CAUFDRNlAl
FISH- GAME

"CONSERVATION OF WILDLIFE THROUGH EDUCATION"

VOLUME 43

JULY, 1957

NUMBER 3

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California Fish and Game Is a journal devoted to the conserva-
tion of wildlife. Its contents may be reproduced elsev/here 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
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viduals must state their affiliation and position when submitting
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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, Ediior
Department of Fish and Game
722 Capitol Avenue
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.

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VOLUME 43

JULY, 1957

d

NUMBER 3

Published Quarterly by fhe

CALIFORNIA DEPARTMENT OF FISH AND GAME

SACRAMENTO

STATE OF CALIFORNIA

DEPARTMENT OF FISH AND GAME

GOODWIN J. KNIGHT
Governor

FISH AND GAME COMMISSION

ANDY KELLY, President
Los Angeles

CARL F. WENTE, Commissioner WILLIAM P. ELSER, Commissioner

San Francisco San Diego

WELDON L. OXLEY, Commissioner T. H. RICHARDS, JR., Commissioner

Redding Sacramento

SETH GORDON
Director of Fish and Game

CALIFORNIA FISH AND GAME
Editorial Staff

LEO SHAPOVALOV, Editor-in-Chief ..^ Sacramento

JOHN E. 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

Food Habits of the Tehama Deer Herd

Howard B. Leach and Jack L. Hiehle 161

Report on a Survey of Bighorn Sheep in the Santa Rosa Moun-
tains, Riverside County

Fred L. Jones, Glenn Fliitner, and Richard Gard 179

An Improved Device for Capturing Deer

Gordon Ash craft and Don Reese 193

The Development of a Vinyl Plastic Subcutaneous

Tag for Trout Robert L. Butler 201

Classification, Variation, and Distribution of the Sculpins,
Genus Cottus, Inhabiting Pacific Slope Waters in
California and Southern Oregon, AVith a Key to the
Species C. Richard Robins and Robert Rush Miller 218

Note

The Reported Emergence of Fishes From

California's Subterranean Waters FjOtI S. Herald 235

Reviews 239

(159)

FOOD HABITS OF THE TEHAMA DEER HERD'

HOWARD R. LEACH and JACK L. HIEHLE

Game Management Branch
California Department of Fish and Game

INTRODUCTION

The Tehama deer herd eonsists of Columbian blaek-tailed deer, Odo-
coileus hemionns colnmbianus (Richardson). These deer are mi.uratory,
summerin<>' within and in the vieinity of Lassen Yoh-anic National
Park and winterinji' alonj;' the foothill slopes bounded by the water-
sheds of Battle Creek on the north and Deer Creek on the south (Fig-
ure 1).

In recent years an increase in herd size has been a source of concern
to livestock interests, which are aware of the competition between deer
and livestock for forage on the winter range. During the period from
1942 to l!t4!l the State of California piii-duised 42,897 acres of range
ill the winlei-ing area, with the intent of improving the range conditions
1liroiig]i tlic control of livestock grazing and by inci-easing the hunter
harvest of deer thi'ough construction of ])ubiic access roads.

The ('alifornia Fish and Game Commission in February, 1950, au-
thorized the collection of four female deer per month (not to exceed a
total of 100). TJie ]iuri)()se was to determine the food habits and to col-
lect reproductive and otlier biological data pertinent to this important
deer herd. In addition, supplementary stomach samples were obtained
from hunter-killed deer. Collections were made by the staff of the
California Department of Fish and Game Disease and Food Habits
Laboratory.

The data presented here represent the results of the food habits study
as determined by the stomach analyses of 96 deer collected from the
"winter range and 37 deer collected from the summer range. In addi-
tion, data gathered in the course of a field investigation conducted at
the same time were incorporated into the study.

The important contribution of the forage production of annuals on
the winter range to the diet of deer is demonstrated by the study.
This phenomenon, undoubtedly, is a major factor affecting the carrying
capacity of the range both for deer and livestock, not only on the
Tehama winter deer range, but also on other deer winter ranges of the
California annual type.

1 Submitted for publication December, 1956. This study was carried out by Federal Aid
in Wildlife Restoration Project California W-2.5-R, "Food Habits Investgatons".
The writers wish to express acknowledgment to Robert A. Gardner, Soil-Vegeta-
tion Survey Project, California Forest and Range Experiment Station, and to Dr.
L. T. Burcham of the California Division of Forestry for their constructive sug-
gestions regarding the original manuscript.

2 — .54385

( 161 )

162

CALIFORNIA FISH AND GAME

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DEER FOOD HABITS 163

THE DEER HERD

The Tehama herd summers at elevations of 4,000 to 9,000 feet and
winters from the edge of the valley floor to the 3,500 foot level. A
movement of deer from the summer to the winter range usually occurs
during the middle of October. This migration is well known to sports-
men, and a heavy concentration of hunting etit'ort is directed toward
the deer during the latter part of the hunting season, which in recent
years has varied from the middle to the end of October. Heavy hunting
pressure on the herd also occurs when the deer are on the summer
range and after the deer have concentrated on their winter range. Much
of the area of summer and winter ranges is relatively inaccessible to
the hunters, and there are years when the deer do not move onto the
lower reaches of the range until after deer season.

This herd produces an appreciable harvest of male deer, and some
of the largest black-tailed deer killed in California. Despite the heavy
hunting pressure and a two-buck limit in effect since 1950, there has
been a noticeable increase in deer numbers in recent years. In order
to obtain a greater harvest the Commission authorized a special hunt
quota for taking antlerless deer from November 1 to November 6, 1955.
In 195(). (luring the last three days of the regular season, antlerless deer
could legally be taken without a quota.

Deer were not always as numerous as they have been during the past
decade. A scarcity of deer in the early days prompted the Legislature
to establish Refuge IG in 1917, to give protection to the deer. This
refuge persists today, despite the increase in deer numbers and the
need of additional hunter harvest of deer.

DESCRIPTION OF THE WINTER RANGE

The winter range is one of the most extensive black-tailed deer win-
ter ranges in California. It comprises over 220,000 acres. Geologically,
the area is of volcanic origin, having been influenced by a series of
mud flows attributable to the volcanism of the Lassen Park region to
the northeast. The watercouses have cut this once continuous volcanic
breccia into steep-sided canyons, exposing prominent rim rocks and lava
outcroppings. The most prominent of these are the canyons of Battle,
Paynes. Antelope. Mill, and Deer creeks. In addition, sheet erosion and
weathering have had the effect of leaving some slopes completely bare
to bedrock, others strewn with basaltic rock, and still others with
enough accumulated soil to support vegetation. Often vegetation will
be found along lateral seams, where soil and moisture conditions are
favorable. A strikingly striated appearance of vegetative growth, char-
acteristic of much of the region, has resulted (Figure 3).

AYork done by the California Soil- Vegetation Survey ^ indicates that
the woodland-grass and grass types of the vegetative cover occur on
the Toomes soils and the brush and dense hardwoods on the Stover
soils. The Toomes soil series is found predominantly throughout the
area and varies in depth from 8 inches to 24 inches. The deeper Stover
soil series (30 inches to 40 inches in depth) supports the brush and
hardwood vegetative types and occurs on colluvial stringers or benches

' This survey is conducted by the California Forest and Range Experiment Station and
the University of California under contract with the California Division of For-
estry.

164

CALll-UKXIA FISH AM) (iA.MK

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FIGURE 2. The woodland-grass cover type which characterizes the lower reaches of the Te-
hama winter range. Deer graze these areas in the fall months for the fallen leafage and
acorns of blue oak. With the advent of early fall rains these lower foothills are the first to
"green up" with the production of annual forage and ore grazed extensively by deer and
livestock. Photograph by Howard R. Leach.

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FIGURE 3.

The underlying soil types are largely responsible for the striated vegetative growth
illustrated here. Photograph by Howard R. Leach.

DEER FOOD HABITS

165

FIGURE 4. The deeper soils of the north-facing slopes support dense chaparral and hardwood

stands. Phoiograph by Howard R. Leach.

tliroiighoiit the Toomes soils, primai-ily on tlie iiorth-facin^i' slopes
(correspondeiife witli I\ol)('i't A. (Ijirdiici-. I!)')?). Figures "2, 8, and 4
illustrate the vegetative aspect of the winter range.

The Tehama \vinter deer range is, like most t'ootluU seetions of
eismontane California, an annual range ty])e in its lower reaches, with
the eharaeteristie range j^roblems inherent in this type. The wide and
irregular changes in forage production from year to year due to cli-
matic Unci nations not only directly atfect the livestock grazing of these
annual ranges but also have an impact on the carrying capacity for
wildlife. Deer are no exception.

STUDY AREA

Public access to the winter range north of Antelope Creek is provided
by State Highway 36 and a number of unpaved county roads. JSoutli of
Antelope Creek the range is less accessible, being limited mostly to
jeeps and other four-wheel drive vehicles. One such jeep road traverses
the rim of Dye Creek and ]\[ill Creek canyons from the valley floor to
the lower edge of the summer range. This road provided access to the
deer over the entire period, and allowed coverage of an area repre-
sentative of most of the winter range. Consequently, this area w^as
selected for studv.

VEGETATIVE DESCRIPTION OF THE STUDY AREA

Figure 5 shows a portion of a vegetative type cover map of the
Tehama Avinter deer range delineating the study area. This map was
drawn from aerial photographs by William P. Dasmann, using the
techniques described by Jensen (1947). It gives the gross characteristics

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rAT.IKOKNIA FISTT AND GAME

of tlie study area relative to the interspersal of the vegetative cover
types.

Essentially, the vegetative complex consists of four broad types, de-
scribed as follows and shown by symbols in Figure 5.

FIGURE 5. A vegetative type cover map of the study area, showing the interspersion of the
vegetative types. Spots indicate where deer were collected.

The lower reaches of the foothills are, predominantly, either a grass
(G) or a woodland-grass (W) type. The woodland is of blue oak with
an understory of annual grasses consisting mostly of soft chess, ripgut
brome, red brome {Bromus ruhens), foxtail fescue {Festiiea megalura),
wild oat (Aveua sp.), nitgrass {Gasfyidium sp.), and wild barley.
These annual grasses are also dominant in the grass type. Some peren-
nial grasses such as needlegrass, Sitanion, Melica, and blue grass are
found but constitute little of the vegetative composition. These two
vegetative types persist throughout most of the winter range.

The annual grasses and such forbs as the filarees, popcorn flower,
hillside lotus, lupine, bur clover, and chickAveeds germinate with the
advent of early fall rains. These are supplemented over the winter aud
early spring months by numerous species of native and non-native
broad-leafed forbs. Above the 1,000-foot elevation the woodland grass
occurs for the most part on the shallow Toomes soils of the hillsides
and swales and more predominantly on the south-facing slopes. Inter-
spersed over the range are areas of chaparral or brush (B). The browse
species characterizing the chaparral consist principally of buck brush,
California scrub oak, w^estern mountain mahogany, silktassel, and
manzanita. Scattered in the chaparral are foothill ash, western hoptree,
poison oak, California buckeye, yerba santa. chaparral honeysuckle,

DEER FOOD HABITS 167

redbud, and California juniper. Many of these browse plants are de-
ciduous. There are occasional thickets of California black oak, but this
oak, together with Brewer oak, is more prevalent at the upper margin
of the winter range. Along the watercourses grow Fremont cottonwood,
western chokecherry, creek dogwood, willow, alder, and California wild
grape. The density of the chaparral varies from thick, mature stands,
noticeably on the north-facing slopes, to opened-up brush fields on the
more shallow soils and in areas influenced by past fire history. The
hardwood stands (H) characterizing the deeper soils consist mostly of
mature oaks, either growing as a dense woodland of blue oak or stringers
of interior live oak along the fissures and lava outcroppings.

One hundred and one line-point forage plots were established to
collect data relative to the ground cover and vegetative composition of
the Teliama winter deer range. An analysis of the ground cover com-
position based on these plots is given in Table 1. It is considered to be
representative of the entire winter range.

Table 2 gives the composition of the vegetative cover as determined
by these same 101 line-point plots and is an index to the relative abun-
dance of the principal plants making up the forage composition.

TABLE 1
Percentage Composition of Ground Cover on ttie Tehama Winter Deer Range

Nonproductive 54.8

Annual grasses and forbs 27.2

Perennial grasses 0.5

Browses (shrubs and trees) 17.5

TABLE 2
Percentage Composition of the Vegetative Cover on the Tehama Winter Deer Range

Annual grasses and forbs 59.9

Perennial grasses and forbs 11.5

Total herbs 71.4

Buck brush 6.3

California scrub oak 4.5

Brewer oak 2.9

Manzanita 1.5

Western mountain-mahogany 1.5

Squaw carpet 1.3

Deer brush 1.2

Poison oak 1.4

Blue oak 0.9

Yerba santa 0.7

California black oak 0.6

Oregon grape 0.5

Western chokecherry 0.5

Silktassel 0.5

Interior live oak 1.0

Ponderosa pine 0.4

Squaw bush 0.4

Creek dogwood 0.2

Other browses 2.3

Total browse 28.6

168

CALIKOK-MA llSli AND (iAME

SEPT. OCT.

