fx layin<i' noted in tliis study was February 1.
A squab was observed in a nest as late as October 18th. Although later
observations were not made, this scjuah |)i-ol)ahly remained in the
nest nn1il tlu' end of October.

BAND-TAII,1.I) IMGEON NESTING 319

Nesting Success

Of the 26 iicstiiius wlii'i'c ()l)S('i-\;il idiis were made, 17 ( ().") percent)
■were considei'od siicccssrul in llcd^iii^' llic s(|iial)s, five; (20 percent)
were niisiiccossfid, and the laic of I'oiir (15 jicrceiit) was unknown. Of
the iinsuceessfnl nests, two were destroyed by avian predation (jays
and raxcns) ; two were destroyed hy unknown causes; one by man;
and one contained a dead (Mnbryo. Mr. I^aidlaw Williams, of Carmel
liiji'hlands, who for numy years lias been engaged in oi-nithological
studies in tliis area, has kindly given the authors additional informa-
tion on mortality factors. Mr. Williams observed a sharp-shinned
hawk (AccijJte?- slriafus) devouring a young squab in the nest, Jnlv
7, lf)4U.

Conversations with Mr. Williams ami other residents in the Carmel
area have brought out the fact that the combination of flimsily con-
structed nests and high winds is i'esponsil)le for considerable losses of
eggs and young birds. ^Ir. Williams has hand reared two young squabs
that have fallen out of nests by feeding them Pablum. Almost every
nesting season young birds are found that have fallen from the nests
due to either Aviiid or the yoinig bird ovei-stopping the nest.

Multiple Nesting

Of unusual interest is the nest designated as Number 1 in Figure 1
and B-1 in Table 2. This nest was found on May 22, 1953. At that
time it contained many droppings, indit-ating that a squab had been
raised in the nest earlier that year. This nest is located in an acacia
tree about 15 feet from the ground. The tree is in a strip between
the sidewalk and street opposite the Carmel Police Station. Members
of the Police Department had observed the nest in 1951 and lf>52.
The officers stated that at least one brood was reared in 1951 and
two broods in 1952.

This nest Avas recheeked on Jnue 5, 1953, when it contained one
eg^'. When the nest was checked on June 8 it contained a downy
nestling. ()n -lune 16 the adult bird was marked on the back feathers
with white airplane dope to enable sight recognition. On June 21, tbe
S(|uab fell from the nest and was banded and weighed. The sipuib
weighed l(i5 grams and was kept until the following day to se(^ if
it was ijijured and then retui-ned to the nest. The squab was last
observed in the nest on June 2fl.

On fJuly 21, the marked adult bird was again on the nest, but the
nest was empty. The nest contained one eug on .Inly 24 ami the
adult was recognized as the l)ird marked on June 1(), Due to other
work it was not ])ossible to follow this lu^stliug closely, but the officers
said the squab was successfully fledged.

On February 18, 1954, an adidt bii-d was on the nest and the nest
contained one egg that appeared to be pipped. ()n I'^ehniary 19 the iu>st
was again cluM-ked and the c^'^ ])roved to be broken rather than
pipped and contained a tlead embroyo in an advanced state of develop-
ment. On iVTarcli 12, the adult hii-d was seen i-earrangiug tlie nest.
The adult was on the nest iiu-ubating one egg on .March 17 and March
23. On Api'il 1 the nest contaiiu'd one s(pial). This nestling was banded
on April 7, and left the Jiest on April 19. The adult was on the nest.

320

CALIFORNIA FISH AND GAME

'"^■O

p.

d -s :s §

_>.

ii^

o e a

1 o2 S

2 If 2

rt

■0

d B 0.

0

2

0

22 S
c d «

0 0

0 0 -*J

0) 4> S

> ^ d

c^ o ** _

4A

t- fc- ^

lO - CO T3
-^ cj fe d • c s

- c -S £ iS ^ -^
5? 0 S =* g ^ d

^

7>

0 0

1

1

^ 1 s -, 1 ^

^00- .-- -3 5 =

^c: s-g d:2 c

d « _S ,^1 "^ ? '•" 0

be
. d

^^

0

G
CO

•o
\^

■0

° ° b

52 «

>,Si Si

£ Z- ^ 'S

—^ in 'n Si

■a cj a; 3

1 "

0 9; c; oi

00 00

t^'-o a<N •< 3 a

tfi (N 10

a

J' d d 0

^ C. 0 00
1* -ti ♦i ♦- *-
0
^ 0 CJ Oi 0

?:• 3 3 3 3

"i 03 01 r! o3
^ G d 3d

■a 3 I- ^ C^ C M

w w-«i w -»: Z

r-l 13 ♦^ *J •„ *J > *;

\-^~ -3aj-dj.Ti ~

■^0-5 -S'^'om'^ -6

33?J

5 ro"

a

S' C G .S

t- 0 0

'S i£ i: «

^333

^ t) -d T3

2:<; < <u\ <

w

z

«<<<;

t3t) l-J t3

3 3

3

3

^^

3

*<-•

3

w

Q

3ms 33C3m

3d33 3r1 333

w

CO

c c c

Cdd3333"C3d

^

u- Q) i*^ t^tM>«*-iC;

V

Wl

^ ^ ^

S S S W ro w rr S i W '

(2

crom mtfiX^cj

w rr w w X 0 vj tfi w

0

(D

wOmm tnrj mcnm

5^

0

t3

000

ooocncoww-. Owe

a;3(Li<D Qjl:^ OOG)

o_i>co c^ 00c
o-ioo o2 coo
3^33 3° 333

3

c c c

ddd<y<aJQ^cjXd3Jd

•Sd-G 2 § § § Sc 23

(N c^ (Mm m

CO •*

>ra

», -^

lO >0 lO lO lO

10 >n

10

-2 a

Q 01

^v, ^^ \^^ ^

-^ ^^ \

O) 10 \

00 1 00 IM O 1 O I

1 1 CO i-H II 0: 1 1

1

1

1 1 1

1 1 1 1 r-l IM 1 1 t CO '

(N 1 -H O) ^ 1 .-C 1

1 1 1 1 -^\ III 1

0 ^^ ^^

to 10 ^^

«

CO 00 00 to t-

t^ 'f

•*

T)

(N ■ C<5 CO

CO CO ^ -^

ira

■* T)l 10

lO lO >o

10 »0 »0 lO

>o

10 "O »o

2 s

\ \^

^\. -1 ^

—■11 1 1 1 <N O

10 1 1 00 II -H cs 1

1

1 1 1

1 1 — 1 1 CO 1 1 IN 1

CO 1 1 , 1 1 rt _(

Csj 1 1 II ,-1

1

1 1 1

1 1 O) Tf 1 1 ~\ . 1 (N 1

-^ w

^ \ ^^

^^

^^^ to ^,

J3

t~ ffl t^

1^ to •* t.

CO

10 to CO

C
0)

U bO bO bO

bt bd bt bo bO be be

be

hf

be bO bC

C wf Kf C 13 C!

ddh.d dd 33

3

c

■>^ 5f C 3 -<^ 3 -"^

~ T3

be

*-tJ

.'-*^

G

C c3 -S ■= cj oj o3 S

CO ra '^ d ^ N ''^ ^ w

d

cS

C-. c-' ^

p-. ff^. 03 ^ 13 13 03 ^ c^. c3 c^.

K 3
03

— 39 5333 —

33g3 33 -233

2

3

•2

-e^'B's-ss ■£

■30? ? cj 0 c -3

oo?tj cjo —00

•3

S

s 3 5 p 5 ^ Z

Bd^d a a ^cd

X

3

c

£mm£^K £

•I

>000 (NOiOOO

iC >o ^ »c to 10 »o >o »o

LO

c^

»c »o »o

«0oi0O-<li00t0i0i0«O

53

CO-tM rHTt<O3(N00

COOS^^i-H ^^ — — — .

-^ T^ 00

COu^(M(NCM-i.-<^lN — -^

III 1 1 1 1 1
III 1 1 1 1 1
III 1 1 1 1 1

CJ ai 0 1 0 Q> Q^ 0

till II III
0) a; ^ 1 II III

1

Cli

0

1 1 1
1 1 t

a» oj 0

1 1 1 1 1 1 1 1 1 1 t
1 1 1 1 1 1 1 1 1 1 1

-3o 0 OOvS.S

c

C g d , d 3 C 3

3 d d 1 II III

c S s

•2

SEE ; E pH £! pH

E E E ; ! 1 111

0

cuEfi.

Q-Cu CuCu

0) ' ai ' '{D<D

>>>>>. [>,>,>,>,

>■>■>■', II 1 1 I

>

>i >i >i

£ £;3 o;3 ' :j;j £ £

s

o

Qj 03 flj ' Qj Qj aj q;

L. t. ^ > t- U U^ U

C; 0) OJ ; J [ III

^

£ £ £

ij

B B U .Si^SwS

Oj OJ U g d g .S .2 .2

e3

-1^

222

22*-'5^.S.'S^^22

CdC ^CGdC

000 »foooo

Ar^ A ■< A A A A

^ "3 ^ "^ '0 '0 "0 *o *o
wik-iwlio 0 f^ 000

'c3
0

0

d d d
000

S S 0 w 0 -< -»; 0 0 § S

CSfNN (NCOCOCOCO

COTj4Tt*CO COt(< -^Tt*-^

>o

CO

CO Tf< ■*

TtlTjlT)<-J<T)l-*-^(<T}<T)<lOlO

^ "2

io»oio tou5»o»oo

»0 >0 'O »0 lO lO IC LO >o

10

10

iO >o 10

10 »o to >o 10 »o 10 10 »o »o »o

^^^>^^ ^^"\^^^-v"^

\

\

"^^■\

-^^^^'^^^^^^^^"-^-^^^

05(N-<J( OSI^COCOCO

COCOCO<N rtOO woot^

CR

00

(N 03 ■>i<

CO— itO02CTCT-*T)<(NC>)C0

C^J'-H^C^J (N^ 'H C^

^ <N

c^ — ,— 1 — eqc^^— , esq

Q o

^\^> ~~^^^^^^^

w\-\ ^^ \^^

^^

\

^ w.

■w-^^\^w^^\

^ »*H

t~oioo oomcotoas

t~ioio>o t~iN eot~o>

<N

10

0> ■* (N

CO-^totototOtotOt-COCO

III 1
III 1

CO (n CO 0} X

-a -S -d -a-a

www

T3 T3 T3

WWII II III
'd "d (C ' II III

d C d G 3

3 d c

C 3 -0 , II III

_rf s! S Si

cj « cj

d d c3 1

o3

3 !d _3 2 ^

3 3 3

3 3—' 11 III

£

M be "5 ;i ~

"3 y sf

M bli^bi' I 11' 111

S5S =S

s s s

£SS 1 I 1 111

COJO '^DC'OCJ

OCJCCJ CC 4)CJ<U

0)

V

0 c; 0

QJQJ^QJQJGJOO^Q;aj

22d ssass

ssss ss ssa

a

a

ssa

sasassssass

tHt-<t-< L4t-<L-<t-<tj

Ut-L-tM t-l(-> (-.t-'I-4

Lri

t4

u t- t^

fc^CCuuCt-CC^-^-

oSoSoQ cdc3^(^<^

cisjojsj o3^ 0j<^03

03

o3

03 ca 03

cdc€cadcQdrarao3da

000 00000

0000 00 000

0

0

000

00000000000

|i

0— o3 ^0 'dOii—

be

030a

0 —I (M CO ■* 'O

^MCO rfiCtpt^OO

0-. — l-H— rt-i rt-lrl

Ol

CO CO T^

tOtOt^OOOli-HrH'Hr-lr-.rH

■< < ■< •< -^

<<<

<<:<«

m

PQ pil

mpi)piipiipiiffl«mfQmm

BAND-TAILED PIGEON NESTING

321

which contained one egg a^ain, on July ft. Tlic sqiKil) lintclicil on
July 12 and was banded on -Inly Ki.

On SeptenilxT 27 ami 2!» an adult witli one c'^'^, was on the nest.
On October 13, the adult was broodint: a squab.

Throughout all of these observations it was possibh' to trll that
it was the same marked pair wliieh was nestinfj and cariu*:- lor the
yonufi-. From these observations it will be seen that a single pair of
pigeons in one season made I'onr nesting attempts and successfully
fledged three birds.

In 1950 an adult bird was seen on the nest for the fifst time Feb-
ruary 9. Some of the marking applied in 1954 was still visible. No

FIGURE 3. Adulf male incubating. Nest located in Monterey pine.
Phofograph by Wallace G. Mocgregor.

322

CALIFORNIA FISH AND GA.ME

ejrg was in the nest. An c^'^' was laid on l"\'bi'uary 11 and this egg
hatelied March L Tlie nestling was banded on March 15, but was
killed by scrub jays ( ApJifJoroDia cnlifornica) on March 22. Xo fur-
thci' activity has been rccui'ded at this nest.

FIGURE 4. Adult male feeding a day-old squab. Phoiograph by Wallace G. Macgregor.

r.A\'n-T.\Tr,Kn imc.ko.v \i:stix(J

323

Incubation

Tliere appoars to be no infontwition <ni tln' iiiciil);it ion period of
the l);iii(l-t;iil since I'.eiidii'o's (]892) statement that "incubation
lasts fiH)in IS to '20 (biys. l)()lh sexes jtssistin'j. " Observations of iiests
in this study indieale that tli(> incubation period avera^-es 1!) (Uiys.

Fifi'ure 3 sliows an aihill nude incubating- a nest in a Abuderey
pine. Observations made of incubatinji' birds eonfinn the work of
earlier worlcers avIio stat(> that the incubation duties are siiared by the
sexes, tlie male iucubatins'- (hiring' tlie day and the female at niglit.

The
down.

Growth of Young

younp' sqnabs wben lialclied are eovei'ed with oran^e-yeUow
The sipiahs are I'vil j)ii.i('oii milk by botli parents. The fol]owin<i-

IS an account

"Th(

this feedinff from the senior author's field notes.

adult wouhl take the beak of the youufi; bird near the base and
<>rasp it in a crosswise ]iositiou ; the adult would then gulp np and
down IS to 25 times. During tlie observations the bird fed five times
with the ft'edings lasting from 30 seconds to two minutes. Three
feedings took place 10 minutes apart." Figure 4 shows an adult nude
feeding a day-old squab. The squabs grow rapidly and by five days
they are almost three times as large as at birth (Figui-e 5). At two
weeks the sqnab weighs half as much as an adult liird and is neai'ly

W

...Oi^^,. ,^^/K

-^'irrrttfTfTfiTrft'

€ -.^'

0

^''-^^A

^

FIGURE 5. Squab, age five days. Phonograph by Wallace G. Macgregor.

324

CALIFORNIA FISH AND GAME

FIGURE 6. Squab, age 14 days. Phofograph fay Wallace G. Mocgregor.

B.\\I)-TAir,i:n I'lOKOX XKSTINO •'{-!'>

full IVathcrod (Fi^'urc (i). Sonic hinls Iriixc the iicst at 1.") days, Wiit
others have bocii observed to remain in thi' nest up to 'Js days. Even
at'tei- the birds l<>av(' tlic nest thi'x arc led at h'ast some (if the time
by the adults.

Tlic youii<>' birds ai-c \'rt\ pi^foii milk lor the first few days and
then a mixture of pigeon milk aud wliatever the adults may be eatinjr.
By the time tlie birds ai-e fiediicd they are jicttiuf? almost straijrbt
regurp-itated adult food and the milk glands have started to resorb.
Thus, by the time the youn^- l)ir(ls are indcpendeid the adults have
little or no evidence of havinji' nnlk <>lauds.

In eonduetinp' cheeks of ])i<>eou hunters in northern ('alit'ornia
durinji' the September season of ID.l'i, appi-oxinuitely (>() percent of
the adult birds liad active milk glands, indicating that many youug
squabs woukl die in the nest if both parents "were shot. At this time
Califoruia had a s])lit season for pigeons, with the first half of the
season running from September 16 through September ;]() and the
second hall' from December 17 through December 31.

The information listed above and the fact that birds were found
nesting in Monterey County into late October resulted in a recommen-
dation that the State be divided into a northern and southern zone,
with an October season in the northern zone and a December season in
the southern zone. This recommendation has since been })ut into effect.

SUMMARY

A nestiug study of baud-tailed pigeons was conducted in the Carmel
area of Monterey County, California, from 1952 iuto 1955. A total of
26 nests was observed, all iu tree locations. In all but one nest, clutch
size was one e<x^. Nesting occurred as early as Fel)i-uary aud as late
as October. Observations on a single pair of birds revealed three
broods successfully fledged during the 1954 nesting season.

REFERENCES
r.fiidirc, Charles

1892. Life lii.storios of Nortli .\nicric;iii l)ii-(is with siifcial rcfcrfMicc (o tlioir
brpcdiiiK hiiliits mikI cufis. \' . S. Nut. .Mus., Spec. P.iill. 1 . p. 122-1 "JT.

Glover, Fred A.

19.5.3. A nestinj; study of the t);iiid-t;iilod piiroou (Cnhiwhn j. fasriatii) in Nortli-
western California. Calif. Fish and Game, vol. 39, no. ?>, p. 397-407.

Griiincll. .Tuscpli

191.3. The outlook for conserving the band-tailt'd pitceon as a stanio liird of
California. Condor, vol. I."), no. 1, p. 2.~i-4(K

Grinnell, .Joseph, Harold C. Bryant, and Tracy I. Stoier

iniS. The Kaino l.irds ..f California. r..Tl<.'l..y. T'liiv. Calif. I'r.-ss, i>. r)7.")-.">SS;.

Leopold, Aldo

19.33. Game maangenient. Xiw Vorl<, Cji.-iilcs Scrilini'i's Sons, (si p.

■Morse, William R.

1949. TIk' liand-taili'd iiigi-ou. Oregon Si. (Jainr ('niiiin. I'.nll.. \'>\. I. ii". 7.

p. 1. ■">. 7.
19.")1. Special rr|i(irl h.-nid i.i ilrd pigeon season. Oregon St. (ianie Coniin.. 7 p.

Xaether, Carl A.

19.39. The l.nok of til.' ].i:;r.iii. Pliil.i., D.ixiil .McKay Co.. 2r.s |>.

326 CALIFORNIA FISII AXU (iA.ME

Xrfl'. .Ii)llliS(ili A.

1'J47. II;il(ils, foud, ;ni(l ci-oiKniiic status ul' tlic l):uul-tail('tl pijjeuii. U. S. Fish
Wildl. Serv., No. Ainer. Fauna 58, 70 p.
Riildle, Oscar

inns. Studii's on the physiology and reproduction in liinls. Anier. Jour. l'hynioI.,
y(d. SO, no. 2, p. 248-205.

Whitman. Charles Otis

IKlll. The liclia\i()r uf pigeons. In: Carnegie Inst, of Wash.. I'uhl. 257, vel. 3,
101 p.

SKELETAL DIFFERENCES BETWEEN DEER,
SHEEP, AND GOATS'

MILTON HILDEBRAND

Department of Zoology

University of California, Davis

INTRODUCTION

Agencies charged with the enforcement of game laws occasionally
must identify frozen, canned, or fresh dressed meat which is believed to
be from illegal deer, but which is claimed by a defendant to be from
domestic sheep or goat. New methods of blood analysis provide reliable
identiiication in many instances, but determinations must sometimes be
made from the skeleton. Only osteologists with access to an adeijuate
collection of skeletons have been able to make the identifications. This
paper has been written to enable any individual familiar with the skele-
ton and the fundamentals of statistics to distinguish bones of deer from
those of sheep or goats without known skeletons for comparison.

Archaeologists have fonnd it difficult to distinguish these domestic
animals by pieces of broken bones. I have discovered no characteristic
which separates all domestic sheep from all domestic goats, but the
skeletons of the two animals differ significantly in the average form of
many characters. Single features serve to identify many individuals,
while combinations of characters make it possible to discriminate nearly
any skeleton.

This study makes only a small contribution to the broader problem
of finding generic differences between the postcranial skeletons of Ovis
and Capra. A search for generic characters is in progress b^' Barbara
Lawrence and one of her students.

I have drawn freely upon the paper on the skeletons of deer, sheep-
goat, and several other animals published by Lawrence (1951) to assist
archaeologists in the identification of l)one fragments. My study, based
on nearly twice as many skeletons, adds criteria not included in her
paper, omits others found to be unreliable, and restates all criteria so
that identifications can be made without comparative material.

Skeletons of 10 deer (Odocoilcus hrwiouus cohimhia)ius), 8 sheep
(Ovis aries), and 9 goats were examinrd. Most of the sheep were of the
Corriedale breed, but the breed of several was not known ; the goats
included Angora {Capra falcnneri) and one or more breeds of Capra
hirca. Each sample included aninuds of both sexes and several age
classes. Uidess otherwise qualified, tlio ratios and descriptions presented
apply to any animal old ('iioiil:1i fdiinnonly to become ;i Imnter's target,
whether fully matnre or not.

The skull and the boues of the feet arc (niiittrd because they are not
preserved with dressed carcasses. Skulls of these animals are not difficult

1 Submitted for publication March, 1955.

( :'.-J7 I

328 CALIFORNIA FISTT AND GAME

to distinguish; Glass (1951) describes several diagnostic characters.
Kil)s are not distinctive. Vertebrae (except the first two and the sac-
rum) are omitted, becau.se it may be difficult to assign an isolated ver-
tebra to its correct position in the spine; comparisons might therefore
be invalid.

I am indebted to the IMuseum of Vertebrate Zoology of the University
of California for the loan of 14 skeletons. Six skeletons were borrowed
from the Department of Zoology, University of California, Davis, and
one from the School of Veterinary Medicine, University of California,
Davis. The remainder belong to the author.

Thanks are extended to K. D. Taber, School of Forestry, University
of California, Berkeley, for valuable suggestions, and, particularly, to
nij^ wife, Viola ^M. Hildebrand, for taking all measurements, calculating
ratios, and drawing the figures.

RELIABILITY OF IDENTIFICATIONS

A bone identification which will be used as evidence in court must
have a high degree of certainty. Several considerations bespeak caution.
Individual variation among different skeletons of the same kind of
animal is greater than commonly realized. It is helpful to compare a
bone of disputed identity with an illustration or known skeleton ; how-
ever, no skeleton, or even single bone, can be expected to approximate
average form in all characteristics. Differences may be detected from
meager material, but the reliability of differences must be determined
from larger collections. Few museums have as many skeletons of deer,
sheep, and goats as were available to me, yet my samples were small.
Ratios usually provide a more satisfactory means of analysis than do
measurements of size, because the latter do not make adequate allow-
ance for exceptionally large specimens, and are subject to more bias
resulting from discrepancies in the sex and age composition of samples.

Many differences betweeu the skeletons of deer and those of the do-
mestic animals, and nearly all differences between sheep and goats, may
be expressed only as variations in the average or typical expression of
characters for which extreme values ovei-lap. Most size differences are
of this nature. Can i-eliable identifications be based upou such criteria?
Yes, often they can if the method of analysis is valid.

Assume we have the scapula of a deer which a defendant claims
is from a sheep. Elsewhere in this paper one can find three ratios
which are useful in distinguishing deer from sheep bj- this bone (see

Tf f C'

— , -7^, and fr, Figure 1;. Suppose that our bone has a value for
D \j U

the first ratio which is two standard deviations fi-om the mean for

sheep ( — =^ 2). The jn-obability that a member of a population would
cr

differ this iiiurli from tin' incaii of the po]iulatioii can be read from a

table of areas of tlic udi'inal curNc foi- \'ai-ions values of — {e.g., Simp-

(j

son and Roe, 1939, page 137). It is found llial tlie diance a sheep taken

from a flock at random would deviate In or more from the mean for

sheep is 4.5 percent. Stated another way, one sheep in 22 Avould differ

so much from the mean for sheep in general.

SKELETONS OF DEEK, SHEEP, AND OOATS 320

NoAV if the second ratio is calculated, and is also 2a from the mean,
the chance that a sheep taken at landoin would at the same time difTci-
this niiicli fi-oni the moan values for each fatio is rodncod to one in (ill.
(The ratios must be indepenth'iit ; the same measui-ement may not be
used in each. If one ratio exju'esses the slender character of a bone it
might not be independent fioni a similar ratio for another bone, because
one animal could be delicate in all its skeleton.)

11" our scapula chanced to deviate i'rom the mean by a like amount in
the third iudei)endent ratio, then the chance a particular sheep would
simultaneously vary so much from all three average sheep proportions
Avould be one in 20;"). By combining meager odds we would have thrown
sofions doubt upon the defendant's claim. The odds re(iuired to estab-
lish adequate identification is, for our purposes, a legal rather than
an anatomical j)roblem, but it will often ])e possible to do much better
than in the above example.

Suppose the disj)uted scapula, claimed to be a sheep bone but really
from a deer, chanced to have values of the three ratios cited which
were equal to the actual means for deer. These differ from the means
calculated for sheep by 3.92, 2.54, and 2.38 standard deviations, respec-
tively. The chances that a particular sheep would vary so much from
mean proportions of sheep are 1 in 10,000, 1 in 91, and 1 in 59, respec-
tively, if the ratios are considered independently. The odds for the first
ratio are such that the defendant's claim is seen to be fraudulent;
however, to carry our example further, let us assume that the spine of
the scapula is broken so that this ratio is not available. The odds of
1 in 91 and 1 in 5i) remain. Taken individually these make the defend-
ant's claim doubtful, but one vrould hesitate to convict on these grounds.
HoAvever, the chance that a given sheep would simultaneously differ so
much from mean sheep values for both ratios is only one in 357.

This analysis indicates that the bone is not from a sheep, but does
not establish it as a deer bone. To do this one must show that its propor-
tions are typical of deer. The text and illustration also provide other
characteristics of the bone which should support our conclusion.

METHOD OF ANALYSIS

The significance of the product of several independent probabilities
is calculated bv an application of chi square analysis as explained by
Kenall (1948, pages 132-133) and Fisher (1936, pages 105-106). The

E

last of the above examples was solved as follows. For the ratios -^ and

G

^, the means for deer differ from those of sheep by 0.33 and 0.19,

respectively. Dividing by the standard deviations for sheep and ex-
pressing the quotients as percentages we derive probabilities (P) of
■5.011 and 0.017. From mathematical tables the natural logarithms of
these parameters are -4.07454 and - 4.50986. Substituting in the e(|ua-
tiou y- = 2S lou', T\ we find tliat the y- value ('(piivalent to the derived
combined probability is 17.16880. Many x^ tables do not include such
large values, but see, for instance, Pearson and Hartley (1954, Table 8),
where it is found that with four deirfiM>s of fi-eedom (two for each
variable) the desired P is between O.OOl and U.005. By interpolation

330 CATJFORXIA FTSTT AXD CAME

(for method see Fisher, Inc. cit.), the required probability is 0.0028,
Mhieh may be expressed as odds of about 1 to 357.

Tlic {lajros Avhich follow jrive for each ratio the number of specimens
ill llic sample (.V). the arithmetic mean (M), aiul standard deviation
(o-). From these parameters the reader can calculate standard errors
of the mean, probabilities, and the significance of the product of several
independent i)r()bal)ililies.

The animals stndietl may be assumed to have about equal variability
in re^^ard to any given ratio. Since the samples are small, it is desirable
to apply to each animal a standard deviation which is a little greater
than the average of the three calculated standard deviations. Thus for

E

the ])elvis ratio ^ of sheep, a = 0.10. but noting that in deer a = 0.29,

and in goats ct ^ 0.22, it would Ix' preferable to use a value of about
0.22 for any of the three animals.

Measurements were made to the nearest millimeter.

SCAPULA
(Figure 1)
Deer and sheep can nearly always be distinguished by the scapula,
and this bone often serves to set deer apart from goats, or goats from
sheep.

The ratio of the length of the spine to its depth separates most deer
from sheep and goats, but does not distinguish the domestic animals.

Ratio jr; deer : .V = 8, M = 4.90, a = 0.32 ; sheep : N = 8, M = 3.92,

(7 — 0.25 ; goats : N = 8, M = 4.00, a = 0.20. The ratio tends to rise
slightly with advancing age.

Dividing the de|)th of the bone's neck by its thickness gives a ratio
Asliicli often distinguishes deer from sheep, but is of less value for dis-

. E

criminating deer from goats, oi- goats from sheep. Ratio-^; deer:

N = 10, M = l.(i(), ,r^().l;5; sheep: .V = 8, 3/ = 1.93, (7 = 0.13; goats:
N = 8,M = 1.79, (7 = 0.14.

The ratio between the depth and bi'cadth of the head of the scapula
separates typical sheep from both deer and goats with reasonable as-
sur-ance. Not many single characteristics of tlie body skeleton are better
for distinguishing sheep from goats; their means differ by about 2.4

C

standard deviations. Ratio — ; deer : N = 9, M = 1.44, o- = 0.08 ; sheep :

//

N = 8, M = 1.63, a = O.O.S ; goats : N = 8, M = 1.43, a = 0.04. The ratio

tends to be lower in large animals than in small (young) ones.

Tlic relation between the length and width of the bone is nearly
identical for deer and goats, but is sufficiently dit!'ei'ent for sheep so
that individual animals having particularly low or high values of the

ratio can be designated sheep or not sheep. Ratio -^; deer : N = 10, M =

l.fil . ,r = 0.0(i ; sheep : .V = 8, M = 1 .4."), ,7 = 0.11 ; goats : N = 8, M =
1.57, ,7 = 0.07.