NOV, DEC.

JAN.

FEB. MAR. APRIL

FIGURE 6. Average monthly precipitation at Red BlufF for the winter months of 19481952,
showing wide seasonal fluctuations in rainfall. Drawing by CliFfa Corson.

CLIMATE

The cliiuato is characteristic of much of the Central Valley foothills
refrion of California. Generally, winter temperatures are mild and
summer temperatures high. A rainj^ winter season is followed by a
rainless summer period. Comparatively gentle rains occur normally in
October and November and increase in intensity during the winter.
Snow occurs occasionally at the lower elevations but melts off rapidly.
At the higher elevations snow remains for varying periods. The rainy
season is often accompanied by fog, which may persist for many days
in succession. Seasonal precipitation both in amount and distribution
is the main factor infliiencing forage conditions on the winter range.

DEER FOOD HABITS

169

In addition to precipitation, the growing period of the annual vegeta-
tion is influenced by dry, cold, northerly winds, which occur frequently
and inhibit the growth of annual A^egetation.

Unfortunately, weather data relative to precipitation, temperature,
and snowfall are not available for the area encompassed by the winter
range. The U. S. Weather Bureau maintains a permanent weather
station on the edge of the summer range at ^Mineral, at an elevation of
4,850 feet, and another on the valley floor at Red Bluff, elevation 341
feet. The fluctuation in seasonal rainfall at Red Bluft' for the winters
of 1948-49, 1949-50, 1950-51, and 1951-52 is shown in Figure 6. The
four winters represented illustrate the variance in seasonal precipita-
tion which is characteristic of the area.

Considering the period of study — February, 1950, to December, 1951
— it may be seen that the later winter months of 1949-50 were months
of relatively high precipitation which had been preceded by an ex-
tremely dry fall period and that the winters of 1950-51 and 1951-52
were wet seasons.

WINIER FOOD HABITS

The food items identified in the 96 deer stomach samples collected
during the winter study period are summarized in Tables 3 and 4. Table
3 gives the volume in percentage and frequency of occurrence in per-
centage of those food items making up 0.1 percent or more of the diet
for any one month of the collection. Table 4 is a supplementary list
expressed in frequency of occurrence in percentage of those food items
contributing less than 0.1 percent of any one monthly period.

A graphic representation of the monthly dietary changes in food
habits in given in Figure 7. The following discussion of the food habits

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NOV.

DEC.

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7

JAN.

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MAR.

13

APR.

8

MAY

FIGURE 7. A graphic representation of the diet of deer on the Tehama winter deer range,
showing the seasonal variations in food habits. Drawing by Cliffa Corson.

3 — 54385

170

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172

CALIFORNIA IlSIl AND ClAME

TABLE 4
Supplemental List of Food Items Eaten by 96 Deer Collected on ttie Tehama Winter Deer Range

Scientific name

Browse

PimLS ponderosa

Pinus sabiniana

Juniperus californica

Torreya californica

Salix sp

Arceuthohium sp

Phoradendron villosum

Umbelhdaria californica

Aesculus californica (fruit) .

FremorUia californica

Chrysothamnus sp

Unidentified browse

Forbs

Lichen

Bryophyta

Pellaea andromadaefoUa--

Polypodiaceae

Selaginella sp

Liliaceae

Polygonum bolanderi

Polygonum sp

Rumex sp ^ —

Amaranthus sp

Mollugo verticillata

Montia perfoliata

Stellaria sp

Silene sp

Carj'ophyllaceae

Ranunculus californicus.-

Athysanus pusillus

Lepidium nitidum

Platyspermum scapgerum.

Thysanocarpus sp

Crucif erae

Saxifraga sp

Lotus sp

Lupinus sp

Trifolium sp

Limnanthes alba

Euphorbia sp

Leguminosae

Malvaceae

Epilobium sp

Godetia sp

Sanicula sp

Daucus pusillus

Lomatium. sp

Amsinckia sp

Trichostema lanceolatum..

Galium sp

Ambrosia psilostachya

Hemizonella minima

Eriophyllum lanatum

Centaurea m^litensis

Compositae

Common name

Yellow pine

Digger pine

California juniper-.
California nutmeg -

Willow

Pine mistletoe

Common mistletoe-
California laurel

California buckeye-
Flannel bush

Rabbitbrush

CoSee fern - .
Fern family _
Club moss--
Lily family -

Knotweed

Dock

Amaranth

Indian chickweed

Miner's lettuce

Chickweed

Catch-fly

Pink family

California buttercup -

Common peppergrass.

Fringe pod

Mustard family _

Saxifrage

TrefoU

Lupine

Clover

Spurge

Pea family

Mallow faiuUy

Willow herb

Godetia

Snake root

Rattlesnake weed-

Hog-f ennel

Fiddleneck

\^Lnegar weed

Bedstraw

Western ragweed -

Wool j'-yarrow

Napa thistle

Sunflower family.

Percentage

frequency of

occurrence

8.3
6.2
5.2
1.0
2.0
2.0
5.2
2.0
1.0
2.0
1.0
3.1

6.2
10.4
5.2
2.0
2.0
6.2
9.4
2.0
1.0
1.0
1.0
7.3
2.0
1.0
1.0
2.0
8.3
15.6
3.1
2.0
7.3
2.0
6.3
2.0
1.0
1.0
3.1
1.0
1.0
1.0
1.0
1.0
1.0
2.0
1.0
1.0
7.3
1.0
1.0
1.0
1.0
7.3

DEER FOOD HABITS

173

TABLE 4— Continued
Supplemental List of Food Items Eaten by 96 Deer Collected on the Tehama Winter Deer Range

Scientific name

Common name

Percentage

frequency of

occurrence

Grass and grass-like plants

Agropyron sp

Avena fatua

Bromus mollis

Bromus rigidus

Festuca sp

Hordeum sp

Stipa sp

Poa sp

Cyperaceae

Juncus sp

Wheat grass..

Wild oat

Soft chess

Ripgut brome

Fescue

Wild barley--
Needlegrass--

Bliie grass

Sedge

Rush-.l

2.0
1.0
42.7
1.0
3.1
4.1
1.0
1.0
5.0
1.0

of the deer on the Tehama winter deer range is based chiefly on the
analysis of the data presented in the above tables and Fignre 6. Admit-
tedly, the sampling is small, but it does give evidence that deer experi-
ence a seasonal dietary change influenced by the availability of forage as
determined hj seasonal climatic changes.

Normally the deer enter the winter range in appreciable numbers
during the middle of October, the does and fawns ordinarily preceding
the bucks. Once they reach the winter range the}- disperse over the area
and remain approximately 210 days. Generally by the first w^eek in May
they have started their return to the summer range. Their exodus from
the summer range is generally attributed to a response to weather condi-
tions. It has been observed that in years of good acorn production the
deer tend to move to the winter range in large numbers earlier than in
3^ears of acorn failure. Upon entering that area contiguous with the
summer range, deer seek out and utilize the acorns and fallen leafage of
California black oak and Brewer oak and supplement this diet with the
leafage of squaw carpet, deer brush, white fir, and incense cedar. They
soon move to the lower elevations and are commonly seen grazing the
woodland areas, seeking the leafage and acorns of blue oak.

"With the advent of the early rains in October and November there is
an immediate response in the germination of the annual forage plants
and the foothills "green up." These annuals normally make little
growth in the months of December, January, and early February, but
in late February and March they grow and mature rapidly. Deer begin
to utilize the growth of green grass and forbs as soon as it makes its
appearance in October and November and continue to use it until the
plants reach maturity in April. The heaviest consumption of herbaceous
food is apparent in March, during the period when it is most available.
However, as soon as the new leafage appears on the deciduous and ever-
green browse plants, an abrupt change to browse consumption is made.

Browse is the staple item of diet and it is obvious that it becomes
the principal means of support in winters of adverse growing conditions
(when there is little production of annual forage) or when an acorn
crop failure occurs. There appears to be a seasonal preference in the
utilization of browse species. Buck brush receives consistent usage by
deer throughout most of the winter. It is the most abundant shrub and

174; CALIFORNIA FISH AXD GAME

one which forage utilization checks reveal to be heavily utilized. Manza-
nita contributes little bulk to the diet, altliou^di it is eaten in every
month of the Avinter. "Western mountain-mahogany is a semideciduous
plant and apparently is most heavily utilized in the fall months and
again in April and May, "when the young growth becomes available.
This is noticeably true of the other deciduous browses such as redbud,
California buckeye, hop-tree, poison oak, and deer brush. During the
months of December, January, and early February, when the herba-
ceous plants are normality retarded in growth, deer rely more heavily
on buck brush, manzanita. scrub oak, silktassel, and yerba santa for
food. Forage utilization checks show" use to be heaviest on western
mountain-mahogany, silktassel, and buck brush. The transition from a
diet composed largely of herbs to one of browse is most pronounced in
April and Maj', when the browse plants sprout new leafage and deer
are quick to take advantage of this source of food.

It has been observed that a 20-day difference in forage growth condi-
tions exists between the 1,000- and 3,000-foot levels. Annual forage
which has reached maturity at the lower elevations in February and
March is in various stages of development at the higher elevations. This
altitudinal response in forage production is most noticeable in the
spring months, when the deciduous trees and shrubs begin to sprout
new leafage. The altitudinal distribution of deer in the winter months,
and especially so during the period of spring migration, is largely in re-
sponse to this phenological change in forage production as it affects both
the palatability and availability of food.

SUMMER RANGE DEER FOOD HABITS

The food habits of the Tehama deer herd on the summer range re-
ceived less intensive study. The limited sampling of 37 deer stomach
samples was represented by 32 deer collected monthly by the Disease
and Food Habits Laboratory staff from July through September of
1950 and 1951 and five stomach samples obtained from hunter-killed
deer. The general locality of the special collection was in the area
southeast of Lassen Volcanic National Park and north of Lake Almanor.
The collection included a sampling of deer from the Childs Meadow,
Wilson Lake, Warner Creek, and Mud Creek areas in Plumas and
Tehama counties.

The summer range of the Tehama deer herd, as described by Grinnell
et al. (1930), reaches eastward around the northwestern base of Lassen
Peak, at least to the vicinity of Manzanita Lake. In the southern por-
tion of the area it has been found to extend down Warner Creek from
Hot Springs Valley to Lake Almanor, and thence eastward to the
vicinity of Westwood.

This summer range, with the exception of the higher elevations, is
predominantly the transition forest climax characterized by a mixed
forest of yellow pine, sugar pine, Douglas fir, white fir, and incense
cedar. The under-storj^ consists principally of mountain whitethorn,
snowbrush, squaw carpet, greenleaf manzanita, chinquapin, bitter
cherry, western service berry, and gooseberry. Where logging and fire
have removed the forest canopy, extensive brush fields of greenleaf
manzanita, mountain whitethorn, and snowbrush have developed as a
subclimax. Figure 8 is a photograph of typical summer range country.

DEER FOOD HABITS

175

FIGURE 8. An illustration of the secondary succession following fire and/or logging operations
in the transition forest climax, typical summer range. Mountain whitethorn and greenleof man-
zanita form the principal subciimax and are important forage species for the deer. Photo-
graph by Howard R. Leach.

Table 5 is a composite summary of the food items eaten by the 37
deer collected from the summer range. Browse made up 72.1 percent
of the diet. Based on the percentage volume and frequency of occur-
rence, the preferred browse plants consisted of mountain whitethorn,
squaw carpet, greenleaf manzanita, western service berry, and snow-
brush. Mountain whitethorn was the most important source of food,
having constituted 33.2 percent of the diet and being found in 78.4
percent of the stomachs. This shrub is an abundant and widely dis-
tributed browse plant in the Sierran montane forest. It occurs from
4,000 to 9,000 feet and may be found growling as a forest understory
or as the dominant in the subciimax montane chaparral which develops
following fire and/or extensive logging operations. Forbs apparently
contribute more heavily to the summer diet than does grass, since
forbs formed 25.1 percent of the diet. Grass, although found in 72.9
percent of the stomachs, contributed but 2.8 percent of the food eaten.
Numerous forbs were identified in the stomachs, but the bulk of this
herbaceous food was unidentified.

Admittedly, determination of the food habits of deer on the summer
range would require considerably more samples of deer stomachs than
the 37 analyzed in Table 5. In such a study consideration would have
to be made of the vegetational changes characterizing the various life
zones over which the deer distribute themselves over the summer
periods. The present limited sampling precludes such an analysis.