SKELETONS OF DEER, SHKKI', AM) COATS

A

3:11

(A

end. view

This an<jle is more tc/pical
of shicep and cjoats but /j>
present in some deer.

Deer Sheep or
Goat

As seen in cross
section this concavity
\ is usually deeper-
than ii) deer.

ee.

Deer Goat

(Cross sections ttirough mi
Edge of the
spine is rela-
tively rounded,
straig/it, and
not curled
under.

Edye of the
spine is usually
curled under
slight It^ but
more so in old
aniinals than
in young adults

Sheep
ddle of spine)
Tlic spine is
usually flallcr
on the edge
than in deer
or goats, and
more curled
under than in
goats

'P

As seen in cross
section this border
is usually more
arigkd than in
sheep and goats

1

I^eer

FIGURE 1. Right scapula, in lateral, anterior, and other views as indicated.

The fi<riii-t' illustrates (littVfoiiccs l)(>tweeii typical dcor and sh(>op or
floats in tlic i'onu of the aiiterioi- boi-dcf of the scapula, tlie sha]io of
the etlj^e of the spine near its niidjioint. jiikI the curvature and thii-kness
of the vertebral border df the hdue. Eai'li of tlu'sc characters is subject

332

CALIFORNIA FISH AND GAME

to individual vai'iation and iki one of lliriii lakcii alone furnishes certain
identification.

Absoliitt! size of tiic shoulder blade, and si raightness of its spine are
not satisfaetory eiiaraeters for the identification of specimens.

This angle and
this elevation are

i/xinablc and do
nut serine to
d/stincjuish the
three animals.

This tuberosity is
usually more buiku
in sheep and goats
than in deer

Deer

Subtle differences in shape
of muscle scors distinquish
tupical deer from sheep
and goats but are not
infallible criteria. .

Presence or absence
of groove here is rot
\a reliable characler

Goat

This ridge Is slightly
higher with steeper
walls in deer than
in sheep or' goats.

\t

These grooves
cue a lit tic
deeper in deer
than in sheep
or goats.

Shteep

FIGURE 2. Right humerus, showing anterior and lateral aspects.

HUMERUS
(Figure 2)

It is always possible to distinguish deer from sheep by the humerus.
In all characters studied the goat is intermediate between the other
a.ninials, but average values lie closer to the sheep. Goats and deer can
usually be separated ; sheep and goats cannot always be assigned.

The humerus is longer in adult deer than in sheep. Selecting animals
old enough to have the distal epiphysis fused to the shaft, seven deer
and eiglit sheep had bones averaging 198 and 149 mm., respectively.
There was no overlap of values, but overlap would be expected in larger
Fam])les. Since the longer bones of deer are also straighter and more
slender, length can be combined with other measurements to calculate
very diagnostic ratios which are not dependent upon age determination.

SKKLKTOXS OI' DKKK. SIIi:i:r'. AM) GOATS .13.'!

Of pjirl iculiii- \;ilii(' is llic i-;iti(i of (i\(M--;i1I lcii<j-tli t(i \\\r iiicnsiii

•e

III' iMir\;it II ri' slinwii in the liuiii'r. IJjilid • dcri': A ^ Id. .1/ 14..),

It

,r=l.l:!; sliccp: .\ S, .1/ U.S. ,r 0.4:!: ^dJlts: .V = H, M =]].],

<r^<).T(). If tlir s1;iii(l;ii(l (lc\i;itinii for the slicc|i s;iiii|)l<' is taUcii more

(•()n"('i-v;ili\cly to l)r O.H. then only ;il)ont 15 sliccp in I.OOO would be

expected to have a ratio as liii:li as 12. 0. yet oidy about 1'J dfr-r in

1,U00 would have .so low a value.

The head of the bone, aud its "zrcatcr tuberosity, are heavier in slieei)

aud ^oats than in (\<'t'r. The ditfereiiee, notieeable to the eye, eau l»e

e.\pressed accurately li\- I'elaliuL'- the breadth of the bead to the Icnirth

of the l)one. Ivatio ' ; ilt-rv : .\ - l'», .)/ ^4.55, <r = 0.24 ; sheep; .\ = 8,

M = 8.4(i, IT = 0.17 ; jjoats: .V == 8, M = 3.83, a = O.l:^ Sli<rhtly hi^dier
A'aliies may be expected in youu^' animals tlian in old.

Over-all lcu<rtb can be divided by least breadth of the .shaft to provide

an cipially useful pi'()p(U'1 ion. Katio y-; deer: .Y = 10, 3/ = 11.4, tr =

0.(i.'): .sheep : .V := 8, 4/ = <).0:5, <r = 0.(i4; <;oats : .V = 8, 4/ = 0.71,

a = 0.32.

A lateral view of the bone reveals the deltoid crest ;ii-chin^- downward
aud foi-wai'd undei' the head. The crest is related to muscle attaeluneiits
and is more evident iu lully mature animals. In sheep and <roats it runs
onto the anterior sni-facc of the shaft relatively closer to the head of
the bone than iu dcci-, so the distance E, shown in the fi<rure, is shorter
thau /•' : in deer E is equal to. or lou^'ci- than F.

DitTereiu-es in the distal ai-ticulatory surfaces, desci'ibcd by Law-
rence (19ol) and shown iu Fi«>ure 2. were found reliable; others she
rcc(\uuized for the head of the bone were unreliable or too subtle to
use with confidence in assi<iui:i.; the bones of my samples.

RADIUS
(Figure 3)
The radius serves to distinguish deci' from sleep or .i:oats, but does
not characterize the domestic animals.

Deer have the longest radius. All adult a:;i:.'.:ds in my samid- la 1
bones measuring 185 mm. or more in total length; for.r were 204 mm.
or more. The bones of one sheep and one goat equalled 185 mm. and
none reached 200 mm. Since deer do not have wider bones, a ratio
of length to width of shaft is a more accurate index than actual length.

Ratio ^; deer : A = 10. 4/ = 10.85, a = 0.62 ; sheep : N = S, M = 8.59,

<r = 0.41 ; goats : N = 8, 31 = 8.43, a = 0.45.

Lawrence (op. cit.) noted that the proximal head of the radius is
relatively broad and flat in deer. This proportion was studied by divid-
ing the breadth of the head by its maximum depth. The following
figures show that there is so much overlap of values that only par-
ticularlv low or high ratios should be considered significant. Ratio of
C to depth of head ; deer : .V = 9. 4/ = 1 .(i!». a = 0.0!) ; sheep : X = 8,
M = l.llO, .r = 0.07 ; goats : iV = 7, 4/ = 1.81, a = 0.08.

5—20860

334

CALU'OKMA FISH AND GAME

Fni'tlicr (linVreiiccs Ix'twccn llip animals are shown in Figure 3. A
reliable t'eatui-e is the picseiiee in slieej) and goats- only of a projecting
lip on the proximal end of the bone. The d(>ptli of a noteh which ac-
coiiiinndates a process of the ulna is h'ss reliable; 1y|)ieal coiitoiii's are

fhis articulately These pits are conspicuously This lip is

surface is smaller wider and. deeper In /characteristic

in sheep and goats sheep and cjoats. / of sheep andgoctts,

~ —' l:)ut is absent //)

deer.

fhis notch tends
to be deeper in
deer, but is not
a reliable
criterion .

This articulutoiu I*
surface is larger
In sheep and goats.

Deer

This suture between
radius and ulna is
more horizontal in
deer.

Til is grooi/e /s alujaijs
deeper in deer than
in sheep and goats

Tile scar showing area of
contact with ulna is
narrower, deeper and less
rounded ai its
proMmai end than in
sheep or goats.

FIGURE 3. Left: distal end view of articulated right radius and ulna.
Right: posterior surface of right radius.

illustrated. The characters figured at the distal head of the bone pro-
vide reliable identification of deer.

The radius and ulna remain free in all but old deer, but fuse in
sub-adult slieej) and goats. The area of contact between the bones,
visible in disarticulated specimens as a scar on the shaft of the radius,
is diagnostic.

ULNA

(Figure 4)

The ulna will neai-ly always set deer apart from the domestic an-
imals. Several characters discriminate the goats of my sample from
the sheep ; I cannot say that these criteria would always be reliable.

Differences in |)ro])ortions of the articulatory surface at the elbow
j(nnt may be e.xpi-essed by relating two measurements shown in the

illustration. Ratio '^, deer : N ■=■ 10, M -- 0.93, a = 0.05 ; sheep : iV =

x>

8, M =: 1.05, (T =: 0.04. Since the mean i'or deer is three standard devia-
tions from the mean for sheep, this ratio will often ])rovide reliable
identification. Measurement ^4 could be taken on only two goats be-
cause in this animal the ulna is usually fused to the radius. Even in
young specimens the suture between the bones is obscured, but the
ulna seems to extend laterally farther under the radius than in sheep
or (leer. N'alues of the ratio for the two measurable goats were 1.25 and
1, •')!). Their mean is nearly seven staii(hinl (h'viations I'l-om thai for
sheep-^a vei-y striking difference.

SKELETONS OF DEER, SHEEP, AND GOATS

335

The olecranon Vie shape of this TIic surface ujIucH

process is straight in process complements articulates u/ith the

deer and usuallif that of the notch radius is narrou/ in

inflected in sheep. shown for radius deer but broad and

in Figure 3. rough in ^/)ccp.

Tfie ulna and radius
are fused here and
elsewhere in most
goats.

/\ notch is evident This end of the ulna

here in sheep and is heai/ier in sheep

goats but not in djeer and qoats and more

concave here than in

deer.

FIGURE 4. Above: anterior surface of right ulna. Below: right ulna in loterial view

(ulna of goat fused to radius).

The olecranon process is deeper in relation to its thickness in deer

(J
than in tlie domestic animals. Ratio — ; deer: .V rr 10, If =r 3.41, a ^

0.22 ; sheep : N = 8, M = 2.75, a = 0.12 ; goats : JV = 8, M = 2.65, a —
0.25. This process is in line with the shaft of the ulna in deer. It is
usually straight in goats, and is usually, but not always, inflected in
sheep. A bone with an inflected olecranon process could not be from
a deer; one with a straight process could be from a sheep.

Diagnostic characters of the shaft are related to the greater tendency
of the ulna and radius to fuse in the domestic animals. Tliese features
are shown on Figure 4, and differences seen at tlic distal end of the
bone are illustrated there and on Figure 3.

METACARPAL
(Figure 5)
The goat is intermediate between deer and slieep in the expression
of most skeletal characters, but the sheep stands between deer and
goat for the most diagnostic features of the metacarpal. The bone offers
sure identification of deer from goats in ;dl but very exceptional in-
stances, and often will disfiueiiish sheep ti-oni the other aninuds.

The metacafpal and metatarsal are each foi-med by the fusion of
two metapodials and are of similar apiiearance. The metacarpal is

330

CAMFOKMA I'ISII AXI)<IA:\IK

O^ Qj

r "^ QJ

- r- ^

^ L ^ "•> n\

-§.5 0 0^

•^^^ q;^

n- f^ ^ f^ v^

QJ §

^ S 3 o

ill I

0

0

~-

C3

Cj

Q.

t^

Q

d

Q)

u

e^

-to

FIGURE 5. Right mefapodial bones, figured in posterior and proximal end

views except as noted.

SKi;i;i:'r().\s (w i)i:ii;. siii;i:i', .wn coats Ji37

i-ccoLiiii/.cd by its shoi-tiT lcii,i;tlij iiiid by ihc cdiiliu iifii' ii'ii nf llic jirox-
iiiial ;ir1 iriil;i1(i|-y siii-facos.

Tlic iiictacarpal n\' dcci- is iiiorc slcndi'f lliaii tliaf of sliccp, and this,
ill turn, is usuallx' more slender than the metacarpal ol" j^oats. This
pi-opoftion is analyzed by relatinu' the h'MLith of the bone to its broadth.

Katio -'^■, door: N = 9, M = 11.73, ,t = O.til ; sheep: .V = 8, M = 8.29,

a = 0.6(i ; poats : .V r= S. .¥ t= 6.46, a = 0.54.

A seeond propoi'tion is ('(pially useful. A cross section of the bone
taken at the niidjioint of its length sho\vs a roupidy cii'enlai- outline in
deer, but is eomprossed from I'l-ont to back in sluM'p. and is e\-en tlattci-

in goats. Ratio ^ ; deer : N = 10, M = 1.07, a = 0.07 ; sheep -. N = S,

M = 1.43, a = 0.09 ; o-oats : A^ = 8, .¥ = 1.63, a = 0.06.

Tlie illustration shows that at its midpoint the |)osterior surface of
the shaft of the bone is eonspieuonsly eonea\-e in deer, lint only sliijlitly
concave in the domestic animals.

INNOMINATE
(Figure 6)

]\rany eharactei-istics of the innonnnate distinguish deer from the
other animals and often permit the distinction of sheep from goats.
However, individual variation is striking and there are sexual differ-
ences, so caution must be exercised.

The ]iul)lic symphysis of deer is longer, but the ischium is also longer
and more slender, so the ischial arch is relativel\' deeper and extends
anterior to a line (straightedge) joining the anterior mai-gins of the
ischial tuberosities. In the other animals the arch just reaches, or falls
short of, this line. By assuming that the line joining the anterior
margins of these tuberosities is about at i-iglit angles to the pubic sym-
physis one can apply this criterion of identification to a single innom-
inate bone not yet fused to its fellow. It should be iu)ted that the sym-
physis is not always exactly in the sagittal plane, and that its plane
may be difficult to determine for young specimens lacking the wedge-
shaped ossification which will foi'iu in the postoi'ioi' I'lid of the suture.

To demonstrate that the ])ubric symphysis is longest in deer, its
length can be divided by the least breadth of the .symphyseal lannis of
the ischium. The ratio is higher for immature animals. The length of the
s,ymphysis cannot be determined from some disarticulated pelves for
which ossification at the postei-ior end of the suture is incomplete.

Ratio—; deer: iV = 6, M = 6.58, a- = 1.52 (but adding 3 paitly grown

animals M = 8.06, a = 2.76) ; sheep: N = 8. M = 3.71, ,r = 0.39; goats:
N = 9, M = 5.29, a = 2.00.

The i-(dationship of two measurements of the ischinm serves to de-

tci-minc mosi (h'cr from the domestic animals, b'atio ; deer: ,V ^ (5.

Vl/ = 1.S5, ,r=r0.16; sheep: .V = 8, M= l.O-l. .rir^O.lI; goats : .V = 7,
M = 1.19. ,r^().14.

838

('Ar.lKOHXIA IMSH AND GAME

.0

c: Co

u

o

■ 2^1

^ O IT) <

d) CO ^ o -

a ^ . o
C C c.

C o ^ ^"^

FIGURE 6. Pelvis, seen in right lateral, ventrolateral, and posterior views.

SKELETONS OF DEER, SHEEP, AND GOATS ."'M!!

Deer liave a iiaiTowcr prKis lli;in do sliccji or nKisf lioats. .Measure-
ments E and F are made \'v the center of the aeetahnhn- notch.

Katio -^; deer: N = 9, M = 2.72. ,r = 0.29 ; sheep : N = H, M = 1.74.
r

c7 = 0.10; goats: X=:S, J/ = l.!»!l. <r = 0.22. This ratio tends to he

smaller in adnlt females beeau.sc in them the pnbic bone is relatively

long (IJ. 1). Taber, MS).

When the iscliial tul)erosit ies are viewed from beliind, a dift'erenee
is seen between deer and shee;) as illustrated; goats are intermediate.

At the level of the posterior margin of the obturator foramen the
ramns of the ischium turns upward more in most deer than in the
domestic animals. Measure // from the rim of the aeetabuliini to a

straightedge placed as shown. Katio — ; deer: N = i), J/ = ii.Ofi,

o- = 1.59 ; sheep : N = S, M = 18.80, a = 3.90 ; goats : N = 8, M =

n.fi4. fr = 1.12.

The pit on the ventral border of the ilium just anterior to the
acetabulnm is broader and deeper in typical deer than in sheep or
goats, but there is some overlap. Measured from its inside lip the pit
is usually more than 4 mm. deep in deer and may be as deep as 7 mm.
In goats it is usually shallower than 4 mm., and in sheep usually less
than 3 mm.

The ramus of the ilium is clearly flatter in deer tlinn in sheep; goats

are again intermediate. Ratio -y ; deer : N =: 10, M = 2.24, a = 0.14 ;

sheep : N = S, M= 1.46, a = 0.12 ; goats : N = 9, M= 1.78, a = 0.15.

The shape of the expanded portion of the ilium is useful in making
determinations. In deer the margin is more convex on its ]iosterior
profile and more concave on its anterior profile than in the domestic
animals. The anterior concavity can be measured as indicated in the
illustration; its depth would rarely be less than 4.5 mm. in adult deer,
or more than this in sheep or goats. Deer: N = 1, M = 6.31, o- = 2.23
(but excluding 2 immature animals 31 = 7.64, o- = 0.90) ; sheep: N =
8, M = 0.69, (T = 0.55 ; goats : JV = 8, M = 1 .56, a = 0.94.

In my material a more or less evident notch marks the anterior border
of the obturator foramen in deer and goats but not in .sheep. This
chai-acteristic sliould be checked in additional specimens.

FEMUR
(Figure 7)
The femur is much less satisfactoi\v than most of the other long
bones for distinguishing the three animals. It is usually more slender
in deer than in sheep. If the ratio of over-all length to breadth of
shaft is less than 10.7, sheep is indicated; if it is over 12.0. deer is
])robable. TJoat is intermediate and sliould not be identified on the basis

of this pi-o|>or lion, b'ntio^ ; deer : ,V - 10, iV = 12.47, a = 0.57; sheep :

X)

N = 8, M = 10.23, (T = 0.49 ; goats : .V = 9, .V = 10.98, .r = 0.77.

The angle between the head of the bone and its greater trochanter
is usuallv less than 90 degrees in goats and deer, but more than 90

340

CALIFUKMA FISH ASD UA.MK

T/y's angle b usually less
than 10° in gout5 and
deer- but n)ore than
90" in sheep.

The slope and length
of these ridcjes are
not reliable criteria.

/} groove or eroded
area here is typical
of gocds, i/ariablc in
sheep and U/OS not
found in deer.

Differences in this
profile ore not satisfac-
tory for ossiyniny
single specimens.

These muscle bcors
are coarser m adcilt
deer than in typical
sheep or guats.

anterior view

Deer

posterior i/teuy

Sheep

/? sharp projection
here is more common
in deer but is variable.

This fore

ilk

his roranien is usually
closer to the proximal
end of the bone; and
more onto its posterior
surface in the domestic
annnalb.

Differences in the lateral
malleolus are not satis-
factoru for assiynin
single specimens.

'9

The development of
these points is not
a basis for identifi-
cation.

anterior vieu/

Deer

Sheep

A/lore of this surface is
seen from this uieu/ in
deer than in typical
sheep or qoats.

FIGURE 7. Above: right femur. Below: right tibia.

SKKLK'l'OXS OI' 1)I:I:K, SIIKKP, AM) (iOATS '.>A]

depTOes ill slii'cp. Sprci iiiciis li;i\iiii;' tlii' angle iii)[)i'<»a('li iiig DO dcgrcfs
should Mill he assigned on this hasis.

Ahove the distal condxlcs arc tiic suiJi'McoiidyJoid ci'csls which mark
the origins of muscles wliicli Ilex the foot. 'IMiese scars, a ridge het ween
them, and a concavity distal Id the hiteral sear are all more [jromineiit
in adult deer than in most sheep and goats; however, muscle scai-s are
in general hii;lil\' variable and I cannot reconunend these featui-es as a
basis for identification unh'ss age differences can be carefully con-
sidered.

All goats of my material have a crease or eroded area in the patellar
gi'oove. One sheep has a similar featui-e ; otherwise it was absent trom
sheep and deer. This unexpected charactei- may prove useriil in dis-
tiuguishing goats, but should be checked in more sj)ecimens.

TIBIA
(Figure 7)

l^ike the femur, the tibia olfers few criteria for distinguishing the
aninuds. It is usually longer in deer. If the over-all length is more
than 27.5 cm. the bone is probably that of a deer (but a large goat
tibia in my material measures 27.2 cm.) ; if it is less than 2'^ em., and is
from an adult aninuil (having ei)iphyses fused to the shaft j it is too
short to be deer.

Lawrence (op. cit.) noted that when the ti!)ia is seen in anterioi"
view, the medial malleolus appears thicker in deer than in otlier animals
(distance C in the illustration ). This is true also of my material, yet its
thickness correlates with, and is not appreciably more vai-iable tiian.
total bone length. Actual measuremeiits for seven adult deer ranged
trom 4.8 to 6 nun., for eight sheep from 4 to 4.."i nun., and for eigiit
goats from 3.7 to 4.8 mm.

Part of the smooth inner wall of the medial malleolus can 1 e s-eii
when the tibia is examined fi-om in front. The extent of the surface
exposed to this view diffei-s with the animals. In no deer was less of the
inner w'all evident than in the specimen shown in Figure 7; selected
sheep and goats show as much of the articnialoi-y surface, but more
often these animals are about as illnst I'ated.

I'ulike the other long bones, the tibia is not appreciably more slender
in deer than in the other animals. The graceful curve of ilu' shaft is
more pronounced in the longer examples, and hence is more character-
istic of deer than of the domestic aninuds.

There is a nutrient foi'anu'u on the lateral surface of the bone about
one tliii'd of its length from the proximal end. In my specinu us. as in
those of Lawrence (op. cit.). this foramen tends to l)c elo>ei- to the
proximal end of the bone, and is also ])laced more onto its posterioi- sur-
face in sheep and goats than in deer, llowexci'. the location of a nu-
trient foramen may be expected to vary and I agi-ee that this feature
does not lu'ox'ide I'eliablc i(h'nt ificat ion.

METATARSAL
(Figure 5)
The metatarsal of i\i'r\- is so (listiiicti\-e that it always sets this animal
apart from sheep and goats. The domestic animals can sometimes be
distinunished bv this bone.

:{42 CALIFORNIA PISH AND GAME

A conspicuous j>:roove marks tlic aiitcrioi- surface of the shaft in doer.
One to one and a half inclies fi-oiii the distal end of the nietatai-sal. this
groove terminates ahruptly in a foram<'u which tunnels throujih the
bone to emerfrc in llie cleft between llic condyles. These characters are
entirely lackinji- in sheep and goats.

That deer have a more slender metatarsal can be demonstrated by the
ratio of over-all length to breadth of shaft. Mean values for deer and
goats differ by about eight standard deviations; only one animal in
10. 000 would be expected to deviate from the mean for its species as
much as the midpoint between the means of the two species. The sheep
is intermediate in its proportions, as was true also for the correspond-
ing metacarpal i-atio. Ratio— ; deer: N = 9, M = 14.34, a = 0.51;

Ml

sheep :N=8,M = 10.26, a = 0.G4; goats : N = 8, M = 8.30, a = 0.74.

The metatarsal of deer is much the longest. My material suggests that
only in exceptionally large sheep or goats would the bone measure as
much as 16 cm. Even juvenile deer usually have a longer metatarsal,
and if the distal epiphysis has fused to the shaft the bone may be ex-
pected to exceed 18 cm.

On the posterior surface of the bone, about half an inch from its
})r()\imal end, there is an evident foramen in deer w^hich is not found
in the other animals.

The metatarsal is formed by the fusion in embryonic life of the third
and fourth metatarsals of the ancestral compliment of five bones. The
second has become a vestigial bone accommodated in sheep by a smooth
articulatory surface on tlie postci-ior face of the head of the metatarsal
(see illustration). This surface could be identified in only four of nine
goats examined, and was not seen in deer.

It was shown that the shaft of the metacarpal is somewhat rounded
in deer, bvit more compressed in .sheep and, particularly, in goats ;
there was also a deeper concavity on the posterior surface of the shaft
in deer bones. These features apply also to the metatarsal, but are
here much less marked.

SACRUM
(Figure 8)

The sacrum of deer differs from that of the otlier animals in the
typical expression of half a dozen characters.

The transverse processes of the first sacral \-ci-tebra extend anterior
to the centrum in all deei-, but oidy in few examples of sheep aiul goats.
It may be concluded that if these processes do not extend anterior to
the centrum the specimen is not deer.

If a straightedge is placed on the lower surface of the sacrum, across
its long axis, and at the level of the center of the fourth vertebra, it
is see]) that the transverse processes extend ventral to the centrum in
deer, but not in sheep or goats (see cross sectional drawings in the
illustration). The ventral surface of this centrum is relatively flat in
deer, and usually convex in the domestic animals.

The vertebrae of the sacrum are more solidly fused together in adults
of deer than in sheep or goats. Sacral vertebra four is often free of the
thii'd, oi' is loosely joined to i1 by a partly opeii suture in the domestic

SKELETONS OF DEEK, SHEEP, AND GOATS

343

THs process extends The neural spnes of
anterior lo the centrum adult deer are reta-
in deer , it usuallu does tivc/y {irmly fused
not e'xtend beyond the
centrum in sheep
and goats ^--....^ " ^

This surface is
broader and flatter
dc(

Vie ironsi/erse proc-
esses extend ventral
to the centrum in
deer.

The '^uentral surface
of the centrurn is
re/aiiuely conuex
in sheep and qoats.

The concauity of the
L/entral profile is not
useful for identification

FIGURE 8. Sacrum, shown in dorsal, right lateral, and cross-sectionol views.

animals. This articulation has i'lisfd in all deer old enough to have a
closed suture between the second and third sacral vertebrae.

Neural spines are also more coalesced in deer than in the other ani-
mals. If the spines are joined as solidly as those figured for deer, the
bone is certainly from that animal; if they are as independent (or
more so) than shown for goat, then the specimen will be sheep or goat ;
if the degree of fusion of the spines is intermediate, identification
should not be made on tliis basis.

Sacral vertebrae two and three tend to have thrii- transverse processes
fused into a broader and flatter surface in deer than in the other
animals. The sacra figured are typical, but many specimens will be in-
termediate between those shown and not assignable by this character.

The bone is shorter in relation to its breadtli in deer and goats than
in sheep. However, caution is necessary in using this proportion because
one or both ilio-sacral joints may be atypical, thus altering the width
measure, and individual variations is marked. The last of the four
vertebrae which usually make up the sacrum is often free of the others.

and is cxc-ludcd from the measure of length. Katio ■ ileer : .V ^ H,

31 = 0.93, a = 0.08 ; sheep : N — 6. M = 0.83, a = 0.03 ; goats : X = 7.
M = 0.95, a = 0.05.

Wide individual variation was noted in the concavity of the ventral
profile of the bone, so this is not a critiM-ifui for species identification.

•.iU

CALIFORNIA FISn AND GAME

Jh/s surface is notched
in deer but flat or
elei/ated m other
anim a/s.

The neural spine extends
posterior to the center
of the ZL/gopophysIs in
deer but not in sheep

Deer

At this level the. neural
spine of sheep is usuallu
thicker than smm. ; that
of deer is usually thiii-
ner than 201 rn.

The area between the
articulatory surfaces is
flat in sheep and goats
but concave in cker.

me transverse canal of
deer tunnels into the
interi/ertebral foramen.

The transverse concJ^
may be absent in
any of the animals.

The transverse canal
of sheep and goats
stops short of the
intervertebral foramen.

FIGURE 9. Left: ventral surface of atlas vertebra. Right: axis vertebra shown in
right lateral and posterior aspects.

ATLAS
(Figure 9)
Only one character of the atlas vertebra serves to distinguish deer
from the other animals. When the bone is seen in either ventral or
anterior view, a distinct noteli or concavity is seen between the anterior
articulatory surfaces in all deer. In sheep and goats these surfaces
crowd closer together and are separated by a flat area or, more often,
by a distinct elevation. A notch also separates the posterior articula-
tory surfaces of the bone iti dcei' ; in the domestic animals this area
is fiat.

AXIS

(Figure 9)

With few exceptions the axis vertebra can be used to discriminate
deer from sheep. The goat falls between the other animals in most fea-
tures and cannot always be assigned.

Deer have the longest neural si)ine (anterior to posterior) ; sheep
have the shortest. In the former, the spine extends posterior to the level
of the centers of the zygopopiiyses, while in sheep it does not reach
so far.

l)i\i(iing the length of the neural spine by the breadth across the

zygopophyses provides a useful pi-npoi-tion. Katio ^; deer: N = 8,

8, M = 1.06, (7

0.07 ; goats : N = 6,

M = 1.88, a = 0.26 ; sheep : X
M = 1.37, a- = 0.48.

The neural spine of deei- is also thinnei- and more blade-like. At its
center it is usually not over one mm. in thickness (but 4i mm. in one

skeijKtons of dekr, sn hkp. and ooats :Uo

old liucki. 'rill' spine of sliccp is iisu;ill\- iiion' tiiaii 4 mm. thick, and
in my saiiiitic never less than '.i iinu. ; goals are intermediate but ap-
])r()aeli deer closer than slie(>]i.

The transverse canal was alisi'ol in luilf of my sheep skeletons, a third
of the fi'oat bones, ami was wanting on one side of one deer axis. AVIien
present it otfers a reliable means ol" setting deer a|)ai-t from the do-
mestic animals, foi- in deer the canal runs forward to join the inter-
vertebral foramen, while it terminates short of this foramen in the
other animals.