176 CALIFORNIA FISH AND GAME

TABLE 5
Food Items Eaten by 37 Deer Collected From the Summer Range of the Tehama Deer Herd

Scientific name

Common name

Percent-
age
volume

Percent-
age
frequency

of
occurrence

Browse

Ceanothus cordulatus

Ceanothus prostratus

Arctostaphylos patula

Ceanothus velutinus

ATnelanchier alnifolia

Castanopsis sempertirens

Ceanothus integerrimus

Alnus tenuifolia

Ribes sp

Abies concolor

Unidentified browse

Librocedrus decurrens

Prunus emarginata

Prunus virginiana var. demissa.

Chrysothamnus parryi

Finns ponderosa

Pinus contorta

Salix sp

Populus trichocarpa

Quercus kelloggii

Total browse -

Forbs

Unidentified forbs

Ranunculus alismaef alius ^

Pedicularis semibarbata

Menyanthes trifoliata

Polypodiaceae

Liliaceae

Lichen

Fungus

Lotus sp

Bryophyta

Polygonum sp

Rumex sp

Eriogonum sp

Chenopodiaceae

Montia sp

Potentilla sp

Lupinus sp

Trifolium sp

Leguminosae

Epilobium sp

Umbellif ereae

Phlox sp

Collomia tinetoria

Taraxacum sj)._

Wyethia sp

Compositae

Total forbs -

Grass and grass- like plants

Gramineae (green)

Gramineae (dry)

Bromus tecU/rUTn

Stipa sp

Carex sp

Juncus sp

Mountain whitethorn-

Squaw carpet

Greenleaf manzanita.-

Sno wbrush

Western service berry -

Chinquapin

Deer brush

Mountain alder

Gooseberry

White fir

Incense cedar

Bitter cherry

Western choke-cherry-

Rabbitbrush

Yellow pine

Lodgepole pine

Willow

Black Cottonwood

California black oak__

Total grass -

Buttercup

Lousewort

Buckbean

Fern family

Lily family

Lichen

Mushroom

Trefoil

Moss

Knotweed

Dock

Wild buckwheat -
Picwecd family- -
Miner's lettuce..

Five finger

Lupine

Clover

Pea family

Willow herb

Parsley family __
Phlox

Dandelion

Mule's ear

Sunflower family.

Grass family

Grass family

Cheat grass

Porcupine grass.

Sedge

Rush

33.2

7.3

6.2

6.1

3.7

2.7

2.7

2.7

2.4

2.0

1.0

1.0

0.9

0.1

0.1

trace

trace

trace

trace

trace

72.1

25.1

2.8
trace
trace
trace
trace
trace

2.8

78.4

45.9

48.6

18.9

45.9

10.8

2.7

8.1

10.8

27.0

8.1

51.4

16.2

5.4

5.4

37.8

5.4

5.4

2.7

5.4

14.0

86.5

5.2

18.9

2.6

2.7

1.0

2.7

0.9

8.1

0.7

8.1

0.5

59.5

0.1

16.2

0.1

2.7

trace

2.7

trace

13.5

trace

2.7

trace

5.4

trace

2.7

trace

5.4

trace

13.5

trace

8.1

trace

21.6

trace

2.7

trace

2.7

trace

2.7

trace

2.7

trace

5.4

trace

8.1

trace

5.4

trace

2.7

72.9
5.4
2.7
2.7
8.1

13.5

DEER FOOD HABITS 177

DISCUSSION

This study of the food habits of the Tehama deer herd was limited
to a period from February, 1950, to December, 1951. "With the possible
exception of the influence of drought conditions on the forage, pre-
ceding the initial collection period in the late winter months of 1949-50,
the winter range experienced a good production of annual forage dur-
ing the winters of 1950-51 and 1951-52. These winters were notably wet.
It is obvious that the availability of food, with respect to the herbaceous
vegetation on an annual type winter range, is influenced seasonally
and from year to year by variations in precipitation and temperature.
These two factors determine not only the quantity of forage produced
but also the quality and time of growth. The impact of good forage
])roducti(jn on the food habits of the deer inhabiting this type of winter
range is well illustrated by the data presented. It is reasonable to be-
lieve that ill winters of fall drought and/or extreme cold weather the
maximum carrying capacity of the range would be severely reduced
and the natural reduction of deer numbers by winter die-off would
result. Tliis phenomenon of reduced carrying capacity is commonplace
knowledge with livestock people, who rely heavilj' on the foothill ranges
to graze their livestock.

A failure or severe reduction of annual forage production would
force deer to subsist more heavily on the browse component of the
range forage. Snow depth and/or extreme cold temperatures would
not only materially affect the altitudinal distribution of deer on the
winter range, but would also have an immediate effect on the avail-
ability of forage upon which they must subsist.

The data available during the study period preclude a discussion
of the effect of an adverse winter on the food habits and survival of
deer. Had a critical winter occurred during the course of the food
habits study, the contrast in food habits data possibly would have
allowed a better understanding of the dynamics of deer survival and
range conditions relative to browse utilization and annual forge pro-
duction on an annual type winter range.

SUMMARY

A food habits study was made of a migratory herd of Columbian
black-tailed deer known as the Tehama deer herd. This herd summers
in the mountains of the Cascade Range and Sierra Nevada and winters
on the foothill slopes of eastern Tehama County, California. The winter
range is an annual type range. Annual grasses dominate the grassland
areas and form the principle understory to the woodland and chaparral.
There is wide variation both seasonally and yearly in the production of
this annual forage, due to climatic fluctuations.

The stomach analyses of 96 deer collected on the winter range reveal
a seasonal utilization of food plants as influenced by the changes in
range forage composition. Browse (buck brush, mountain mahogany,
manzanita, scrub oak, silktassel. and many deciduous browses) appear
to constitute the staple item of diet throughout the winter. The leafage
and acorns of the deciduous oaks contribute heavily to the early fall
diet, and as the foothills "green up" with the advent of fall rains,
the deer graze heavily on the annual grasses and forbs. This annual

178 CALIFORNIA FISH AND GAME

forage production reaches its height in March, when it contributes
most to the diet of the deer. Early in April tlie browse plants begin
to leaf out with new growth and the doer appear to switch entirely
to a browse diet.

The summer food habits of the Tehama deer herd are represented
by 37 deer stomach analyses. The preferred browse species consist of
mountain whitethorn, squaw carpet, greenleaf manzanita, western
service berry, and snowbrush.

REFERENCES
Burcham, L. T.

1956a. Historical backgrounds of range land use in California. Jour. Range

Mangt., vol. 9, no. 2, p. 81-86.
1956b. Historical geography of the range livestock industry of California. Doc-
toral Thesis, Univ. of Calif., 416 p. (Typewritten)
Gilman, James H.

1949. Annual report on Tehama winter deer range. Calif. Dept. of Fish and
Game, 44 p. (Unpublished report)
Grinnell, .Joseph, .Joseph Dixon, and .Jean M. Linsdale

1930. Vertebrate natural history of a section of northern California through
the Lassen Peak region. Univ. Calif. Publ. Zool., vol. 35, 594 p.
Hiehle, Jack L.

1951. Annual report on Tehama winter deer range for the period .July 1950 to
.July 1951. Calif. Dept. of Fish and Game, 50 p. (Unpublished report)
Jensen, Herbert A.

1947. A system for classifying vegetation in California. Calif. Fish and Game,
vol. 33. no. 4, p. 199-266.
Talbot, M. W.. H. H. Biswell, and A. L. Hormay

1939. Fluctuations in the annual vegetation of California. Ecologv, vol. 20, no. 3,
p. 394-402.

REPORT ON A SURVEY OF BIGHORN SHEEP IN THE
SANTA ROSA MOUNTAINS, RIVERSIDE COUNTY'

FRED L. JONES

Game Management Branch

California Department of Fish and Game

and

GLENN FLITTNER and RICHARD GARD

University of California

Berkeley

INTRODUCTION

During the month of July, 1953, a survey of desert bighorn sheep
(Ovis canadensis nelsoni) was conducted in the Santa Rosa Mountains
of Riverside County by personnel of the California Department of Fish
and Game and the T^niversity of California. The TTniversity personnel
were empkn-ed by tlie Riverside County Fish and Game Commission.
Primary emphasis was placed on State Game Refuge 4-D.

The main purpose of the surv'ey was to establish the distribution
and numbers of bighorn sheep. Additional data on herd composition,
food habits, range condition, water, parasites, and domestic stock were
gathered.

Since 1953 periodic cheeks have been made of the areas covered in
the original survey and of several new areas by Richard Weaver, Floyd
Vernoy, and Bert Craig of the California Department of Fish and
Game in the course of developing and improving springs, and also by
the senior author.

Acknowledgment is made to Mr. Arthur Nightingale, at whose camp
we stayed during the survey in 1953, and who gave us considerable
assistance in our search for information.

METHODS

An aerial reconnaissance of the Santa Rosas was made in June, 1953,
preliminary to the field work. By this method several springs, tanks,
and sheep trails were located, aiding the authors in becoming acquainted
with the topography and drainages.

Inasmuch as the entire Santa Rosa bighorn range could not be cov-
ered in the allotted time, it was planned first to survey the refuge and
then to expand north and south until time ran out. Canyon bottoms,
sidehills, and ridge tops were coursed in search of water sites, bighorn,
and sign.

1 Submitted for publication Augnst, 1956. This study was financed by Federal Aid in
Wildlife Restoration Project W-41-R, "Big: Game Studies", and by funds made
available by the Riverside County Fish and Game Commission.

( 179 )

180

CALIFORNIA FISH AND GAME

SANTA ROSA MOUNTAINS
BIGHORN RANGE

Bighorn Summer Range
rv\\^A^\\] Bighorn Winter Ronge

we>'"

FIGURE 1. Map of the Santa Rosa Mountains area, showing bighorn range.

Drowing by C/iffo Corson.

DESCRIPTION OF AREA

The Santa Rosa ^Mountains form a southeastern extension of the
San Jacinto Mountains, being separated from them by Palm Canyon.
The highest summit, Toro Peak, is 8,716 feet in elevation. Secondary

BIGHORN SHEEP SURVEY

181

summits range from 4,000 to 8,000 feet. The foothills rise from the
Coachella and Borrego valley floors near sea level.

Higher slopes, above 4,000 feet, are covered with chaparral, piiion
pine, and juniper. Lower slopes support a desert scrub type of vegeta-
tion composed in part of the following species: maguey {Agave con-
sociata), Spanish dagger {Yucca schidigera), ocotillo {Fouquieria
splendens), California juniper {Juniperus calif ornica), cactus {Opun-
tia sp., Echinocactus sj)., Cevens sp.),catclaw {Acacia Greggii) , eY>liedva
{Ephedra sp.), briekellbush {Brickellia sp.), wild buckwheat {Eriogo-
wwmsp.), white ratany {Krameria (7 rai/i), black sage {Salvia mellif era) ,
white sage {Salvia apiana), bee-sage {Hyptis Emoryi), goatnut {Sim-
mondsia chinensis), rush bebbia {Behbia juncea), ground cherry
{I'liysidis crassijolia ) , l*arry condalia {Condalia Parryi), creosote busli
{Larrea divaricata), French tamarisk {Tamarix gallica), honey mes-
quite {Prosopis chilensis), screwbean mesquite {Prosopis puhescens),
desert-willow (Chilopsis linearis), squawbush {Rhus trilohata), and
galleta grass {Hilar ia rigida).

Some species that generally grow close to water are : palm ( Washing-
tonia filifera), cottonwood {Popidus Fremontii), willow {Salix sp.),
water-wally {Baccharis ghdinosa) , arrowweed {Pidchea sericea), horse-
tail {Equiseiuin sp.), connuon reed {Phragmites communis), cattail
{Typha latifolia), and rush {Juncus sp.).

The lower slopes are steep and cut with many canyons. In many
places, as in Guadalupe Canyon, Deep Canyon, the mouths of Magnesia
Spring Canyon and Cat Creek, and at the junction of Ebbens, Grape-
vine, and Dead Indian creeks, hard rock strata have been cut, forming
perpendicular cliffs and waterfalls.

FIGURE 2. Larrizo Creek area, looking north across the Palms to Pines Highway. A ewe and
ram can be seen near the middle. Photograph by Fred L Jones, Sepfember 16, J 954.

182 CALIFORNIA FISH AND OAME

Ou the east slope of the Santa llosas a zone of undulating, low-angle,
open terrain extends southward above the 1,500-2,000 foot level and
below the mountain summits to about Guadalupe Creek, interrupted
only by Deep Canyon (Figure 2). To the south the canyons become
deeper, and the ridges attain greater stature.

BIGHORN DISTRIBUTION

At present the bighorn occupy the lower country, generally below
4,000 feet. Figure 1 shows the winter and summer ranges to the best
of our knowledge.