SUMMARY

To enforce game laws, it is sometimes necessary to determine if
eei-tain bones at hand conld be from a deer. The parts of the skeleton
likely to be used as evidence of illegal hunting are those retained in a
dressed carcass. Sheep and goats are most often related to deei- in this
way, and archaeologists sonu'times have occasion to distinguish sheep
from goats using only skeletal fragments. This paper deserihes diag-
nt)stie features of 12 bones from limbs, girdles, and spine which nearly
always permit certain discrimination of deer from these domestic
animals, and often set sheep apart from goats.

(Mear-eut criteria for identification are best. Thus a large foramen is
found in the metatarsal bone of deer which is never present on bones of
the othei- animals. Such characters are welcome but few; differences in
relative size or proportions must usually suffice. Compared to sheep,
deer have a more slentler femur, a thicker olecranon process, and deeper
trochlea. Measurements are more concise than adjectives in, describing
differences of this sort, and ratios are preferable to linear measures,
because they are not vitiated in exceptionally large or small animals.
Individual variation is striking and many species dift'erences can be
expressed only as variations in the average form of characters for which
extreme values overlap. Statistical analysis is therefore desirable to
establish the reliability of determinations. By calculating the signifi-
cance of the product of several independent probabilities it is often
possible to combine relatively small odds to establish reasonably certain
identifications. The method is described.

Deer can nearly always be distinguished from the domestic aiiimals
by the scapula, humerus, radius, and metatarsal; the axis, atlas, sacrum,
and ulna are usually satisfactory. The femur and tibia are least dis-
tinctive. Goats are intermediate between the other animals in most
skeletal characteristics, but differ markedly from deer in features of
the metacarpal. Sheep are best set a])art from goats by characters of
the metacarpal, scapula, pelvis, and ulna.

REFERENCES
Fisher, Konalil A.vIiikt

1936. Statistical methods for resc.'irdi workers, (ilh cil., rev. mid ml. IMiiilmrKl:,
Oliver niul P.oyd. :!.''.n p.

Glass, B. P.

1951. A ke.v to the skulls of Xortli AiniM-icaii iii;uiim:ils. Minneapolis, Burgess
Publ." Co., 54 p.

Kendall, Maurice George

1948. The advanced thenry of statistics. London, Ciiarles Griffin and Co.. T,td.,
vol. 2, 5131 p.

;U() CALIFORNIA FISH AND GAME

Lawrence, B.Trbara

1951. Post-cranial skeletal cliMrnctcrs of deer, pronffhorri, and sheep-goat, witli
notes on Bos and Bison. I*t. 2. Harvard T'niv., Peabody Miis. Aiikt.
Archaeol. and Etlmol. Pajiers, vol. 35, no. 3, p. 0-43.

IVarson, Egon Sliarpi', .nnl II. O. Hartley (eds.)

inr)4. Biometrika tallies for statisticians. London, CainlM-idjre T^ni\. Press, vol.
1, 2.3S p.
Simpson, fJeorge (laylord, and Anno line

193!). Qnantitative zoology. N. Y., McGraw-Hill Book Co., Inc., 414 p.

NOTES

WHITE STURGEON WITH SEVEN ROWS OF SCUTES

The primary (liH<iiiostic I'liaracter generally {^[hcii lor the r;iiiiily
Acipeiiscridae is tlie five rows of bony jilates (scutes, siiieids) eovei-iiip:
the body: one dorsal, two latei-als. two veiitrals. As a rule numerous
irregularly placed stellated pi-jcklcs occur between the I'ows of scutes
but are of a diflPerent shape and also, because of their much smaller
size, are rather inconspicuous.

In the course of taji<i-in<;' of white stui'i^con (Aciix iis( r Innisinon-
tauns) in the Columbia River at Prescott, Oregon, duriiiL; the winter
of 1950 and later, a few individuals of this species which possessed
not five, but seven, rows of scutes were found. The additional rows
occurred on each side between the dorsal and lateral rows (Figure 1).
In some individuals the additional scutes were about the same size as
the lateral ones and in a few others were somewhat smaller. In the
Columbia River specimens the number of scutes in the additional
rows was between 20 aiid 30 and intermediate between the number of
dorsal and lateral scutes.

At least tw^o dozen such unusual specimens were secured by Mr.
C. E. Graham, Prescott, Oregon, moorage owner, Dr. ]\I. A. Kenney,
Rainier, Oregon, and the autlior. Mi-. Graham, an old-time sturgeon
fisherman in the Columbia River, stated that lie liad seen seven rows
of scutes on white sturgeon before. The author has examined a large
specimen of white sturgeon with seven rows of perfect scutes caught

FIGURE 1. Dorsol view of a white sturgeon from ttie Columbia River, Oregon, showing the
two extra rows of scutes between the normal dorsal and lateral rows. Photograph by the author.

( 347 )

348 CALIFUKXIA FISH AND (iA.ME

above Bonneville Dam by Mr. John Bronghton, a Colnmbia River
eonnnereial fislu'riiiaii. wlio also stated tliat he had cauLiht sevei-al such
fish before.

Out of approximately 7.000 ^vhite sturgeon examined and tagged in
the Columbia River, about 2") fish or approximately 0.3 pereent
possessed seven rows of scutes. I)ni-iiig the tagging of 1.003 white
sturgeon in the Saeramento-San Joaquin Delta in lU'A by tlie Cali-
fornia Depai-tment of Fish and Game, one fisli with seven i-ows of
well developed scutes was found (letter from J. B. Kimsey, IMarch
15, 1955). AVhether or not such individuals oeeur among the other
representatives of the stui-geon family is not known. — Alexander D.
BajTxOv, 6820 Jenuiuf/s Avenue, Mihvnul-ie, Oreejon, June, 1955.

A RECORD OF AN ATTACK BY A LEOPARD SHARK
{TRIAKIS SEMIFASCIATA GIRARD)

On February 10, 1955, while ex])l()ring the bottom of Trinidad Bay
in Humboldt County, California, a professional diver, John Adams,
was repeatedly attacked by a fish identified as a three-foot leopard
shark. Adams was equi])ped with an A(|ua-TiUng and was moving
slowly along the bottom when he was struck squarely on the left side
of the head by the fish. Just after the first strike Adams noticed that
his nose had been bleeding. Some of the blood was draining into his
mouth from Avhence it Avas being exhaled through the exhaust tube of
the "lung". These blood-tainted bubbles fi-om tlie exhaust were ap-
parently detected by the shark, since it was next observed making a
pass at the rising stream of bubbles, snapping at them as it turned
belly up. A few seconds later a strike, made directly at Adams' face,
was warded off and the fish disappeared.

This incident should serve as a warning to divers that even small
a)ul ordinarily harmless fishes may become dangerous under circum-
stances when the diver assumes to the fish a suggestion of food. —
John W. DeWitt, Humboldt State College, Areata, California, May,
1955.

SECOND KNOWN SHARK ATTACK ON A SWIMMER
IN MONTEREY BAY

On December 7, 1952, a young man .swimming at Pacific Grove,
California, was attacked by a large shark and died as result of his
injuries (Bolin, 1954). A second attack, this fortunately not fatal,
occurred at almost the same spot at about noon on February 6, 1955.
The latter incident so aroused some of the citizens of Pacific Grove
that a $50 reward was offered for the first man-eater {Carcharodon
carcharias), also known as great white shark, to be taken in Monterey
Bay. On ^Mar-ch 0. one month after the second attack. Jack Daniels and
Bill Tomlinson, who were hunting basking sharks off' Moss Landing,
shot and killed a 10^-foot Carcharodon carcharias which was feeding
on a dead basking shark that they had anchored offshore. The fishermen
received the reward when they brought the man-eater to Pacific Grove,
and it was then kiiuUy given to the Hopkins Marine Station for study.

The victim of the second attack, James F. Jacobs of Santa Cruz,
California, received oidv minor scratches to his right foot and shin

KOTES 340

and hniisi's on hotli 1cl;s. IIowcNcr. tlic ri^lit fi'ont ankle I'c^ion of
liis Ijlack mbbcr skin (li\inu snit was torn cxlcnsivcly. and flic foot
of tho frannont luin<^' onl.\ hy a stnfdl piocc of llic lied. 'I'lif left leu- of
Ins snit had two tears. In the attack, the shark removed two wool
soeks fi'oni .Jaeobs' ri^lit foot and ent two holes in the onter wool sock
on the left one. The shai-k tore away the heavy rubbei- swinifin from
the rijiht foot, and Jaeobs, who saw it drop, has the impression that it
was etit into two i)ieees. The left swimfin, which w'as still on his foot
when lie was pnlled fi'om the watei', re('eiv<'d seratelies and deeji
ents from the auimars teeth. The deepest cut, a narrow slash with
parallel marks of tooth serrations on its walls, was about three-eighths
inch deep; a slij>htly curxcd gouge, abotit one-half ineh long, was one-
eighth inch wide and a little over one-eighth inch (lee|). These values
seem small until the resilient nature of gum rnbber is considered.

In attempting to reconstruct the attack from the information given
by -laeobs and studying the left swimfin, it a])pears that the major
sci-atehes and gashes were made only by the lower teeth. The upper
jaw possibly was kept from ocelnding with the lower by the angle of
attack and the bulk of the swimfin. It is appi'oximately 20 inches long,
tapering from three and one-half inches to one-half inch in depth, and
from 4 to 10 inches in width. There are three reinforcing ribs: one on
each vd'^e aiul one in the midline. The deepest liortion of the swim-
fin. if it wei-e ])ushefl far into the angle of the jaws, could possibly
keep the mouth open and the weaker posterior teeth could not cut
through the heavy rubber. The right swimfin was probably cut in two
by the .stronger anterioi- teeth avIkmi the shark released its; initial hold
and allowed tlie swimfin to slide forward in the mouth to the region
of greatest cutting ]>ow(M'.

The striated markings which characterized the cut surfaces of the
swimfin pointed to either Cdvchdrodnn carrhnn'ns or the blue shark,
Frionacc (jlauca, as the probable aggressor. These are the only two
large, local sharks with marked serrations along the tooth margins.
When the marks on the swimfin were compared to the teeth and jaws
of the 10^ -foot man-eater which was landed March 6. the serrated
tooth edges matched closely the striations of the major gashes, and
the cuts were in almost perfect alignment with the teeth of the lower
jaw, the teeth being closer together than the marks, indicating that if
the attacker had been a man-eater it was probably greater than lOJ
feet in length. This tends to coiToborate Jacobs' statement to the
Pacific Grove police that the depth behind the eye was 2^ feet
and the length was 15 to 20 feet. These proportions fit C. car-
cliarios better than P. r/Jnuca, which, according to the figure in
Rigelow and Sehroeder (1948), Avould have to be about 30 feet in
length to achieve the same depth behind the eye. Jacobs further stated
that the shark was blue on top and whitish-gray undtUMu^ath. colors
which better fit the blue shark but are also a color phase (slatt>-blue)
of the man-eater. "When shown a series of plates in "California Sharks
and Rays" (Roedel and Ripley, 1950) Jacobs unhesitatingly identi-
fied C. carchan'as as his attacker. "While P. fiJaucn cannot b«> cleared
completely as the cidprit, the circumstantial evideiu'c points with a
fair degree of certitude to C. carcharias in tliis instance.

350 CALIFUKMA IISII AND GAME

Tlie attack on Jacobs took place as he was swimming in water 20
feet doop, abont ono-qnarter mile from shore, sontheast of the tip
of Point Auloiic (known locally as Lover's Point). A moderate surf
was running, with waves to a height of five feet, the water was clear,
and the day sunny with scattered clouds. Unfortunately, the water
temperature was not taken after the attack; however, the temperature
at the Hopkins ^Marine Station, approximately three-quai'ters of a mile
to the southeast, Avas 52.9 degrees F. at 9 a.m., PST. The temperatures
of the previous week varied between 52.0 degrees F. and 53.6 degrees
F., and these temperatures are about average for the season. During
the preceding week, strong northwesterly winds, not unseasonable
for February, had occurred.

While the attack took place only a short distance from the one in
1952, the physical conditions do not closely a]jproximate those described
by Bolin. Then the temperature was 55.2 degrees F. at the Hopkins
Marine Station, the surf reached a height of eight feet, the water
was murky with phytoplankters and runoff from the land, and the
day was characterized by intermittent sun. It appears that there are
no strong correlative physical factors which could have induced the
attacks, and the evidence seems to point to random movements of a
large predator acting independently of local physical conditions.

A possible common factor in both attacks might be the presence of
blood or blood serum in the water. In the case of the fatal attack,
the victim had been "body surfing" (riding on the waves by the
use of the outstretched body as a surfboard) and coming in over
rocks. George Fraley (a former commercial diver and president of
the "Sea Otters," a" local skin diving club) had talked to the victim
and noted that his body had numerous welts and scratches caused by
this activity. Fraley warned the boy that the presence of blood in
the water might lead to danger from a large predator. Bolin examined
the body of the victim after the attack and found no marks other
than those caused by the shark. However, the cuts of the teeth could
have so mari-ed or overshadowed any breaks in the skin present before
the attack that they would not have been apparent at the time of
examination.

In the case of the second assault, Jacobs had speared a two-foot
cabezon (Scorparnichthyf! marmoratu!^) and had placed his bleeding
catch in a web-lined float. He immediately resumed fishing and scout-
ing for another kill, when the attack took place. This was 10 to 20
minutes after he had speared his fish. He was close to his float Avhen
he felt the strike and pressure on his feet. His first thought was that
a friend was i)laying with him, but, as the pressure increased, he
looked down and saw the shark. He kicked, and the shark left, appar-
ently content with the swimfin and the two socks from his right foot.
Why the shark did not return to pursue the attack has caused some
conjecture. Even tliough Jacobs was soon aided by other swimmers and
removed from the water into a rowboat, there was still time for further
assault. Possibly his complete rubber suit prevented the shark from
sensing any animal odors when it made its attack, the flavor of the
rubber garments being far different from the fish blood it had detected
and followed to the swimmer. The victim of 1952, on the other hand.

NOTES 351

liad been garbed in swiiiiniiiii:- fi-iiiiks find swimfiiis. and tlio seepage
of his scratches could have Ix'cii tlic attractaiit. In this case, tlie shark
returned repeatedly to slash and tear at his prey, i)robahly lured
lime after time by wound seepage. Tt is noteworthy, in this connection,
tluit all the wounds wei-e conceuti-ated on the lower extremities; ii(»
bk'niishes appeared above the waist.

There have been two other instances of large sharks api)roacliiiig
skin (livers who have speared fish in the Monterey area. George Fraley
told me that on two occasions in Xoveniber, ]954, members of the
"►Sea Otters" speartishing off the Hopkins Marine >Station were
approached by sharks. Tliey identified one as the man-eater, C.
carcharias, and the otlier as the salmon m- mackerel shark, Lamva
fUtropis. These men have trained themselves in uiidei-watcr ohscrvntion.
and I believe their observations to be accurate.

From these experiences and the incident mentioned by DeWitt
(1955), it appears that shark attacks pose a real threat to the skin
diving enthusiast. There is strong probability that witli the present
increase of s])earfishin<i- along our coast the number of attacks by
sharks will increase. Those individuals who indulge in this sport
should exercise ever}^ precaution to avoid seepage of blood, fish juices,
or other attraetants.

REFERENCES

Ripelow, Henr.v B., nnd Willinin C. Sohrocdfi-

3048. Fishes of the western North Atlantic. ('h:i|it, :'.. Shnrks. Senrs Fumnl.
Mar. Res., Mem. 1, p. r>!)-r,4G.
Bolin, Rolf L.

in."i4. Report on a fatal attack by a shark. Pac. Sci., vol. 8, no. 1, p. lO-'-lOS.

DeWitt, John W.

1955. A record of an attack l).v a h'op.ird shark (Triakis semifasicain Girard).
Calif. Fish and (J.iiiic vol. 41. no. 4. ji. .•',48.

Roedel, Phil M.

1953. Common ocean fishes of the California coast. Calif. Dept. Fish and Game,
Fish Bull. 91, 184 p.

Roedel, Phil M., and Wm. Ellis Ripley

1950. California sharks and rays. Calif. Div. Fish and Game, Fish Bull. 75, 88 p.

— Thomas A^. Fast, Hopkins Marine Station of Stanford University, Pacific Grove,
California, May, 1955.

REVIEWS

The Complefe Book of Sfriped Bass Fishing

Hy lli'iiry Lyuiaii and I'raiik Wddliici- ; A. S. Jiarncs ami < 'iiin|iaii\ , Xfw ^'ll^k,
1!);"54 ; xii + 242 p.. illuslrali'il willi IN lialf-toiit? and linr iliawiiifjs. $4.

Tins \(ilnni(' is a welcome addiliuii In the t;i-(i\vinj; series on striped l)ass an^'lin;;.
All lioiiuli the jacUet chiinis if is the first t'ull-len;,'tli hook devofed exclusively to this
suh.ject, several ntheis ha\e preceded it, iiichulins oiie on Afhiufic striper tishin;; h\'
Kodnian (11144). an<l another on Pacific Coast sfriped hass fishiiii; h.\ Leon Adams
(li>.-.4i.

Fisliinji tackle of all fyiies for every variety of Atlantic Coast sfrijier an>;Iinj; is
described in defail. (Jood photographs are Sfiici'onsly scattered fhrouKli the text, alonj:
with a series of pen and iak drawinjis illnsf rating: methods of rif;;;inK tackle and
usinjc various baits.

The book is well written and well ai'ranK<'d. It reflects the authors' strong' interest
in striped bass anslin<r in many jileasaut ways.

'I'he many details about specialized Atlantic Coast anfrlinR will he of only passin;;
interest to most I'aoific Coast anj;lers.

It is unfortunate, in some respects, that the authors ha\e nivcu so little attention
to the bass tliemselves .-ind to the various problems of striper management and con-
servation. Many of their readers would probably have welcomed nioi-e information
aloii^ those lines. — Alex ('nllioun, Culifoniia Depart luciit of Fish mnl (limn .

Homemade fishing

By Verne E. Davison; The Stackpole Co., Harrisburf?, Pa., 105.''.; xvii + 20.'. ]>..
illus. $4.50.

The author has been closely connected with small tisli pond research in his 12-ye.ar
association witli the ()klahoma Fish and <!ame Commission. ;ind more recently in
ills capacity of regional biologist with the Soil Conservation Service in the South-
east. In this area he fell under the influence of Prof. Homer S. Swingle and Dr.
E. V. Smith, in their application of agricultural methods to arpiiculfure. His ])ond
management principles stem from the findings at the Alabanui Polytechnic Institute.

Although somewhat oi.timistic in his st;ind that anyoii(> can create "the world's
best fishing pond" in his own backyard through lu-oper management, .Mr. D.ivisou
does impart some practical advice regarding pond site selection, water supply, initial
stocking, the ensuing population composition changes, the fishing, and physical and
chemical (through fertilization) m.anagement of the water. Basic laud agricultural
)irinciiiles. as related to tlie hydrobiology of ])on<l fish production, are clearly ex-
plained. Pond owners are eucour.-iged to fish their jionds hea\ily. discourage rooted
atpmtic plants, fertilize with miner.-il comiiounds, juul use some control over the
liopidation composition of their i)ond. In this latter instance tlie author wi.iild
sanction the use of seines and derris to control this crop.

The combination of l;irgemoufh black bass and bluegill is designated as the only
satisfactory one for small inuids. with words of warning against the use of cr.-ippie
and bullheads. We know now that oflnn" comi)inations with forage minnows li.-ive
been successfully tried. The writer does discuss, briefly, ciddwater jionds stocked with
rainbow trout. Brackishwater or saltwater pond culture is discouraged.

I think that many of Mr. Davi.son's reconunendations to the sin.-ill itond owner
are exemplary, and some could be api)lie(l to other w;irmwafer fish nuinagemeTif
practices. This is a book that will start m.iuy of us dre;iniing of a '"tishin' hole" of
our own. — /'. 1. Ddiiiihis, Ciilifiirnid Ihiinrtniriit of Fish niid (liiiiic.

Ti'ap a and lis Culture

I'.y Pierre Chimifs. /".• FAO Fisheries P.ulletin. vol. S, no. 1, .lan.-M.ir., I'.l."..". ;
11. ii. 1-.".;!. illus

Tilapia is a rel;iti\c lu'wcomiT ii> the tiild of \\>U cultur.-il liter.-iture. Even in the
Far East, whei-e jiond culture h.is existed for thous.-inds of ye.-irs, this genus li;is been

r

354 CALIFORNIA FISH AND GAME

used only since .iImhii (lie tinic of AVorld War II. Its tolerance of high temperatures
and relatively IukIi levels of salinity, and its great adaptability as a fish for pond
culture, has caused its serious coiisiderat imi for introduction into problem waters in
this country and .-liirnad. Only three of the api)ro\imately '.»l known species liave
been studied extensively from the tish cultural point of view.

This annotated bibliography, with its !)4 citations from the world literature, is the
most complete that this writer has seen. It has extensive comments under three
general headings in luldition to the bibliography: I. Tilapia in Open Waters, II.
Biology of the Triucipal Species Raised in Fish Culture, and ITT. Methods of Rais-
ing Tilapia.

The members of the Tilapia group are warmwater fishes that do l)est when tem-
peratures range near 90 degrees F. They spawn at about 70 degrees F. and die if
temperatures fall below about 50 degrees F. They are well suited for the tropics and
since much of this area is in perpetual need of more protein we are destined to hear
more and more of Tilapia culture. Their use in the United States will be restricted
largely by their temperature requirements and sporting finalities. We have not yet
reached the point where extensive pond fish industries are necessary to provide
Ijrotein.

This interesting and timely paper is recommended to anyone interested in finding
out more about Tilapia. A useful note by the editor of the FAO Fisheries Bulletin
aids the reader further by directing him to the world's principal libraries and docu-
mentation centers w^here the papers listed in the bibliography may be obtained. — ■
J. B. Kimsey, Califortiia Departtiient of Fish and Game.

Streams, Lakes, Ponds

By Kol)ert L. Coker ; University of North ("arolinn Press, Chapel Hill, North
"Carolina, 1954 ; xviii + 327 p., illus. $G.

This is a book for which many whose major interest is in the management of our
surface water resources have waited. It brings together in one concise volume a
wealth of information concerning the complex picture of life existing in our streams,
lakes, and ponds. The author has reflected his wide personal knowledge of and ex-
perience with the subject and has portrayed it in an interesting and readable form.

The book is written in three parts in a natural sequence — Water and Its Content,
Running Water, and Still Water — each of which is a separate entity, but which
together form a well coordinated and concise story of life in fresh water (as opposed
to life in the .sea).

The chapter on stream pollution is very interesting and points up the importance
of keeping our surface waters clean, i.e., "well oxygenated, free from silt — untainted
by germs of disease of man or animals or by substances toxic to man, beast, fishes
or the living food of fishes."

The book is neither a textbook for students nor a reference for specialists, but is
a pleasantly readable and comprehensive review of life in inland waters, recom-
mended to those engaged in fisheries management, as well as to instructors in the
field of general zoology, sanitarians, and fish culturists. — Harry A. Hanson, Califor-
nia Department of Fish and (ianie.

The Disfrlbution and Abundance of Animals

V,y H. (J. Andrewartha and L. C. Birch; University of Chicago Press, Chicago,
1954 ; XV -f 782 p., illus. $15.

Here is a textbook of ecology which asks the fundamental questions: AVhy does
an animal inhabit so much and no more of the earth? Why is it abundant in sonu'
parts of its distribution and rare in others? Why is it sometimes abundant and
sometimes rare? In seeking answers to these questions the authors critically analyze
the ways in which the environment influences an animal's chance to survive and
nmltii)ly.

Both .-luthors are Australian workers, are experienced in their field, and have
accumulated and coiniiiled. in very orderly fashion, an enormous volume of fact
and theory pertinent to the field of animal ecology. They h;ive intentionally organized
their material in an unorthodox though logical manner. Rather than emphasize the
traditional study of animal communities, the authors place their emphasis on species
l)oi)ulations and the actions of the environment on these populations. An attempt is
made to .set up a basic pattern of inciuiry which should l)e applicable in the study of
any species.

HEVIKWS ■'^')i')

The book is (li\ifl<'(l into five k<'Ii<'>'''i1 topics ciicli coiiljiiniii;; iiiirncroiis siilxlivisioiis.
Tlie introductory \)nvt includes discussions wiiicii cxiu-css some of liu' ;iutliors' ol)jfc-

tions (o certiiin "jn-inciiilcs" ol' cc(do;;.v wliicii liicy t'c<d li.-nc 1 n Jicccptcd dou-

ni:itic;ill,v (in the l))istl. 'I'lie next two pjirts tiike up tlie piiysi(do^ic;Ll jispects of
ecidii;;y ;ind an aiiiilysis of tlie four hasic coniiioiu'uls of tin- enxironiuenl (wealluT,
food. otiuT animals, and a place in which to live). The fourth part of the liook is
entitled "The Numbers of Aninuils in .Natural Populations." Here is a series of well
documented, long-term population studies is analyzed according; lo the lufthods and
reasonint: outlined in the first three parts of the book. The last portion of the volume
is devoted to the frenetic aspects of ecology and their implication in e\olution.

The entire book is well illustrated and amply documented. < "onsidei-ing the complex
nature of the material, it is written in a very clear and easily followed manner. The
authors have avoided the use of p(uiderous. complicated definitions which they feel
have plagued so many works on animal ecology iu the past. However, this is not a
simple, easily read book — to be lightly scanned at leisure. The very miture of the
material precludes this possibility. It can i)r(d)al)ly be most effectively used in group
or seminar discussion. To obtain the most from the book recpiires considerai>le
thought and concentration and a real desire to learn.- Ihniil ('. ■ftintiih. <'alifonii)i
Dcjxtrimf'ni of Fish and Game.

The Wesfern End of Lake Erie and Its Ecology

l\\ Thomas 11. I.anglois; .1. W. Ivlwards, I'ulilisher, Inc., Ann .Vrbor, .Mirhig;in,
l!>r)4; XX + 47'J p. ; 72 figs. $10.

This book, as the title indicates, is a very composite study of the interacting forces
of nature in a l.^tiri-sqiuire-mile "key area" of Lake Erie. Its aims are to supi>ly the
basis for a fishery nuuiagement program for Ohio's part of the lake. I feel very
humble in reviewing a work of this magnitude, which covers siune Kttl ye.-irs of data
collections and 60 years of intensive study by the author.

It is extremely gratifying to see research starting with the basic factors affecting
the welfare of the individual organisms and working up to the endpoint of the evalu-
ation of the commercial and sjiort lisliery. By the understanding of why the biota at
the unicellular end of the food chain tluctuates, we can more easily underst:.nd the
resultant fish population dynamics. The author presents aderpiate evidence, by inclu-
sion of many personal observations and data on the many phases of ecology collected
by other workers on Lake Erie, with which to formulate a s^ound fisheries manage-
ment program. Some of the topics covered are climatology, bottom studies, pliysic.-il
and chemical character of lake water, origins of turbidifying materi:ils. bac(eri:i.
algae, aquatic phanerogams, protozoa, annelids and other worms, mollusks, crusta
ceans, aquatic insects, reptiles and amphibians, fishes, birds, and mammals. The fish
life of the Lake Erie drainage includes .some lU species. Notations are made on col-
lections and life histories of many.

I'nder fish management methods six major headings are covered : regulated
cropping, artificial propagation, iiollution control, facilitation of migration of anad-
ronious and catadromous fishes in tributary streams, improvement of land use prac-
tices, and research.

In addition to the very "meaty" material contained, some 4^0 references are cited,
making an excellent bibliography. This book shoidd be in the liitrary of all warm-
water fishery research workers. — /'. .1. I>(,ii(jjas, Cnlifoniin nrimiliHriit of Fish niul
Gil me.

Water; A Study of Its Properties, Its Constitution, Its Circulation on the Earth,
and Its Utilization by Man

By Cyril !S. Fox; The rhilosoi)liical Liiu-ary. liu-., .\ew Y-.rk. 1'.>.".L': xxvii + 14.**
p., illus. with 4 text figs, and '2'> pis. $8. To.

This book is an attempt to gather together a considerable amount of information
on the subject of water. The scope of the book is not as broad as the title suggests
and the principal emphasis is almost entirely on geology, rather than on the engi-
neering or social aspects of the world's w:iter iiroblems. This undoubtedly reflects the
author's background as Director of the tJeological Survey of India and tends to
reduce the usefulness of the book for readers who are interested in a more broad ami
basic reference book on the subject.

This book contains a c<insiderable amount of numerical dat;i on water. i>arti<MiIarl\
on rei)resentative surface and underground waters. These analyses are interesting
but lengthy and there seems to be a rather inordinate amount of repetition that could

356 CALIFORNIA FISH AND GAME

Lave been eliminated without loss. For ex;uni)]e. on i).i;;c .">, it states that sea water
contains 'A~> percent salts and the same infoiinatinn can he found six otlier times in
tlie liook. .More interest would lie ^'enerated l)y liavinf; less empliasis on niinilicrs
and more cniidiasis on topics of ;,'iMnT;il intcii'st, siicji as ii'ri;,'at ion clc\ i']o|inii'iil ami
mctlMxls of water utilization.

Tile hook is short — only 14S ])a};<'W. The rc\ lew copy suffered hadly as a result of
rather poor printing and six jjajres were blank. It is lioiied that this mechanical fault
was corrected in the press run. I'nfortunately the editing; that is needed to make this
a valuable, recommended refi'rence or textlxxdj will apparently have to wait for a
new edition. — A*. .1/. I'liul, Vulijurnitt Depart nwnt oj Fhh and Gaiiie.