Water is the primary factor determining bighorn distribution. In
winter, when moisture is widespread and the climate is cool, the range
is more extensive. In summer the Santa Rosa sheep have not been
found by us ranging farther than about one mile from available water.

The upper limit of the range is roughly determined by the chaparral
line. The lower limit is indeterminate, although the sheep generally
remain in the mountains or close to them. They often feed in the dry
washes, and according to local residents it is not uncommon for them to
come down into cultivated fields as far as two miles out from the base
of the mountains during winter. We have an account of one being killed
along the Southern Pacific Railroad tracks some years ago.

Old residents state that in the early days the sheep were found higher
in the mountains than they are now. Although little of the higher
areas appear to us to be suitable range, it is possible that the greater
numbers reported prior to the die-off in the 1920 's (which will be
discussed later) required greater range, and so had dispersed upward.
It is possible that livestock use in the higher elevations has contributed
toward restricting the sheep to their present altitudinal distribution.

Owing to water localization, some areas receive heavy summer use.
Magnesia Spring Canyon, Cat Creek, and Carrizo Creek were the only
sources of summer water located north of the Palms to Pines Highway
on the eastern slope, and were supporting all, or nearly all, of the sheep
in that sector in Julj^, 1953. No sheep were ranging in the intervening
country on Ebbens, Dead Indian, and Grapevine creeks at that time.
Although we covered little of the country north and east of Sheep
Mountain, the same situation appeared to exist between Deep Canyon
and Agua Alta Canyon, with only a few sheep occurring along Guada-
lupe Creek.

Sheep range continuously from Agua Alta Canyon to the southern
end of the mountains. It is probable that there are sheep on the Palm
Canyon side of the Santa Rosas, but we have been unable to survey
that area or to obtain any information on it.

NUMBERS AND COMPOSITION

Old residents told us of a severe die-off in the early 1920 's. Many
carcasses were found in the canyons and around waterholes by pros-
pectors. The general expression of opinion is that there were more
sheep in the Santa Rosas prior to this period than there are now, but
that the number is greater now than at any time since the die-off.

Table 1 lists the sheep actualh^ observed in July, 1953, with all pos-
sible duplications eliminated, and the number estimated in each area.

BIGHORN SHEEP SURVEY

183

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184

CALIFORNIA FISH AND GAME

Population estimates for the areas visited were made on the basis of
sheep seen, the relative numbers of rams to ewes and of young to adults,
and the amount of fresh si*rn observed. If the fresh sifrn appeared
to be greater than the observed animals could be expected to make, a
conservative addition was made. No estimate was possible for the Palm
Canyon area, although we feel certain that sheep are present there, if
water is available.

The estimates for the areas south of Sheep Canyon are based pri-
marily upon spot checks made since the original survey. These figures
are supported by information from local residents whose judgment
proved to be sound in other areas that we covered more completely.

We believe that all of the estimates are conservative and have little
doubt that the total population is larger.

The 86 sheep estimated to be in the refuge comprise 25 percent of
the total estimated in the entire Santa Kosas, and the 60 in Borrego
State Park, 17 percent.

From Table 1 it may be seen that the Cat Creek-Magnesia Spring
Canyon area immediately to the north of Refuge 4-D, and the area
south of the refuge to Borrego Park, contain the bulk of the sheep.

Unquestionably some two-year-old rams are included in the ewe
classification, since identification of sexes by horns cannot be made with
certainty before three years of age. Figure 3 shows a f-curl ram. The
scrotum of a two-year-old ram must be seen for positive identification,
and it is not always possible to make such an obserA'ation. Information
given us indicated that the 1051 lamb crop was extremely small. Since
these lambs were 2-year-olds in 1953 we believe that possible mis-sexing
of two-year-old rams was of slight importance, due to a low percentage
of them in the population.

li&URL j, A iriiKc-quarter curl ram, Santa Rosa Mountains. Photograph by
Fred L. Jones, Sepfember 16, 1954.

BIGHORN SHEEP SURVEY

185

The ratio of rams to ewes observed was 109 :100 ; of lambs to ewes
18:100; and of yearlings to ewes 27:100. Exactly two-thirds of the
rams had a three-quarter curl or better ; that is to say, they were in the
older age classes. Table 2 gives the computed number of each sex and
age class in the estimated populations based on the observed ratios. The
animals observed make up 40 percent of those estimated.

TABLE 2
Computed Number in Each Sex and Age Class of Estimated Populations

Area

Young
rams*

Old

riunst

Ewes

Yearlings

Lambs

Totals

Refuee 4-D _ _

13

7

31

2.5
14
02

33
20
84

9

.5

23

C

4

15

86

.50

Rest of Santa Rosas

215

Totals in Santa Rosas

51

101

137

37

25

351

* Less than three-quarters curl.
t Three-quarters curl or better.

The observed sex ratio indicated an overabundance of rams. Since
bighorn sheep are polygamous, they do not require a ram for every
ewe for breeding purposes. During the mating season only a few ewes
of a band are in heat at any one time, and the rams consequently give
these much attention (Jones, 1950).

The optimum sex ratio in deer is often said to be about 25 :100. It
may be more than that in bighorn, but 50 :100, or one ram for every
two ewes, should be ample for breeding. It is certain that fewer rams
than there are now would be able to breed all the ewes. Table 3 gives
sex and age ratios found in other herds of desert bighorn.

TABLE 3
Sex and Age Ratios Observed in Desert Bighorn Herds

Area

Num-

Year

Ram:

Lamb:

Yearling:

ber

Ewe

Ewe

Ewe

obser-

ved

1953

109:100

18:100

27:100

141

1938

89:100

22:100

11:100

20

1955

57:100

29:100

8:100

95

1951

121:100

65:100

28:100

154

1952

137:100

43:100

11:100

110

1953

123:100

50:100

17:100

156

Reference

Santa Rosa ^Mountains

Death Valley

Death Valley

Arizona

Arizona

Arizona

Dixon (1939)
WeUes (1955)
Russo (1956)
Russo (1956)
Russo (1956)

Ewes normally produce single lambs, twins being rare (Jones, 1950).
The observed lamb:ewe ratio of 18:100 is the lowest reported in any
survey of desert bighorn for which we have data. Jack Miller, an old
resident, told us that there was a large lamb crop in 1949, no lambs or
very few in 1951, a large crop again in 1952, and a very small one in
1953. "We have certainly confirmed this last report and our observed

4—54385

186

CALIFORNIA FISH AND GAME

yearling: ewe ratio of 27:100 in 1953 tends to confirm this statement
for 1952. This yearliiij; ratio compares favorably' with those found
olspAvhere.

The lamb crop appears to be closely tied in with the pix'cipitation for
the year. In wet years there seem to be larger lamb crops than in dry
years. A deficienc}' of vitamin A in the diet of ewes, resulting from
a shortage of green feed during dry j'-ears, may preclude ovulation or
may cause the death of lambs in the uterus or the birth of weak lambs
which soon die (Miller, 1942). A shortage of water and green feed may
result in inadequate quantity and quality of milk for suckling lambs,
or tlie lack of proper feed for weaning lambs. We found two one-month
old lambs dead (aged after Demming. 1952).

Competition with adults for insufficient drinking water may work
to the detriment of the lambs, and prolonged searches for water up and
down dry canyons (which we found them doing) may tax the lambs
severely.

Other states have met the problem of reducing competition between
old and young sheep by holding periodic special hunts for a limited
number of mature rams. In 1954 Colorado allowed the taking of a few
surplus ewes also. Table 4 summarizes information on bighorn hunt-
ing seasons compiled by Dr. Helmut K. Buechner, University of Wash-
ington. For purposes of comparison it is estimated that there are 2,000
to 2,500 bighorn in California (Jones, 1955).

TABLE 4
Summary of Special Ram Hunts Held in Other States

State

Period

Number of
hunts

Average

estimated

populations

Average

number of

permits

Average
number of

rams
harvested

Arizona

Colorado -

Idaho

Montana

Nevada

1953-55

1953-55

1946-55t

1953-55

1952-54

1954-55

1905-55

4
3
6
3
3
2
32

3,025t
5,000 1
2.500
1,100
1,500
160
2,300

20
196
44
39
53
14
87

9
61*
16
26
14

New Mexico

9

Wyoming

36

* In 1954, 62 rams. 14 ewes, and 3 lambs were harvested.

t .Some hunts were held prior to 1946, but were not reported on.

t From U. S. Fish and Wildlife Service, 1956.

We collected remains of 10 carcasses and found old weathered bones
of many others in 1953. Two of the carcasses were lambs born in 1953
which died in the spring at about one month of age. Another was a
lamb of the previous year which died in the fall at about six months
of age. None of these appeared to be predator kills, since the carcasses
and bones were intact. A predator would have torn up the carcasses
and eaten most of the soft bones of the younger lambs. Two other car-
casses were six-month-old lambs that had died several years previously,
and five were adults. With the exception of one old ram found high
on a ridge, all carcasses were found near waterholes.

BIGHORN SHEEP SURVEY 187

Remains of various additional carcasses have been found at random
since the 1953 survey. Of particular interest was the location in April,
1955, of one adult ram that had been dead about six months to a year ;
and one lamb, three adult ewes, and two adult rams that had been dead
from one to two years around Intrigue Spring in the north end of
Borrego State Park. Three were along the water which runs below the
spring for 150 yards and the rest were within a half mile of the spring.
None of the bones was gnawed, as would be done by any predator, and
there was no evidence of gunfire. Apparently these animals had died
during the summer while concentrated around the water, which is the
only supply in the area.

WATER

These bighorn prefer fresh water to drink. We found foul or alkaline
water to be generally unpalatable to them, although they frequently
took it when slightly stagnant. "When springs are sanded over the sheep
will dig down with tlioir front hooves in an attempt to reach water.

Sheep often bed down at a spring and drink at any time during the
day. We observed them traveling to water most frequently early in the
morning and in the evening after sunset. One night about 3 a.m. one
was heard coming to a spring at which we were camped.

Even the most casual inspection of bighorn range in the Santa Rosas
brings out the fact that water, or lack of it, is a prime factor limiting
the numbers of animals. We therefore concentrated on locating and,
wherever possible, visiting every waterhole in the area surveyed. A
few additional springs have been located since the 1953 survey.

The existence of most springs depends upon rainfall. During years of
subnormal precipitation, such as 1953, many otherwise reliable springs
dry up. Canyons now dry may have water in other years. Cloudbursts
occur periodically in all of the canyons. There may be several during
one season or none for a decade. The effect on springs is unpredictable.
Some may be scoured out and made available ; others may be buried
under several feet of sand. Consequently, spring developments must be
constructed so as to have the least susceptibility to damage by cloud-
bursts. Summer rains are usually of such short duration, and the runoff
so rapid, that springs are not benefited.

Since water is a critical factor for bighorn, and for other game as
well, we strongly recommended in a preliminary report in 1953 that the
program of water development in this region be accelerated. We be-
lieved that moderate sums spent on additional development could well
raise the carrying capacity of the Santa Rosas many fold.

Weaver, Vernoy, and Craig have visited all of the water sites located
in the original survey, in addition to several located by themselves.
Each water source has been evaluated for possible improvement and
wherever practical to do so development work has been accomplished.

Results of this development work have varied due both to the vagaries
of the bighorn and the spring flow. As an example. Magnesia Spring
was used heavily in July, 1953, though the spring was diurnal, run-
ning only at night and drying up in the daytime. In March, 1954, a
tank was blasted out of a rock face below the spring. In July, 1954,
the spring itself was completely dry. though the tank was full. While
sheep were using it readily, they were far fewer than in the previous

188

CALIFORNIA FISH AND GAME

summer. In December, 1954, the tank was still full. In September, 1955,
it was nearly dry and no water was available for game.

A striking example of the benefit of water development in opening
np sheep range was afforded on Upper Dead Indian and Grapevine
creeks. In Jnly, 1953, it was found that sheep were not using the ex-
tensive area in the upper reaches of these canyons. Since no available
water was found, it was assumed that the absence of sheep was due to
the lack of water. High in each of these drainages a permanent
seep supplying only enough water for insects was found. In March,
1954, these seeps were developed to provide tanks of drinking water
by blasting and digging out existing sandfilled basins in the bedrock.
Bighorn accepted these immediately and in July, 1954, each tank was
receiving considerable use. In this case previously untenable summer
range was immediately transformed into suitable habitat by the de-
velopment of drinking water.

FOOD HABITS

Table 5 lists plants that we found to be eaten by bighorn. The plants
listed as very important are the ones supplying the bulk of the forage.
Use on forbs may well be of greater importance than our observations
revealed, for cropping on them is less obvious.

TABLE 5

Plants Observed Eaten by Bigtiorn in the Santa Rosa Mountain:

Common name

Scientific name

Importance

Horsetail

Equisetum sp. . . -

slight

Ephedra-

Ephedra sp. .- .- .

moderate

Cattail .