Analysis of Development

Edited by lienjamin II. Wiilier. ran! A. AN'ciss. and Viktor IIanil)Ui ;:er ; W. 1',.
Saunders Company, Philadelphia. ]U7,~, ; \\\ -\- 'J'.i', p., 24.S text liss. .$!">.

One may presume that analysis is the ohjecl of a book entitled Aiialijyi^ of Dervl-
opment. This is the case, but the introductory statement by the editors stresses
synthesis. Although these terms are antithetical, the book approaches the suh.iect by
both methods. This is a presentation of the analysis of growth, from i)roduct'on (»f
gametes, their union, and subsequent differentiation to completion of the organism
prior to independent living ; and this l)ook is also a synthesis of work in the field of
embryology, composing the contributions made by former workers and workers of
today.

Twenty-nine pajiers by twenty-eight authors, guided by ;in editor-sponsored out
line, have resulted in understandable style differences and occasional loss of unity,
which are not apt to be found in a book by a single author. The editors noted this
and also pointed out the impossibility of a single author writing such a book for the
field of embryology. This book is a .synthesis of knowledge of the principles and
mechanisms of development, as the authors state ; clearly it is also an analysis of the
develoi)ment of living organisms. The science of embryology is diverging, nvn must
specialize, the labor must he divided. The terminology, the language of the field,
grows hut the oiijiortunity for broad principles diminishes. The true synthesis seems
remote.

An interesting chapter on methods and techniques poses problems that might well
be consideied in other fields. The problem of technique of investigation in embryology
is one of observation or interference ; both require the determination of natural
environment. Interference with the developing embryo makes results suspect. But
observation is affected by prec'onceplions. Normal environment is difHcult to deter-
mine, since many organisms have wide ranges in environmiiii : the emhryo also has
an internal as well as external environment.

Each author is a S])ecialist and leader in his field. <^nly fellow workers in the
author's special investigation could aibMiuatsdy criticize lis contributory paper. The
reviewer suggests, howevei-, that a common fault s;'ems to be an absence of i)road
interpretation and neglect of summation. There are exceptions to this, a notable one
being the paper on teratogenesis. A i)ai);'r by Weiss concludes with a thoughtful dis-
cussion of summary in developmental proces.ses. He believes summary is not possible,
for we must "clai-ify complex mechanisms of their operation" as well as "establish
and reconfirm the \;ilidity of general laws and princ'))les." With this it is difhcult to
argue; nevertheless, summarization is a standard practice in most journals and a
very valuable custom it is. Technical i);;pers are addressed to small .-indiences iuit
there are workers in tields hoih allied .-nid ili\erse who i]|-olii through the niediuin
of summary.

The endiryologists who conl rihuleil to this hook are working at ihe limits of knowl-
edge in their respective tiidds ; future devtdopment. following their guidance, will be
expressed in applied science. There has stddom been an exceiition to this in the iiast.
This book m.ay serve as a reference work on emhryological research for workers in
ai)plied fitdds. To these people it will indicate the direction specialists are now taking
toward understanding embryogenesis. The mi'thods, demonstrations, and interpreta-
tions may providi' stimulus to those confronted with i>rohlems in production ;ind
assist design and interjiretat ion of their experiments. The hook adds to the knowledge
of the ways of living organisms.

To read this hook is to be imi)ressed once ag.-iin that this is the fundamental branch
of the science of living organisms. Research in Ibis tiidd (d)serves development .-it its
Iirimary le\-el and interrelations in their most complex state. It is not strange that
therefore new problems multiply wiili each explanation.

KI'.VIKWS :!.)<

.\ii<ili/sis of l)ii<hi)nii< III ciiNcrs I hi' I'nlliiwin;; ticlils ;iiii| llii- wuikcrs ciiiit rilnilt-
li.'ipcis williiii till' rriiliii'Wiirk nt" tlii'si' si'ctinii liriiiliiifrs ;

I I'rulilrins, Coiiccpts jiiid Tlii-ir IlisloTy

II Mi'tlinds and Ti'chiiiinics

III ('I'lliil.ii- Structure and A<'tivit\-

I \' Till' Xiiclous and Cytnplasin in I )('\ I'lupnu'iil

\' i;nilir\ n;;i'ni'sis : I'roparat ion I'liasi's

\' I Miiilii'viiLcrni'sis : I'ri)i;ri'ssivi' 1 )ilTi'ri'nt i;i t ion

\ 11 Spi'cial N'rrti'hratc < >r};aiio};(Mi('sis

\ 111 l']n('r;;.v I'L\clianf;>' and Enz.vnio Dcvclopnu'Ml KiiriiiK i;ndpivoi;''ni'sis

IX Ontofjcny of Ininiuiiolo};ical Properties

X ( )nto{,'iMi,v of Kndocrine Corrrlat ion

XI I >i'ti'rniination of Size

X I I Mi'iainorpliosis

XIII Ki'j;i'neratiou

X I \ Teratogenesis

This bonk is a wise addition to an iiist il ii( ion lihi-aiy. Allhoui;h u'l'm-ral workers in
the hioloKical scionecs will have only oeasional nse for it. enihryolojiists will find it
essential to their woi-k.

Advances in exjuTinient and thouKhl are rajiid and new summaries in the field of
einhryolofiy undouhteilly will appear, altlmujjh probably some time will elapse before
another book of such scope and importance is puldished. — Roheit J\. Hell, California
hrparinicnt (if Fish and Game.

introducing Sea Shells

r.y K. Tucker Abbott; I). Van Xostrand Co.. Ini-., Xew York, lit.V. ; vi + G4 p.,
19 line drawings. 10 plates (6 in cnhn- im-luding covers). $2.50

Popularity of shell collecting, a fast -growing Anieric:tn hol)by. is the first subject
of discussion in this little booklet. Thence. i;ne is given an insight into the ri>m:intic
l>asts of some of the world's most priceless shells. Succeeding chapters are entitled
"Hunting your own shells." "Exploring our color plates." "Preparing shells for the
collection," "Strange seashells", and "A collector's list of .shell names, dealers, and
clubs."

Perhaiis the most v.ihiable information in the entire (14 pages is that containnl in
the chapter on preparing shells for the collection, a subject the importance of which
cannot be too highly stressed. Similarly the li.sts of shell clubs and dealers should
prove useful to those hobbyists who currently lack this knowledge.

A number of undesirable items and jissorted inaccuracies tend to limit the value
of the book — especially for the beginning collector who would be unable to sepanite
the chaff from the wheat. A true conservationist would be horrified to find that the
author has advocated, in at least four places, turning over stones and boulders in
search of shells without once informing the reader of the need to replace these
natural communities original side up. On page 23 one is led to believe that in
California only the red abalone is protected by law— whereas eight .species 'of .•ib:i-
lone-s found within the borders of the State are currently regulated (from the sports-
man's point of view) by such methods as: a single aggregate bag limit, four size
limits depending upon si)ecies, a closed sea.son, the need for a valid fishing license,
and so on. In addition, these laws are subject to change from year to year as the
need for greater or less protection is determined. The author failed to warn the
amateur collector of a basic rule of thumb which should always be applied i)rior
to field activities in any state, namely: check the latest fish and g;ime laws of the
state in question.

It is obvious from his statements on pai^r 1'.) that the author is neither a skin
diver himself nor is he well versed in the art ot" skin diving. It is doubtful if more
than 1 percent of the tens of thous.-inils of enthusiastic skin divers in the Tnited
States are physically cai)able of remaining submerged for more th.-m one niinute
l)er dive, let alone "two — or even three".

A number of other errors and inconsistencies are .scattered through the book. Even
if one overlooked them, the paper covers, very large type, and Imited information
make the asking price for this booklet niucli too hi-h. John E. Fitih. C'nlifoinlti
Department of Fish ami (lauiv.

358 CAL7FT)RXIA FISH AND GAME

North American Moose

liy Riiiidolpli L. I't'tei'snn ; riiixorsit\- <<( 'rnmnto I'ross, '^I'ofoiito, litrirt ; xi + 280

p., 4;! iihotns. .'i;i2.r>n.

The existing liirr.ituic nii inniisr is rdinhiiicil with Dr. Peterson's persDiiiil work
in Ontiirio to coiniiusi' tllis \oliniii'. I»;it;i frum tin' Sc;ili(liii;i\i;ni couiitrifs arc used
freely for eomparative iniritoses.

Current knowledge of moose, including taxonomy world-wide, from i)rehisforie
times to the present is presented. Chapters are devoted to reijroduetion rate; general
life history ; behavior ; food habits ; habitat ecology ; relations with other animals ;
diseases, parasites, and insect pests; accidents; population studies; management;
taxonomy ; and general history.

Population data show that moose ha\-e decreased in Xew England and the western
fJreat Lakes states in recent years, while encouraging increases have taken place
in the western state-s, Canada, and Alaska.

In general, moose have a low reproductive rate, but reasons for it are poorly under-
stood. The basic cause is probably a lack of variety of good foods. White-tailed
deer competition has kept moose num])ers down in many areas. Populations reach
maximums in early successional stages and decrease as forests reach maturity.

Two appendices are included. One. "A Study of INIandibular Tooth-Wear as an
Index to Age of Moose", by R. C. Passmore, R. L. Peterson, and A. T. Cringan,
presents age classes derived from grouping jawbones of similar wear. The lack of
known-age material is clearly i)ointed out. This aging data should be useful to
technicians.

The other appendix, "Studies of Moose Antler Development in Relation to Age",
by A. T. Cringan, presents little that can be put to practical use.

Animals existing in low-density populations, such as moose, compound the normal
difficulties of obtaining size and age composition of herds, rate of annual production,
and range carrying capacity. To a person familiar with current research efforts on
such abundant big game as deer, the generall.v small samples referred to and the
lack of much basic data in this book will be glaring. However, the work that has
been done on moose is amalgamated, and profitable avenues for future investigation
are clearly pointed out. — Fred L. Jones, California Department of Fish and (lame.

M Is for Moose

Bv Anton Swanson ; Vantage Press, Inc., New York, 1954; 64 p., 35 illustrations.
■$2.50.

The author is a contractor whose lifelong hobby has been the study of wildlife,
his chief interest being moose. The author's experience has been mainly with moose
in Sweden, where he spent his youth, and in Ontario, where he has lived for over
30 years.

This short book covers briefly the main phases of life history and management and
even contains information on souvenirs which can be made from moose antlers.
Of special interest to the reviewer was the section on the moose and its management
in Sweden.

The pi-ofessional wildlife workci- will find this book lacking in detail and will feel
that much has been left out in regard to life history and management. The layman
who may have :i genei-.-il interest in the species will find this book an easily read,
quick source of iMfortnation on the subject. — C. M. Ferrel, Galifoniin Drpartnirnt
of Fish and Gaiiic.

INDEX TO VOLUME 41

.\riiirti!<< r t ni iisiiidii t<l ii lis, .'(47-348

AcipciisfridMc, ."MT-.'WS

Adennxtoniii jnsriruUiiutn, .jT-TS

AIf:ilf;i. 57-78

AiisHiij; : ^cc fishins. sport

Artriiiixiii tridcntata, 57-78

Atheriiiops : affinis. 279-284; affinis

affinis, 270-284
Attack ; shark, leopard, 348 ; shark,

whito. 348-351

B

It.ijkiiv . Ali'xaiulcr I >. ; ^\'hit(' stur«poii
with seven rows of scutes, .">47-.">4S

r.arT-aeiula ; ("alifornia, 1(')7-17<>

Hass ; larftcinouth black, aii;;liiif;- esti-
mates. lt)-.32: striped, angling
estimates, 19-:!2

f'.at!i\ tli('riiin<;rai>h protile, 79-90

I'.erry, S. Stillmaii ; On recent Cali-
fornian occurrences of the rare
octopod Octjthoe. 177-181; The
male flapjack devilfish. 219-224

Rigeye tuna ; see tuna

liighorii sheei). 131-14.",

I'.issell, Harold I)., Bruce Harris, Helen
Strong, and Frank .James ; The
digestibility of certain natural
and artificial foods e.aten by deer
in ("alifornia, 57-78

Bissell, Harold I)., and Helen Strong ;
The crude protein variations in
the browse diet of ('alifornia
deer, 14.5-155

P.itterl.rush, 57-78

Jiroad, Robert I).; see (langniariv and
Broad

C.-ihezon. .348-.351

Capra ; falconerl, 327-340 ; hircn. .327-.34G
Carchnrodon cnrcharias, .348-351
C'arpelan, I^ars II.; Toler.'ince of the
San Francisco lopsm«>It, Atlir-
rinops affinis affinis. to condi-
tions ill salt-]>rodn(ing jionds
bordering San Fr.ancisco Bay,
279-2S4
Catch records: catHsh. while, L'lil 209;
freshw.-iter ;mgliiig estimates for
1953, 19-:!2; sardines, 2S5 293 ;
tuna, yellowHii. HI ".>s ; lima. i)ig-
eye, 91-!t8
('.illish : angling estimates. I'.)l'>2; white,
2<;i 2t;'.i

( 'il liisliiiil IIS I iiii iriiliis. lis ll'.l

Census; I'ismi. cl.-iiii. 2(i'.l 21 1

Census methods; deer, 22.5-22.S

< 'rra tiiiii i/.iii shiislii, 18.3-187

("lianiise, r>~ ~S

Chandler, ll.iiiy I'ii.\ la iider ; in me -

riain, 248
Clam, I'ismo; census, 209-211
Cieini'iis, Harold 1',.; Fishes collected in

the ti-opical eastern I'aeilic, 1952-

.53, 1 CM fit!
Cohnnha fiisriatn nionilis, .''.1.5-.">2(>
Columbia River; tagging of white

sturgeon, 347.348
Coot; nesting studies, 2!)5-314
Coots, Millard; The I'acific lamprey,

Entosphcnns triiUntatiis. .above

Copco Dam, Siskiyou County,

California, 118-119 '
Coots, :Mill;ird; see Dill. Coots, .■ind

Dollgl.'lS

Crappie ; angling estimates, 19-.32
Cryptobia iorreli, 183-187

Dasmaiin, Raymond F., and Richard D.
Taber ; A comparison of four
deer census methods, 225-228

D;iugherty, Anita E. ; see Felin, M.-ic-
(Jregor, D.-iugherty. ."ind Miller

Deer; browse diet. 14.5-1.5.5; census
methods. 225-228; Columbian
black-tailed. 225-228; foods. 57-
58; skeletal characteristics, .327-
:!4(i

Demiiig, ( ». v.; Rearing bighorn landis
in captivity. 131-143

Develoiiment ; early, barraciid.-i. 1t)7 17<>

Devilfish, flapjack, 219-224

D.Wiii, .John W. ; A record of an at
lack by a leopard shark (Trinkis
srni ifasciii ta ( ! i i-a rd ) , .348

Diet; deer, .57-7S. 14.5-1.55; opaleye,
20:'.-208

Dill, Willi;iin A. ; resign.ation of, 247

Dill, William A., Millard Coots, and
Philip A. Douglas; The status of
the pumpkinseed, l.ipmni.i flilt-
liosiis (Linne), in California,
157-1.59

Diseases; lambs, bigiiorn, ll'.l 11.".;
liroto/.oan, 1S.'!-1S7

Distribnlioii ; piinipkinseed sniilish, 157-
159

Dorosoinn crprdianiini. ll.''i-ll(i

Dol\. I\ennetli F. ; see Lassen, Doty anil
Sa uceiniaii

( 359 )

;;(■)()

C'AT,IK()RXT.\ FISTT AND (!Ai\IE

I till. Coots :iii(l

I>(Mi;:l;is. riiilip A. : .•

I >oii;;l;is
1 MicUs ; nesting,' studies. 2'.iri-;!14

lOcoiioiiiic v.iliics; sec tisliiiif:, spo
I-'fTfTi^ : .•illoti-iof;iiatli aiul syiiciitoi

fislics, lO.'Mon : l.aiT.-KMKla,

170; expiTiiiH'iilal liatchinj;'.