Typha laii folia . _.

slight

Grasses - - __-__-__

Unidentified (at springs)

Hilaria rigida . .

slight

Galleta arass

moderate

Palm... - -- - - -.

WashiTifftonia filifera

J uncus sp. - -.

slight

Riosh - . -

slight

Maguey .. -

slight

Stream orchis - _ .

Epipactis Qifjantea _

slight

Willow..

Salix sp

slight

Cottonwood -. - . -

sUght

Wild buckwheat- -

EriogonuTn sp. -

slight

Desert apricot .

slight

White ratany . _

Krajneria canescens .

high

Palo verde

Oercidium TnicrophylltiTn . .

sUght

Honey mesquite . . -

Prosopis chilensis. .. . . . ..

shght-moderate

Catclaw . ..

Acacia Greggii .

moderate

Littleleaf rushpea

slight

Silver bush

Argythamnia lanceolaia - . ..

moderate

Bernardia

Bernardia myricaefolia

Euphorbia sp.

Larrea divaricate .

moderate

Spurge . .-

unknown

Creosote bush. ....

slight

Goatnut .

Simmontisia chin^nsis

high

French tamarisk . .. .

Tamarix gallica

slight-moderate

Barrel cactus.. .. .

EchinocactuB cylindraceus

slight

Phacelia . ..

unknown

Black sage . .. . . .

Salvia mellifera

slight

Bee sage . .

Hypiis Efnoryi -- --

high

Groundcherry ..

slight

Desert-willow .

Chilopsis linearis

slight

Water-wally .

Rush bebbia

moderate

Incienso

Eficelia faritiosa

slight
slight

White burgage .

Franseria dumosa .

BIGHORN SHEEP SURVEY

189

Table 6 lists plants occurring in the Santa Rosas which we did not
observe used, but which have been found to be eaten by bighorn else-
where.

TABLE 6

Plants Occurring in the Santa Rosa Ivjountains Recorded as Bighorn Food Elsewhere

Common name

Scientific name

Locality

Reference

Fluffgrass

Triodia pulchella

Arizona _ .

Arizona

Unknown

Arizona

Arizona

Arizona

Arizona

Russo (1956)

Elepha nt tree --

Bursera odorata. .-

Russo (1956)

Rhus trilobata

Seton (1929)

Ocotillo

Cactus

Brickellbush- _ _ „

Fauquier ia splendens

Opuntia sp., MammtUaria sp

Srickellia californicd

Russo (1956)
Russo (1956)
Russo (1956)

Bush encelia

Encelia frutescens

Russo (1956)

In two areas there was evidence of overbrowsing on certain forage
plants. Goatnut was used heavily in a liaK-mile radius around the
upper Cat Creek spring and water-wally, rushes, and young palms
were taken heavily at the spring. Some of the use on the goatnut was
by rodents, but most of it was by sheep. Tamarisk was highlined, and
catclaw was browsed heavily in the wash above jMagnesia Spring. Plants
growing at several of the springs, such as cattail, horsetail, and various
grasses and forbs, were grazed extensively on occasion. Distribution of
these is limited to waterholes, where heavy use is to be expected.

DISEASES AND PARASITES

We were told of a large scale die-off in the early 1920 's. One man
stated that the bighorn lost their hair while still alive. He believed that
domestic sheep ranging from the valley floor had transmitted some
disease to the bighorn. The loss of hair could have been due to scabies,
which has been carried to bighorn by domestic sheep in other areas
to their great detriment.

"We observed one yearling sheep that had a violent coughing spell,
and a fawn deer that was barely able to walk, but we have no idea of
the causes.

Samples of 41 fresh pellet groups were collected in July, 1953, for
examination for eggs and larvae of internal parasites. The examination
was made by John Azevedo of the California Department of Fish and
Game Disease Laboratory, Project California W-35-R. The results were
as follows : No organisms were found in 36 ; tapeworm eggs of the
species Monezia were found in one; nematode eggs of the species
Nematodirus were found in two, of the species Gongylonema in one,
and an unidentified nematode larva in another. The occurrence of eggs
and larvae was much lower than Azevedo has found in other herds of
bighorn in California. In June, 1955, five additional samples were
sent to Dr. Erling R. Quortrup, Director of the San Diego County
Livestock Department, to be examined for lungworm larvae. He found
lungworm larvae in two and strongyle ova in three.

190 CALIFORNIA FISH AND GAME

INTERRELATIONSHIPS WITH OTHER ANIMALS
Range Competition

Domestic stock are present in tlie higher elevations, generally above
the bighorn range. About 20 wild horses range around Asbestos Moun-
tain. These apparently do not range lower than the upper limit of the
bighorn winter range. An undetermined number of cattle range in the
brush above the bighorn range.

About 30 domestic goats live in Tahquitz and the adjoining canyons
that drain into Martinez. As they are in bighorn range, have sim-
ilar feeding habits, and heavily use the area, it appears that they
adversely affect the bighorn. It is said that other goats that have been
wild for years run on the bighorn range, though we did not see any.

A few domestic burros are present in Martinez Canyon. They use the
waterholes there and in the lower portions of other incoming canyons.
It is possible that their presence restricts bighorn use.

Deer are present in small numbers above the bighorn range. They
adhere to the chaparral, generally above 4,000 feet. We observed three
during our search for sheep, all at high elevations. We found some old
deer sign and a few shed antlers as low as 2,400 feet, indicating winter
range limits. Deer numbers are said to be far less than they once were.
Competition with bighorn is very slight. At some of the higher springs
on the north facing slopes, such as in Boulder Camp Canyon, deer and
sheep are using the same springs.

Antelope ground squirrels occur throughout the sheep range and
take considerable forage in some areas. Use on juniper was noted espe-
cially, and on goatnut around the upper Cat Creek Spring.

Predation

We observed no positive indications of predation on bighorn,
although some undoubtedly occurs. Mountain lions are present to the
north on Mt. San Jacinto, but seldom come into the Santa Rosas. One
was reported by U. S. Army surveying personnel in February, 1955, at
2,000 feet between Martinez and Toro canyons. This is in the middle
of the sheep range. Jack Miller observed a large shepherd-dog-like
animal, which he believed to be a wolf, chasing his goats in Tahquitz
Canyon some years ago. Bobcats are common in the chaparral, but
are of rare occurrence in the bighorn habitat. We saw considerable sign
at the head of Deep Canyon on the Cactus Spring Trail (at 4,000 feet),
while Miller has seen only one at his cabin in Martinez Canyon (at
3,000 feet) in 25 years.

Coyotes are common in the sheep habitat. We observed six during
our survey and found abundant sign. Although coyotes have been
known to kill adult bighorn (Russo, 1956), their depredations are gen-
erally directed at newborn lambs.

Poaching

Poaching constitutes a drain on the bighorn population, although the
extent of it is difficult to assess. Dismembered carcasses along with
cartridge cases and other evidence of illegal hunting are encountered
occasionally.

BIGHORN SHEEP SURVEY 191

Continuing patrol effort throughout the bighorn range is necessary to
keep poaching at a minimum.

SUMMARY

A survey of bighorn populations in the Santa Rosa Mountains was
conducted during July, 1953. Additional information was gathered
through periodic visits to the area through 1955.

Bighorn range on the lower slopes, generally below the 4,000 feet
contour. The estimated population for the area was set at 351.

The ratio of rams to ewes was 109 :100, of lambs to ewes 18:100, and
of yearlings to ewes 27 :100. Two thirds of the observed rams had a
three-quarter curl or better.

It is believed that availability of water is the most important limit-
ing factor in bighorn distribution in this area, since bighorn are
restricted to the immediate vicinity of springs during the long dry
period.

Forage plants in the vicinity of water were found to be heavily
utilized.

REFERENCES
Deming, O. V.

1952. Tooth development of the Nelson bighorn sheep. Calif. Fish and Game,
vol. 38, no. 4, p. 52.3-529.
Di.xon, Joseph S., and E. Lowell Sumner, Jr.

1939. A survey of desert bighorn in Death Vallev National Monument, summer
1938. Calif. Fish and Game, vol. 25, no. 2, p. 72-95.
Jones, Fred L.

1950. A survey of the Sierra Nevada bighorn. Sierra Club Bull., vol. 35, no. 6,
p. 29-76.

1955. California bighorn inventory. Calif. Dept. of Fish and Game. (Unpub-
lished report)

.Miller, R. F., G. H. Hart, and H. H. Cole.

1942. Fertility in sheep as affected by nutrition during liio hreeiling season and
pregnancy. Univ. of Calif., Agric. Exp. Sta., Bull. 672, 31 p.
Russo. John P.

1956. The desert bighorn sheep in Arizona. Ariz. Game and Fish Dept., 15;; p.
Seton, Ernest Thompson

1929. Lives of game animals. New York, Doubleday, Doran & Co., Inc., vol. 3,
part 2, p. 410-780.
U. S. Fish and Wildlife Service

1956. Inventory of big-game animals of the United States, 1954. Wildl. Leaflet
376, 3 p.
Welles, Ralph

1955. Report of the 1955 summer survey of bighorn in Death Valley National
Monument. Calif. Dept. of Fish and Game. (Unpublished report)

AN IMPROVED DEVICE FOR CAPTURING DEER'

GORDON ASHCRAFT and DON REESE

Game Management Branch

California Department of Fish and Game

INTRODUCTION

Big game trapping and tagging programs liave always exhibited a
(•ommon weakness: the inability to capture the animals efficiently and
economic-ally. Field men are constantly experimenting with new de-
vices and techiii(|ues in an eff<n't to improve efficiency. Factors to be
considered are portability, adaptability, safety, cost of materials, and
simplicity of operation. The "Improved Deer Snare" has proved very
adaptable to tlie various site factors and situations that present them-
selves in most trapping programs.

CONSTRUCTION

A detailed description of the fabrication of the various parts follows.
Cut the rubber used to motivate the device and to anchor the safety
rope from discarded truck tubes. Cut, fold, and tie two strips approxi-
mately 6 to 10 feet long and 6 to 8 inches wide. Lash back the material
with a modified sheet bend at either end, to facilitate the attachment
to the anchor point and nooses. Tie a sliding loop in either end of a
25-foot length of quarter-inch green nylon rope. Procure additional
rope, about 30 feet, for use as linkage between the rubber bands and
the snare loops. This rope will serve as a secondary static line, for
additional strength.

Cut the heads from two 20-penny nails. Score a third nail lightly
near the head with the cutting portion of lineman's pliers and attacli
one end of the leader material very securely in the notch thus created.

TABLE 1
List of Materials Used in Making the Deer Snare *

Item

Materials

Approximate
cost

Discarded truck tire tubes- -

Rope - >-

SLxty feet of 3/16-inch diameter O.D. Nylon

Three 20-penny common nails

Three yards of 6-lb. test nylon fish leader

One N^o. 10 cow or sheep bell -_

$1.80

Nails- -- - - - --

.05

.06

Bell

.50

Total cost

S2.41

Tools needed for setting trap are as follows: hammer, axe, machete, lineman's pliers, and tree climbers (op-
tional).

1 Submitted for publication June, 1956.

( 193 )

194

CALIFORXIA FISH AXD GAME

This will serve as the tripper. Details of construction are shown in
Figures 1, 2, 3, and 4. Table 1 is a list of materials needed for con-
struction.

Safety line

Cow or sheep be

Tree or post

Trigger Rubber

Safety Rubber

Trigger holding nails

Trigger nail

Trigger trip string

Snare loop

FIGURE 1. "Improved Deer Snore", showing details of construction. Drowing by Cliffa Corson.

DEEE CAPTURING DEVICE

395

OPERATION

To make a typical setting, select a spot having a tree or post two
feet or less from the trail or well-used passage. Secure one end of the
actuating rubber at a height of approximately 15 feet. Drive the head-
less nails into the tree (or post) approximately 12 inches from the
ground on the trail side. Space the nails approximately one inch apart
in a horizontal plane and leave 1^ inches protruding. Tie one end of
the extra rope (static line) to the lower end of the actuating rubber,
stretch the rubber to the maximum, and attach the trigger nail to the
rope at the level of the two headless nails. After engaging the trigger

FIGURE 2. Typical snare set. Photograph by William Aumai

196

CALIFORNIA FISH AND GAME

FIGURE 3. Snare trigger mechanism. Photograph by William Auman.

nail below these stub nails, pull the rope gently to the trail, maintain-
ing the tension on the rubber. Attach to the exact center of the snare
rope, enabling the loops of the snare to be spread fully in the trail,
without any slack in the line. Pass the free end of the static line
through the loop of a small bell (as used on sheep) and secure through
two loops of rubber near the butt of the tree, encircling the trunk.
Two or three feet of slack is allowable in this line.