242
Kl Xino. 79-90

l']mliryol(><;y : han'acuda. 1(>7-I7fi
Entosiilirniis tritlrnidl iix. IIS-IIU
ExocoetuH ; ohtiisirostris, 103-10");

tans, 103-105

rt

irnath
107-

roli-

Fast, Tlioiiias N. ; Second known sliarlc
att:aok on a swininicr in Monterey
Hay, 3-48-351

Feder, Howard M. ; The use of vital
stains in markinp: Pacific Coast
starfish, 245-24<)

Feeding; lambs, l)i;;horn. 131-143

Feeding hal)its ; opaleye. 203-208

Fflin, Frances E., John MacGregor,
Anita E. Daugherty, and Daniel
J. Miller ; Age and length compo-
sition of the sardine catch off
the Pacific Coast of the United
States and ^lexico in 1954-55,
285-293

Fish tagging ; see tagging

I-'ishcs ; eml)ryonic characters. 10.3-105;
south San Francisco P>ay, 271-
278; tropical eastern Pacific.
161-166

Fishing, commer<-ial ; Pacific mackerel,
107-112

Fishing, sport ; .-ingling estimates, 1953,
l!l-.32; economic values, 5-17

Filch, John PI ; Age composition of the
southern California catch of
P.acific mackerel for the 1953-54
season, 107-112; Results of the
l!l.-.4 Pismo clam census, 209-211

Klyiiiglishes, 103-105

Food studies ; subsurface tuna, 99-101

Forage ; deer, 57-78, 145-155

Gangmark, Il.-irold A., and Robert D.
T'road ; Experimental hatching of
king salmon in IMill Creek, a
ti-ibutary of the Sacramento
River, 233-242

German, E. R., and J). A. Lal^'aunce;
A Comment on the us<' of red
tags on fish. 119-120

(lirellu ni(;ric<nix. 20.3-208

Gizzard siiad, 113-116

Goats; skeletal characteristics, 327-340

Grain, 57-78

H

Il.irris. I'.ruce; see I'.issell. llai'i'is.
Strong, and .fames

Her;il(l. I'^arl S.. and l>on;ilcl .\. Simp
son; Fluctuations in aimndance
of certain fish<'s in south San
Francisco Bay as indicated by
sami>ling at a trash screen. 271-
2TS

Ililiiebrand. Milton ; Skeletal differences
between deer, sheep, and goats,
327-340

Honey Lake Valley ; nesting studies of
ducks and coots. 295-314

Hume, Lloyd C. ; Rainbow trout spawn
twice a yeai'. 117

Hunt. K. (i., and A. E. Xaylor ; Nesting
studies of ducks aiul coots in
Honey Lake Valley, 295-314
Hunting areas; waterfowl, 3>3-55
Hydrostatic pressure tests; rainbow
trout, 243-244

I

Ictaliints catiifi, 2(51-269

Iwai, Tamotsu ; The pharyngeal p(»ckets

of the .sardine, Sardinoiis c<tcrulca

(Girard). 113-116

James. Frank ; s(>e liissell. Harris,
Strong. .iikI .lames

.lennings. Feenan ; see Wooster and
.Tennings

.Inlil. Ivolf ; Xotes on the feeding habits
of subsurface yellowfin and big-
eye tunas of the eastern troi)ical
P.-icific Ocean, 99-101

K

Kozlik, Frank M. ; Waterfowl hunting
areas ojjerated by the California
Department of Fish and (!ame.
33-55

L

LaFaunce, D. A.; see German and
LaFaunce

Lassen, Robert W., Kenneth E. Doty,
and Ruel D. Sauccrman ; Sexing
day - old ring - necked pheasant
chicks iiy a color characteristic,
229-2.31

L.inghlin. John; Ki\er otter noted east
of Sierra n crest, 189

lA'puiiiis jiilihoxiiK, 1.57-159

Longlining; tuna, 91-08

Ijoithotes cepedianus, l<l.">-l(l5

Liitra canadensis, 189

INDKX

361

M

.M;ic;ml;iy. Kdw.-inl I,.; i-ct ircnii'iil nf,

IL'I
M.i(( i ri'^or, .lolin : sci' I-'diii. .M;ic

<lri',i;ipr, I »;ni};licrl \ . .-iikI .Miller
.M:ici;rc-iii-. \V;ill;ifc ( ).. ^iinl W;illnii .M .

Smith; Xcstiii;; .-iiid n'lirndiici inn

i>r the Ii:iiul-t;iiltMl ])i^'r((ii in

('.ililurnia. 8ir>-;]12(;
.M.ickcici, Pacific; age ooiniiosition. 107^

1].:
M.I 1 kin-; starfish, 24r)-24()
.Mc( 'amnion, Geoi'sc W. ; see Pelgen and

McCammnn
Mi.mation; catfish, wliitc, 2(Jl-2()'J
Mill Creek; hatching, king salmon, 233-

242
Miller. Daniel .F. ; sec Felin, Mac-

Gregor, Daiighorty, and Miller
Munterey P.ay ; shark attack, 348-351
Mortality ; king salmon eggs, 233-242

N
N.iylnr. A. E. ; see Hunt and Naylor
\eothititniis niacropteriis, 91-98, 99-101
Nesting ; band-tailed pigeon, 315-326
Nesting studies ; ducks and coots, 295-

314

O

Oak, live, 57-78

Oceanographic observations; eastern
tropical Pacific, 79-90

Octnixxl. 177-181

Oci/thoe tuberculula, 177-181

OdocoUeiis ; hemionus, 145-155 ; henii-
onvs coliinihinniis, .57-78, 22.5-228,
327-346; hemioiiux hemionus, 57-
78 ; virginianus, 57-78, 145-155

Onrorhi/nchu.H f.shninjfscha, 2.3.3-242

Opaleye ; feeding habits, 203-208

Opistlioieuthis californUtna, 219-224

Oiton, Grace L. ; Early developmental
stages of the California barra-
cuda, aS p h y r n e n a argent ea
(Girard). 167-170; Separation of
eggs of synentogiiatb and allo-
triognath fishes in ear]\ endir\-
onic stages, 103-105

Otter, river, 189

Oris: dries, 327-346; niniiilriisis, i;;i-
143

racitic lamprey. 1 is 1 111
I'lniithiiinnis sil,i. '.\\ '.IN, ;i;i |(i|

I'elgen, David K. ; Keonumic values of
striped bass, salmon, and sfeel-
head s|>orl fishing in < 'aliforni:i,
5-17

i'eken. David K.. and C, -ge \\ . .Me

('amnion; Second prf)gre.ss report
on th<' lagging of white catfish
i fcliiliinis riiliis) in the S;icr;i
mi'iiloS.-in .foaiinin Delta, 261-
2f>'.»

I'lnisiil II IIS cdIiIi iiiis. 22".I2."1I

Pheasant ; ring neek. 22'.t-2.''.l

Pigeon, hand-tailed, :;i5-:;26

J'istisler orlirfireiis. 245-246

I'listophorn. 18;M,S7

I'nriniiiilophoriis diego, 107-112

Postal card survey; see Skinner. I'.i :;2

Protozoa, 183-187

Punipkinseed sunfish, l.")? 15'.»

I'lirxliiii t riilciitiild . ."iT IN

Q

Querriis trisli;:enii, 57-78

Peproduction ; band-tailed pigeon, 315-
.•;26

Reviews; All about: brook trout, from
Maine to California, 123; The
American angler, 195 ; American
game birds of field and f(uest.
197; Amphibians •■ind reptiles of
western North Americ;i, 192;
Analysis of development, 356-
357 ; Animals of the southwest
deserts, 125; The complete book
of striped ba.ss fishing. ."..").•{; The
comiiosition of scientific words.
2."i.".-254 ; Conservation law and
administration, 255 ; The distri-
bution and abundance of animals,
354-355 ; The duck hunter's
manual. 125; The elementary
chemical composition of marine
organisms. 101 ; Elements of
ecology, 251 ; Family Circle's
guide to trout Hies and how to
tie them. 191-192; Favorite flies
and their histories, 249 ; Fishes
of the western North Atlantic.
I'art two. Sawfishes, guit.irfishes,
skates, rays and chimaeroids,
194; Forestry. 194; Homemade
fishing, ."^'i:; ; Introducing sea
shells, 357; M is for Moose. 35.S ;
The mammal guide, 193 ; Mam-
mals of California and its coastal
waters, lO.'i ; The nafur:il regula-
tion of anim.'il numbers. 2."i2-2.">.'! ;
North American moose. '.\r>S ;
Principles of gener.-il ecidogy.
250-251 ; Sea - birds, 1-23-124 ;
Spinning in America. 249-25(1;
Sleelhead to a fly. 196 197 ; Stray
feathers from :i iiird nnin's desk.
25;!; Streams, lakes, ponds. 354;
Tilapia and its culture. 353-354;
Timber in vour life. 254 ; Water:

362

CALIFORNIA FISH AND GAME

A si ml \ iif il s ]ii'i]]irrl ii's, its
coiisf it 111 inn. ils ciicuhil idii (111
llic (■.-irlli. ,■111(1 ils III ili-/.:i1 inn liy
liiMii. .■'>.■!."> I ;."•(> : Till' w.iy In liillr
;llli;lili^'. I'.iri; The wi'sl cili cliil nl
I,;ik(' P'ric ;iii<l ils rcnln^.v. .">.>.>;
Wli\ iinl siirvixi' V. '2T)4-2~>~) :
Wilillilr iii.iiiiiui'iiK'iit. Miliinii' II :
Fur lit'iirtTs, \\;i tcirnwl .iiul tisli.
124

Kihhniifishes, 103-105

Knwlcy. William E., Jr.; llydmstat ic
prcssui-o tests on rainlmw trniil,

Unsscll, Kiudlay E. ; Multiple caudal
spines of the round stingray,
Urohafis JuiUrri, 218-217

.Sacramentu-San Joaquin Delta ; tagging

of white catfish, 261-269 ; tagging

of white stursenn. 847-348
Sage, big, 57-78
Sagebrush, 57-78

.Salinity; eastern tmidcal Pacific, 7'.)-'.l(l
Salmo gairdneii. 117, 243-244
Salmon; angling estimates, 19-32; king,

cxperiment.-il hatching, 233-242;

sport fishing values, 5-17
Salt ponds; San Francisco topsmelt,

279-284
Sampling; south San Francisco Bay

fishes, 271-278; topsmelt, 279-284
Sai-diiies ; catch cnniposition 1954-55,

2S5-29:! ; pliarviigi'al pockets, 113-

116
i^didinops caenilca, 113 11<>. 2S5-293
Saucerman, Ruel D. ; see Lassen. Doty,

and Saucerman
XcorpdCH ich fh .(/.v in <i riii o i<i I ks, 348-351
Sexing; pheas.ant chicks, 229-231
Shark ; attack, 348 ; 348-351
Sheep ; skeletal characteristics, 327-34(1
Sliiinada, r.ell M.; see Wilson ami

Shiiiiada
Simpson, Dniiald A.; see Herald and

Simpson
Skinner, John E. ; California state-wide

angling estimates for 1953, 19-32
Sinilh, Walton M. ; see Macgregor and

Sniilli
Spawning; I mill, raiiilmw, 117
tiphyraena argent ea, lt)7-176
Sport fishing ; see fishing, sport
Stains ; marking starfish, 245-246
Starfish ; nnirking, 245-246
Steelhead ; Jingling esliiiiales, 19-32;

spnri lisliiiig \aliies, 5-17

SI iiifii'ay ; rniiml. 21.*'> I'l 7

Striped bass; sport lisliing values. 5-17

Slrniig. Helen; see l'>issell and Strong;

see P.issell. Ilanis. Slrmig. .-111(1

James
Sliirgenii; while. ;>47-'>4S
Sucker; Klamath small-sc.-iled. llS-119
Siiiifisli ; angling estimates, 19-32

T

Taber. Kicliard D. ; see Dasmann and
Taber

'ragging; callisli. white. 2C)l-2(i9; red
tags, 119-120 ; sturgeon, white,
347-348

'rein]ieratures ; e.-istern tropical Pacific.
79-90

Tnlisnielt. 279-284

T i(ii-hipienis spp., 103-105

'I'r'idkis scmifasciata, 348

Trout ; .n n g 1 i n g estimates, 19-32 ;
diseases, 183-187; hydrostatic
pressure tests. 243-244; rainbow,
19-32, 117, 183-187, 243-244

Tuna ; bigeye, 79-90, 91-98, 99-101 ;
feeding habits, 99-101 ; longlin-
ing, 91-98 ; oceanographic obser-
vations, 79-90; yellowfin, 79-90.
91-98, 99-101

u

( rohdlis hiiUeri, 213-217

w

Wales, J. II. . and IL Wolf; Three
|irotn>;oan diseases nf trniil in
("alifornia, 183-187

Waterfowl ; see hunting areas

Williams, (ieorge C, and Doris C.
Williams ; Observations on the
feeding habits of the opaleye,
Girella nigricans, 203-208

Williams. Doris C. ; see \\'illiaiiis and
Williams

Wilsnii. Robert C, and I'xdl .M.
Shimada ; Tuna longliniiig: Ue-
sults of a cruise to the eastern
tropical Pacific Ocean, 91-98

\\n\{. IL; see Wales and Wolf

Wnnsler. Warren S., and Feenan Jen
iiings ; Exploratory oceanographic
observations in the eastern tropi-
cal I'acitic, January to March,
1953, 79-90

\Clinw1iii I una ; see tuna

jtrilllfil ill TATirORNlA STATr PRINTING OFllCE

20860 7-55 5,200