Arrange the trigger nail to allow" easy extraction from its position
beneath the two stub nails and tie the free end of the leader material
to some substantial object on the opposite side of the trail, at approxi-
mately the same height as the nails. This constitutes the trip string and
should be taut at all times. Use a tree, bush, or rock if possible, since
the animals hesitate to pass a stake or freshly cut limb ; test the action

DEER CAPTURING DEVICE

197

FIGURE 4. Variation of the typical deer snore, using a fulcrum pole in connection with a tree

site setting. Photograph by William Auman.

of the device, reset, spread the loops to their maximum in the trail on
either side of the trip string, and cover them lightly with dust.

As stated above, the "Improved Deer Snare" is highly adaptable.
A small amount of imagination and ingenuity will enable the operator
to make sets in nearly any situation. One variation, representing the
key to success in treeless areas, is illustrated in Figure 5. The same
components — rope, truck tubes, and nails — are used. The nails con-
stituting the trigger mechanism are driven into a large stake driven
into the ground near the trail. This peg should not protrude more than
14 inches and should be well hidden or camouflaged. A light, stiff pole
approximately six feet long is used to supply the upward motion needed
for raising the nooses around the animal's legs. The pole is butted
against a stake driven firmly into the ground as the pole lies at right
angles to the trail. This stake should protrude approximately two feet
above the ground to provide fulcrum for both pole and rubber. The
pole is held down by the rope and trigger arrangement at the end
nearest the trail and prevented from moving away from the trail by the
fulcrum. The rubber is attached to the pole at midpoint, stretched to
the maximum over the top of the fulcrum and secured to a post placed
directly in line at the proper distance behind said pivot point. Lateral
motion of the pole is prevented by a slender guide stake on either side
at midpoint. The static line is threaded through a bell, tied to the

198

CALIFORNIA FISH AND GAME

trail-eiifl of the pole, affixed to the trigger nail, and tied to the noose
rope, Avitliout slack. This line is linked to the rearmost stake by the
second rubber band.

The objective is to capture an animal by snaring one or two legs
in a loop of rope. Considerable experimentation indicates that there

FIGURE 5. Snare set at a treeless site. Photograph by William Auman.

is less possibility of injuring an animal or allowing it to escape if it
is held by the legs rather than the neck.

Once caught, the animal is prevented from applying full impact to
the restraining ropes by the resilience of the rubber which activates
the device. A bell attached to the restraining rope signals the operator,
who remains in the vicinity, servicing a number of snares. If the snares
are used in a belling program the snare bell should be of different
tonal qualit}^ than the deer bells, in order to avoid confusion.

The snared animal, if approached cautiously, keeping the anchor
point between the operator and the animal, will usually throw itself.
Care should be exercised to prevent the accumulation of slack line,
which the animal would use to develop momentum, resulting in shock
to the animal and the device. Once the deer has thrown itself, or has
been pulled to the ground, it is possible for the operator to crouch at
the animal's back, placing one knee on its neck, and bind its feet. This
immobilizes the deer for purposes of examination or marking.

DEER CAPTURING DEVICE 199

Several sets may be made by one man in a day's time, if necessary,
though a two-man team can work more efficiently. A team of operators
can set out from 10 to 20 of these snares, depending upon the number
of suitable locations presented within a radius which is determined by
the distance the snare bell can be heard. As a deer trips the snare and
becomes caught in the loops, the activity causes the bell attached to
the static line to set up a clamor. The possibility of injury to the animal
or escape increase in proportion to the amount of time it remains in
the trap. Therefore, the operators should remain within earshot of all
the operating snares.

The possibility of injury to snared animals is slight if the devices
are closely attended. The authors snared a total of 62 deer during the
period August 2 to September 1, 1955, (35 man days in actual opera-
tion) without injuring an animal.

The snare is partieulai-ly adapted to use on well-traveled trails lead-
ing to waterholes, salt licks, bedding grounds, and foraging areas.
Baiting is sometimes helpful in attracting tlie deer to a certain spot,
although usually this procedure is not necessary for successful opera-
tion.

THE DEVELOPMENT OF A VINYL PLASTIC
SUBCUTANEOUS TAG FOR TROUT'

ROBERT L. BUTLER

Inland Fisheries Branch

California Department of Fish and Game

INTRODUCTION

California has one of the largest catehable rainbow trout {Salmo
gairdneri) programs in the United States. Approximately eight million
such trout are planted each year by the California Department of Fish
and Game at a cost exceeding one million dollars. These trout, weighing
from four to six to the pound, are planted in readily accessible streams
and lakes having high angling pressure. The policy followed by the
Department is to plant only water from which a return in excess of 50
percent of the planted fish may be expected to reach the angler's creel.

The magnitude and costs of the catehable trout program are so great
that it is imi)ortant to make the best possible use of the hatchery prod-
uct. A special project was begun in 1955 to develop better methods for
evaluating the results of catehable stocking under various conditions,
with a view to improving the program.

The great majority of planted waters have multiple access points, if
not continuous roadside contact. The difficulty of censusing such waters
is readily apparent, since it is virtually impossible to obtain a complete
creel census or to obtain truly random samples.

Since dependence had to be placed in large part ou voluntary returns,
a marking program involving the removal of certain fins also could not
have been expected to yield adequate results. The majority of fishermen
seldom recognize the absence of fins and less frequently report such
marks.

A new approach was therefore sought. The most promising one in-
volved estimating the percentage of recapture through voluntary
returns of numbered tags by anglers, supplemented by creel census.

Any tagging study involving large numbers of fish requires a tag that
is easy to apply, is readily seen by the angler, and has no adverse
effect on the fish.

The jaw tag (Kounsefell and Everhart, 1953, p. 279) has been widely
used on catehable trout, but has revealed serious weaknesses. For exam-
ple, in various studies tag loss has been demonstrated, tissue destruction
has been apparent, and growth has been impaired. There was no ques-
tion but that time spent in developing some other tag for catehable trout
would be time well spent.

1 Submitted for publication March, 1957. This work was performed as part of Dingell-
Johnson Project California F-14-R, "Evaluation of Catehable Trout Stocking",
supported by Federal Aid to Fish Restoration funds.

(201)

202 CALIFORNIA I'^ISII AND GAME

A series of tests with various tajrs was carried out at three different
hatcheries and results and modifications of these tests were made simul-
taneously in the field. The Eipper tag, a thin arrowhead shaped metal
piece to "which is attached a round dangler, gave a very high tag loss
in the first tests. Heincke stud tags prepared from acrylic plastic were
tried in tAvo positions on the trout. Retention in a dorsal position was
good over a period of 185 days ; however, an unsightly wound developed
around the tag. It is possible that modifications of these or other types
of tags would have yielded satisfactory results in the end, but prelimi-
nary experiments with vinyl plastic subcutaneous tags looked promising,
so a major effort was made to develop this tag for use on catchable
trout.

Metal subcutaneous tags have been used on livestock for several j-ears.
A special tool has been devised for inserting these tags in the ears of
cattle and sheep. The first known subcutaneous tagging on fish was done
by Le Cren (1954). His tagging of char involved placing metal tags in
locations near the dorsal fin. Although the incision at the point of tag
insertion was visible and the tag could be felt beneath the skin, it could
not be seen through the pigmentation of the back. An electronic tag
detector device was developed for application with the above tag (Moore
and Mortimer, 1954). Bertelsen ^ of Denmark has been using plastic
subcutaneous tags on plaice. Of 1,150 young plaice tattooed for detec-
tion and tagged for identification, 438 were reported returned, with
only two of these showing tag loss.

The belly was chosen as the best location for the subcutaneous tag on
trout. There is little if any black pigmentation here to obscure the tag
and through careful tagging procedures the tag can be placed above
the white guanine pigment layer (Figure 1). Inasmuch as most sports-
men turn the fish belly up in cleaning, the likelihood of their seeing the

Personal communication. Dr. E. Bertelsen, Danmark.s Fiskeri-og- Harunders</>g-elser,
Charlottenlund Slot, Charlottenlund, Denmark.

xJS^Bk '

mttk

FIGURE 1. The location and appearance of a vinyl tag recently placed beneath the belly skin
of a rainbow trout. Photograph by Edwin P. Pisfer, January, 1957.

SUBCUTANEOUS TAG FOR TROUT 203

tag in this position is greatly increased. This section of the body experi-
ences minimum body movement and therefore irritation due to body
movement is minimal. Induced mortality through blood loss or inter-
ference with vital organs may be expected to be minimal because of the
fat deposits and lowered blood supply to this part of the body. It was
found that this tag location permits ease of handling fish during the
tagging operation.

TAG DESCRIPTION AND PLACEMENT

The tags are prepared from soft upholsterer's yellow vinyl plastic.^
The tag is 2^ to 3 mm. wide, 20 mm. long, and 0.018 inch thick. It is
believed that a thickness between 10 and 15 thousandths would be more
desirable. The tag is rounded at both ends and care is used in the manu-
facture to avoid am' ragged ends or sharp projections.

All tags have been marked with Vinyl Stamping Black number
104X5A4 of the California Ink Company, San Francisco, California.
The tags have a number plus the word "Return" on one side and on
the opposite side is printed "C. F. G. Sacto." to indicate return to the
California Department of Fish and Game, Sacramento. Two studies
have used reward tags and these have been prepared with the words
"Reward $5" on one side instead of the number. During the first year of
field application several tliousand tags were prepared by hand. It is
now possible to obtain these tags from Howitt Plastics Company, Route
2, Box 61-B, Molalla, Oregon.

The principal tool for tagging is ])repared from a Bard and Parker
No. 7 scalpel handle fitted with a No. 12 blade (Figure 2A). These can
be obtained from any surgical supply house. The handle is filed down to
a flat blade 1 mm. thick, 4 mm. wide, and approximately 30 mm. long
(Figure 2B). The handle is then bent to form a U-shape (Figure 2C).
The end opposite the No. 12 blade, which is used to form the tag pocket,
is finely sharpened on the outer surface. The tool is then butfed to a
high polish, so as to permit a smooth and rapid operation of tag pocket
formation.

Subcutaneous tagging would be quite difficult without placing the
fish under anesthesia. Trieaine methanesulfonate (M.S. 222) is being
used very effectively for this purpose. A concentration of ^ gram/gallon
at 50 degrees F. prepares fish for the operation within 30 seconds.
Four to five fish are anesthetized at a time. As the temperature increases
it is necessary to reduce the number of fish exposed to this concentra-
tion for a given time or else reduce the concentration to something less
than one-half gram per gallon. At this concentration fish should not be
exposed to the solution for periods in excess of three minutes. At lower
concentrations or lower temperatures trout may be held longer without
harm. Each situation reciuires some experimentation and adjustment.
Chlorobutanol has been tried and, although very effective in relaxing
the fish for some hatchery operations, such as spawning, excessive move-
ment occurs during an operation of this type.

3 The plastic was obtained from three different manufacturers. In each case its exact
physical properties and chemical composition could not be obtained. As these prop-
erties may influence retention, toxicity, etc., it is necessary to test these qualities
for each lot of plastic used.

204

CAIJFORNIA FISH AND GAME

C.E.C.

FIGURE 2. The steps in forming the subcutaneous tagging tool: A, Bard and Parker No. 7
handle for a No. 12 blade; B, The handle is filed down to a flat blade 1 mm. thick, 4 mm.
wide, and approximately 30 mm. long; C, The handle bent into a U-shope with the end oppo-
site the blade sharpened on the outer surface. Drawings by Cliffa Corson.

A narrow incision perpendicular to the body axis and not exceeding
4 mm. in length is made left of the sagittal line and slightly anterior
of the ventral fins (Figure 3A). In making the incision it is necessary
that the No. 12 blade be held nearly flat, not exceeding 30 degrees in
elevation from the body axis. Possibilities of cutting the musculature
are thereby greatly reduced and the resulting tapered incision seems
to facilitate more rapid healing than could be obtained with a perpen-
dicular cut.

The pocket for the tag is started by carefully inserting the opposite
end of the tagging tool (Figure SB), followed by pressing forward
and gently upward simultaneously (Figure 3C). Care must be exer-
cised to avoid bruising the tissues or cutting through the skin. If the
tool is properly sharpened and bulfed the white pigment is cleanly
separated from the outer skin layer with moderate upward pressure.
The tag is picked up with flattened forceps, immersed in isopropyl
alcohol, and immediately pushed to the forward end of the pocket
(Figures 3D and E). Any excess alcohol is not removed from the tag.

If the incision is made as above certainly less than one percent of
the fish tagged will show any signs of bleeding. Bleeding generally
indicates improper placement, poor incision, and/or poor pocket prep-
aration. It was found that incisions made on the right-hand side of
the belly generally cut blood vessels. No check was made to determine
what effect tagging on the right-hand side of the body would have on
mortality.

Although the above procedure may seem tedious and time consum-
ing, it was found that within half a day of tagging all field men were
capable of tagging one fish per minute. "With good tools and a eon-

SUBCUTANEOUS TAG FOR TROUT

205

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206

CALIFORNIA FISH AND GAME

venient working situation 100 tags per hour may be considered opti-
mum tagging speed. Chi square analysis of tag returns from various
taggers has indicated the technique is effectively learned in a short
time. Significant differences in numbers of tags returned from specific
lots of trout tagged by different individuals could seldom be demon-
strated,

TAG RETENTION

The initial experiments with a broad tag (5 x 15 mm.) were carried
out by making tag placement in two different manners. A longitudinal
placement, similar to that now used, was made. A second placement was
made by locating the tag transversely across the body in the same area.
In the transverse tagging half of the tag lay on one side of the incision
and the other half of the tag was fitted into a pocket on the opposite
side of the incision. After 135 days all transverse tags were lost. Tags
in the longitudinal position showed a 24 percent tag loss with only 30
percent good retention. Most of those retained at the end of 69 days
were poor (Table 1).^

TABLE 1

The Quality of Subcutaneous Tag Retention With Different Tag Sizes/-'
Placement, and Treatment of the Incisions

Troatment'of
incision

Num-
ber of
fish at
end'of
study

Tag retention quality

Lengtli

of

study

in days

Tag width and placement

Good

Fair

Poor

Lost

Broad (longitudinal)

Broad Ttransverse)

Broad (longitudinal)

Narrow (longitudinal

Narrow (longitudinal)

Nothing

Nothing

Alcohol

Alcohol and terra-
mycin

Alcohol and sulfa-
merazine

.50
19
10

9

10

15

2

10

8

10

7

1

16
1

1

12

15

0

0

0

69
69
61

61

61

* Broad tags (j mm.) ; narrow tags (3 mm.).

Field application of the broad longitudinal tag was made in Castle
Lake in 1955. Tag loss increased during the first 69 days (Table 2).
With a tag loss of 57.8 percent during the 42-69 day period, it was
obvious that such a tag would not be very desirable.

From the first experimental work, early tag loss was found to occur
through the open incision. Tag losses occurring after the second week
Avere related for the most part to blister formation. The blistered tissue
became weakened through what appeared to be poor circulation, fol-
lowed by necrosis and tag loss. It was believed at the time that blister-
ing was caused through one or more of the following factors: (a)
bacterial infection introduced with the tag and/or tagging tools; (b)
toxicity of the tag itself; (c) irritation induced by the stiffness of the

* The terms good, fair, and poor are descriptive of the state of tissues enclosing the
tag. Good retention : the skin covering the tag is normal in appearance. Healing
has been complete. Fair retention: tissue repair at the incision has been complete:
however, the skin covering the tag is raised in a blister. Poor retention: the inci-
sion has not healed, the tag is partly extruded, and or the skin covering the tag is
necrotic. Tag loss soon follows this condition.

SUBCUTANEOUS TAG FOR TROUT

207

TABLE 2

The One-season Returns From 980 Marked and Tagged Fall-spawned Rainbow Trout
Planted in Castle Lake, June 15, 1955 *

>i umber of days after planting

Tr»tal«

0-13

14-27

28-41

42-69

70-140

Tas lost

8

(6.7)

112
(93.3)

44
(30.3)

101
(69.7)

62
(40.. 5)

91
(59.5)

37

(57.8)

27
(42.2)

17
(45.9)

20
(54.0)

168

Tag retained

(32.4)

351
(67.6)

Totals

120

145

153

64

37

519

* Numbers in parentheses are percentages based upon the total returns within each time series.

taj?; (d) blocking- of circulation ihrough the covering tissues by the
width of the tag.

The contents of the typical blister were examined from two blistered
tag areas, Practically no bacteria were found. In examining the lateral
and deeper tissues around the tag incision, there was no obvious tissue
infection. Inflammation was local. In these studies two groups of fish,
longitudinal control and transverse control, were included. Both groups
underwent anesthetization, incision, and tag pocket formation. The
only difference between these control groups and the tagged groups was
simply that the control groups were without tags. None of the controls
developed blistering or showed any signs of irritation. Complete healing
was noted within two weeks.

Simultaneously with the above work on the bacterial aspects, two-
year-old brood stock were tagged at one of the hatcheries. The plastic
tags used were covered with a three percent aureomycin ointment.
Blisters occurred in almost every instance and if bacteria were causa-
tive in blister formation aureomycin was ineifective. Tags on this group
of 60 brood stock fish, although slightly larger (6 x 17 mm.), were well
retained two years later in 8 out of 10 fish examined March, 1957
(Figure 4).

Tests were run to determine whether or not blister formation was the
result of toxicity and/or tag irritation. To check these factors it was
necessary to obtain a soft material of known low toxicity from which
tags could be made. The closer the flexibility of the material could ap-
proach that of tissue the less likelihood there would be of irritation.
Various silicone products meet these requirements. They have a very
low order of toxicity (Rowe, Spencer, and Bass, 1948), and are substan-
tially without effect on the human body (McGregor, 1953). As a group
they maintain a very pliable state over a range of temperature far
beyond the temperature range of water.

A silicone, ''Silastic" S-97-11, was obtained from the Dow Corning
Company for this test. Individual fish were tagged with "Silastic"
and w'ith the vinyl plastic subcutaneous tag. Observations made 19 days
later showed typical blistering around both tags. Blistering, therefore,
was not due to toxicity and if due to irritation this factor had been
reduced as much as possible.

208

CALIFORNIA FISH AND GAME

FIGURE 4. The presence of a two-year-old tag indicated by pigment deposition prior to its
removal. Phofograph by George Bruley, February, 1957.

The last aspect of experimentation concerned itself with broad
(5 mm.) vs. narrow (3 mm.) tags. During field work several hundred
fish were tagged using two tag sizes : 3 x 20 mm. and 5 x 15 mm. Ran-
dom observations of these fish 15 days later at the hatchery, just prior
to planting, indicated the narrow tag to be superior to the broad tag
(Table 3). Four tags had been lost from the broad tagged group and

TABLE 3

The Quality of Subcutaneous Tag Retention as Determined From Random Observations of
Tagged Fisti Held for 15 Days Prior to Planting

Tag size in mm.

Tag retention quality

Good

Fair

Poor

Lost

Narrow (3 x 20).
Broad (5 x 1.5)..

10
14

3
16

there were no observed losses in the narrow tagged group. Chi square
analysis of tag returns in these two groups six months later demon-
strated a significant difference between the returns of narrow and broad
tags. The narrow tags returned at a significantly higher rate.

SUBCUTANEOUS TAG FOR TROUT

209

At the time the third hatchery experiment was begun, conclusions
concerning bacterial infection had not been established. A series of
tests was made, using both broad and narrow tags, in conjunction with
specific treatment for the incision. Three types of treatment were made :
isopropyl alcohol alone, alcohol and terramycin, and alcohol and sul-
famerazine. In each instance the tag was dipped in alcohol and those
receiving terramycin or sulfamerazine were then covered with either
of these substances. Subsequent observations made on the third and
seventh week and the last day of the experiment (Table 1) indicated
that alcohol alone was effective in greatly improving tag retention.
Tags in each instance were not accompanied by blistering and most
of them could be verv cloarlv read through the skin. There was no
sign of irritation. The alcohol may hasten the healing and formation
of adhesions around the tag and incision.

MORTALITY

Values for the effects of tags on mortality have been obtained from
tagged and untagged fish held in hatchery raceways. Those trout with
longitudinal tags showed only one dead fish out of 56 for a period of
69 days, compared with two dead fish from the longitudinal control
group (Table 4). There is no way of knowing what happened to the
missing fish. Loss may have been tlirough prcdation or escapment into
other sections of the raceway. The transverse tag group had one death,

TA3LE 4
The Comparative Mortality of Tagged and Untagged Trout for a Period of 69 Days

Type of tagging and control

rNumber of fish

Missing

Known dead

Longitudinal tag

Longitudinal control

Transverse tag

Transverse control-.
Unmarked control-,.
Fin clip control

56
56
23
23
5
15

compared with no deaths in the transverse control group. One death
was observed in the fin clipped control group.

It has been observed consistently that rainbow trout feed readily
after tagging. As soon as they regain their equilibrium following an-
esthetization they are very active. Short-term observations of mortality
have been made in the various hatcheries where the tagged fish have
been held a few days prior to planting. From 1,200 fish tagged for one
plant, one loss was observed during the three days prior to planting.
This loss occurred during the actual tagging operation. Of the 980
tagged at Castle Lake, three losses occurred during tagging. No addi-
tional losses were noted five days later when planted.

It seems that the initial mortality from tagging is related directly
to the tagging procedures and it has been observed that if temperatures
of the anesthetic solution are appreciably higher than the hatchery

210

CALIFORNIA FISH AND GAME

water, they may adversely affect the fish. It is important that the tem-
perature of the anesthetic solution be maintained as near as possible
to that of the hatchery water. At the recommended concentration of
^ gram/gallon at 50 degrees F. too long au exposure to the solution
may cause death. Once gill movement ceases completely care must be
used to return the fish as rapidly as possible to the hatchery water. The
higher the concentration of M.8. 222, tlie higher the temperature, and
the longer the exposure to the anesthetic, the greater danger there is

of losing fish.

GROWTH

Inasmuch as most of the returns from catchable trout are obtained
in the first two or three weeks after planting, there is little opportunity
to determine the effect of the tag on growth after planting. Values that
liave been obtained were taken from fish in the hatchery experiments.
The mean growth increment is that recorded for all the fish at the
conclusion of each experiment (Table 5). In the 135-day test the mean
growth increment was 1.8 inches and the range was from 1.4 to 2.3

TABLE 5
The Growth of Longitudinally Tagged and Untagged Rainbow Trout From Two Hatchery Experiments

Type of tag or control

Broad yellow vinyl..
No incision control-.

Incision control

Narrow yellow vinyl
Broad yellow vinyl..

Number of fish
at end of study

12*

50

50

10

15

Mean growth
increment

1 . 8 inches
1 . 8 inches

1 . 6 inches
1 . 8 inches

1 . 7 inches

Length of study
in days

135
61
61
01
61

* Poaching following 69 days of protected retention greatly reduced the original number of 56 trout.

inches. In the Gl-da}" test, in which controls were run with the narrow
and broad tagged groups, there was no striking difference in the mean
growth rates, although fish with the broad tags had a much wider range,
0.7 to 2.6 inches.

Growth obtained from four-year-old brood stock tagged two years
])revious]y was all positive. These fish, originally 14^ to 16 inches long,
showed growth increments from 2| to 5 inches for the two-year period.

It may be concluded that in the hatchery, growth is not hampered
Ijy tlie tag nor by the incision. Inasmuch as these values were obtained
under hatchery conditions, they may not be applied with certainty to
lake or stream conditions. Conclusive testing should be done if a long-
term tag return is expected.

APPLICATION TO FIELD PROBLEMS

During the past two years the vinyl subcutaneous tag has been used
on groups of trout planted in sevei'al different tj'pes of waters. The
publicity given to these studies has varied from virtually none to very
intense. The fact that greater numbers have been returned voluntarily
than have been checked or picked up by creel checkers indicates that
the tag is readily seen.

SUBCUTANEOUS TAG FOR TROUT 211

Studies on catchable trout using this tag will be published upon at
a later date.

DISCUSSION

The subcutaneous tag is demonstrating its usefulness in the catch-
able trout program in California. The tag is readily seen by sportsmen.

Tag loss has been shown not to be a serious consideration in the high
angling intensity waters. Since 80 to 90 percent of the trout to be taken
are removed within a period of two to three weeks, definitive measures
of long-term tag loss cannot be obtained.

The tagging method developed is yet to be used in a long-term tag-
ging study.

Desirable qualities which have been demonstrated for this particular
type of tag are: (1) the tag does not affect growth rate of rainbow
trout under hatchery conditions and may be expected not to hinder
growth under lake and stream conditions; (2) the tag and the tagging
procedures described are not of themselves effective in increasing
mortality; (3) fish may grow without outgrowing the size of the tag;
(4) the tag will not catch upon any object in the stream or lake; there-
fore, it is particularly valuable where tag loss may be expected through
netting or contact with heavv brush.

'■fe

ACKNOWLEDGMENTS

One of the problems of inserting vinyl plastic tags was overcome
through the valuable suggestion offered by E. D. Le Cren of the Fresh-
water Biological Association, Ambleside, Westmoreland, England. He
suggested the use of a philatelist's tool for tag pocket formation. The
idea was adapted to the modification of the Bard and Parker blade and
handle. The interest shown by C. S. Kabel of the California Department
of Fish and Game was productive through his suggestion of using the
Bard and Parker surgical instrument. J. H. "Wales of the California
Department of Fish and Game kindly cared for one group of experi-
mental fish and examined the blisters found on tagged fish for
bacterial infection. His interest and active participation in this respect
and at Castle Lake were gratefully received. Sincere appreciation is
expressed to E. Bertelsen for the encouragement and presentation of
unpublished data concerning vinyl plastic tag application on plaice.

The chi-square analysis on narrow vs. broad tags made by H. K.
Chadwick of the California Department of Fish and Game was valuable
in emphasizing a more careful examination of narrow tag application.

Without the pleasant and very cooperative assistance of California
Fish and Game hatchery foremen A. J. WeaA'er, L. C. Hume, and R.
Bliss, the experimental hatchery work would have been virtually im-
possible.

SUMMARY

A subcutaneous vinyl plastic tag has been developed and is in current
use in California. The tools and method of tagging are described and
it has been demonstrated that the technique can be effectively learned
in a relatively short time. Optimum tagging rate may be expected to
be near 100 tags per hour.

212 CALIFORNIA FISH AND GAME

Hatchery experimentation and field observations have demonstrated
the following: (1) blistering ma}^ be the most serious problem related
to tag- retention; (2) blistering is not due to vinyl plastic toxicity, bac-
terial infection from the tools and/or the tag, or tag irritation; (3)
differences in mortality between tagged and untagged fish could not be
demonstrated; (4) the growth observed with various groups of hatch-
ery fisli was not hampered by tag application; (5) the experimenta-
tion has demonstrated that of the various methods and materials used
the narrow tag plus isopropyl alcohol provides the best tag retention;
(6) the tag can be seen b}- fishermen. Greater numbers are returned
voluntarily than are checked or picked up by creel checkers.

No long-term tests in lakes and streams have yet been conducted
to determine how effective the tag is for such fisheries.

REFERENCES
Le Cren, E. D.

1954. A subcutaneous tag for fish. Jour, du Conseil. vol. 20, no. 1, p. 72-S2.
McGregor, R. R.

1953. Silicones in pharmacy. Pharm. Internat., vol. 7, p. 24-26, 63.
Moore, AV. H., and C. H. Mortimer

1954. A portable instrument for the location of subcutaneous fish-tags. Jour, du
Conseil, vol. 20, no. 1, p. 83-86.

Rounsefell, George A., and W. Harry Everhart

1953. Fishery science : Its methods and applications. New York, John Wiley &
Sons, Inc., xii -f- 444 p.
Rowe, V. K., H. C. Spencer, and S. L. Bass

1948. Toxicological studies on certain commercial silicones and hydrolyzable
silane intermediates. Jour. Indust. Hvgiene and Toxicologv, vol. 30, no. 6,
p. 332-352.

CLASSIFICATION, VARIATION, AND DISTRIBUTION OF
THE SCULPINS, GENUS COTTUS, INHABITING PACIFIC
SLOPE WATERS IN CALIFORNIA AND SOUTHERN
OREGON, WITH A KEY TO THE SPECIES'

C. RICHARD ROBINS

The Marine Laboratory, University of Miami

Coral Gables, Florida

and

ROBERT RUSH MILLER

Museum of Zoology, University of Michigan

Ann Arbor, Michigan

TABLE OF CONTENTS

INTRODUCTION 213

METHODS 214

KEY TO SPECIES OF COTTL S OF THE PACIFIC SLOPE OF

CALIFORNIA AND SOUTHERN OREGON 218

SYSTEMATIC ACCOUNT 219

Cotfus tenuis 219

Coitus asperrimus 223

Coitus princeps 224

MATERIAL EXAMINED 225

NOTES ON THE CLASSIFICATION OF OTHER SPECIES 226

Coitus klainathensis 226

Coitus aleuticus 227

Coitus gulosus 228

Coitus asper 229

ZOOGEOGRAPHICAL IMPLICATIONS 229

SUMMARY 231

LITERATURE CITED 231

INTRODUCTION

The sculpiiis of the genus Coitus comprise one of the most perplexing
groups of North American freshwater fishes. Variation is so marked
and often so haphazard that interpretation of the species limits is fre-
quently difficult. Moreover, many of the numerous nominal forms are
based on a few poorly preserved specimens and inadequate descriptions.
Perhaps for these reasons, few systematic contributions on Coitus have
been made in recent years. That by Bailey and Dimiek (1949) added
substantially to our knowledge of western sculpins when they separated

1 Submitted for publication February, 1957. Contribution from The Marine Laboratorj',
University of Miami, Number 176, and from tlie Museum of Zoology, University of
Michigan.

( 213 )

214 CALIFORNIA FISH AND GAME

their new species, C. huhdsi, from C. rhotheus and discussed briefly
other western forms.

Sculpius form an important component of the stream fauna through
much of the West. They are particularly common in colder waters and
are often reported in conjunction with studies on trout and salmon.

The acquisition of new material and a review of type specimens
at the United States National Museum and at Stanford University
now permit a more complete discussion of the several species from
the upper parts of the Klamath and Pit rivers of Oregon and Cali-
fornia. In addition, we present a key to the species of Cottus now known
from the Pacific drainage of California and southern Oregon and dis-
cuss their nomenclature.

The existing difficulty in identifying the numerous species has led to
considerable confusion in the literature and so our primary purpose
at this time is to provide an acceptable classification at the species
level. The delineation of subspecies awaits more exhaustive studies, both
in the field and laboratory.

Leonard P. Schultz, Ernest A. Lachner, and Robert H. Kanazawa
liaA-e kindly made available material and working space at the United
States National Museum. George S. Myers, Margaret Storey, and Nor-
man J. Wilimovsky similarly aided us at the Stanford Natural History
^luseum. Edward C. Eaney loaned specimens from Cornell University.
To Reeve M. Bailey and Carl L. Hubbs appreciation is expressed for
permission to use field notes. Joseph H. Wales obtained the excellent
series of C. asperrimvs from Hat Creek. We are indebted to "\^adimir
Walters for notes on the types of C. alcuticus and C. shasta at Stanford
University.

The following museum abbreviations are employed : CU, Cornell Uni-
versity; UMMZ, University of Michigan Museum of Zoology; SNHM,
Stanford Natural History Museum; and USNM. United States Na-
tional Museum. The number of specimens and their range in standard
length is given in parentheses after each museum number.

METHODS

Counts and measurements have been taken by the methods described
by Hubbs and Lagler (1947, p. 8-15) except as noted below.

Dorsal and Anal Soft Rays. Hubbs and Lagler (1947, p. 10) recom-
mend that the last two elements of the dorsal and anal fins be treated
as one ray. Upon dissection it is found that in many groups of fishes
the last two rays stem from a single interspinal series, whereas each
preceding series supports only one ray. This method of counting is
well founded in such groups. In the genus Cottus, the dorsal and anal
rays are unbranehed in most species and. as described above, the last
two rays are often close together and stem from a single interspinal
series (Figure 1, B). Occasionally, however, this last interspinal bears
three very closely approximated rays, the first well developed and the
last two splint-like (Figure 1, C). In each instance the rays which arise
from the last interspinal are considered one. An important variation
occurs when the last ray is well developed, well separated from the
preceding ray, and stems from its own interspinal (Figure 1, A). Were
this a rarity or of equally probable occurrence in all populations, then

SCULPIN STUDIES

215

the arbitrary rule of counting the last two rays as one would introduce
only a negligible and random error. Unfortunately, this is not the
situation, for when this variant occurs it is likely to involve the entire
sample. Moreover, it is the prevalent condition in such forms as C.
tenuis (Figure 1, A; also Evermann and Meek, 1898, p. 83, fig. 6)
and C. cognatus (Greeley, 1928, pi. 7), and is unusual in most other
species (Figure 1, B and C), as exemplified by C. liubhsi (Bailej^ and
Dimick, 1949, pi. 1). Careful dissection reveals that a very small
and completely imbedded splint ray is often present in the variants;
sometimes the last ray is truly single. To count the last two rays as
one under these circumstances leads to apparent population differences
of one ray, a large variation in this feature, when no such differences
actually exist. To count every ray, no matter how small, leads to an
equally biased result. In this study the last two (or three) rays are
counted as one only when they stem from a single interspinal. With
experience, the true situation is easily seen by noting the degree of
separation of the rays : they are very close together when they stem
from a common interspinal and equally far apart when they arise from
separate interspinals. If there is uncertainty, an incision is made along
the bases of the last several rays to expose the interspinals. In this
manner unbiased counts may be made with little chance of error.

FIGURE 1. Second dorsal fin of Coiius, showing the variation in enumeration
of rays (for clarity, segmentation is omitted). A, every element is counted; B,
last tv^o elements counted as 1; C, last three elements counted as 1.

Pelvic Bays. Every ray is counted no matter how rudimentary.
Since a hidden spine is closely appressed to the first soft ray in all
forms of Cottus, this element is not enumerated.

Branching of Soft Bays. In those species in which the pectoral and
pelvic rays are branched, the branching occurs typically at or near the
tips. The lower rays of the pectoral fin are never branched and the first
ray of the pelvic fin often is simple. Branching is correlated with
growth of the fins; thus, young individuals may have simple rays
(though flattened and fraj^ed at the tips), whereas branched rays are
characteristic of the adult. Usually only a few elements are involved in
the branching of dorsal and anal rays.

Lateral-line Pores. The first pore counted is the first one behind the
union of the opercular membrane to the body. When the lateral line is

216

CALIFORNIA FISH AND GAME

developed outo the tail, the last pore counted is the one directly over
or iramediateh' in front of the posterior edge of the hypural plate. The
lateral line is termed complete when it reaches this point. When it is
incomplete, everj' pore is counted. This count is reliable on adult fish
onl}'; in the present study no fish smaller than 40 mm. in standard
length was used.

Ilead-pore Arrangement. Useful and important differences (unpub-
lished) have been noted by the writers in their studies of this group.
Although we are most concerned with the infraorbital and preopereulo-
mandibular canals, in this study we present in Figure 2 a schematic

AN PN

S0_L-5 L-l

FIGURE 2. Schematic drawing of the head-pore arrangement in a hypo-
thetical species of Coitus. A, lateral view. AN, anterior nasal pore; PN,
posterior nasal pore; SO, supraorbital canal; L-1, L-5, lateral-canal pores
1 and 5; PM-5, pore 5 of preoperculo-mandibular canal. B, dorsal view.
N, nasal openings; lO, infraorbital canal (IO-1 = pore 1); L, lateral
canal; PM, preoperculo-mandibular canal; OP, opercular canal (often lack-
ing); ST, supratemporal canal; COR, coronal pore; INT, interorbital pore.

SCULPIN STUDIES

217

outline of the entire cephalic canal system of a hypothetical species of
sculpin. It is hoped that these features will be investigated by other
workers and that this outline will serve as a basis for comparison. The
terminology is after Hubbs and Cannon (1935) as modified by Illick
{1956). In contrast with Illick {op. cii., p. 206, fig. 1) we enumerate
the pores of the lateral canal from the attachment of the gill mem-
brane forward (the more anterior pores are absent in many species)
and distinguish between the pores of the lateral and infraorbital canals.
For a critical review of this topic the reader is referred to Illick
{op. cit.) and the references cited therein.

Infraorhital Pores. The pore which often is present on the lateral
canal at its junction with the infraorbital canal is not counted.

Freopercnlo-mandihular Pores. Every pore below the lateral canal
is counted. In most species the two preoperculo-mandibular canals do
not unite on the chin and each has 11 pores, as indicated by the formula
11-11. In some species these canals are united on the chin and share a
median chin pore (pore count — 10-1-10, Figure 3. A). In others the
uppermost of the two post-maxillary pores, behind the corner of the
mouth (Figure 3, B; also pi. 1 in Bailey and Dimick, 1949), is absent
(pore count = 10-10). When both conditions exist the pore count is
expressed as 9-1-9.

6-7(P0ST MAX.)

5 A

FIGURE 3. Ventral view of chin of Cof/us. A, postmaxillary pore double, chin pore single,
count = 10-1-10. B, postmaxillary pore single, chin pore double, count = 10-10. Combina-
tions of these conditions yield counts of 9-1-9 and 11-11.

Prickles on the Body. The presence or absence of prickles on the
l)od3^, and their distribution, has been used to characterize almost every
species of Cotiiis. The considerable constancy which this feature
exhibits within most populations undoubtedly led to its employment as
a systematic tool. So much variation occurs between populations of most
species, however, that its geographic constancy in one or two forms,
such as C. prince ps, is difficult to evaluate. A correlation with habitat
has been noted. Within a given species, samples from mountain brooks
are naked or have many fewer prickles, whereas lake populations ex-
hibit the most extensive prickles. The first prickles to appear and the
ones lost last are those mesial to the pectoral fin.

218 CALIFORNIA FISH AND OAME

Preopercular Armature. This feature behaves much like the body
prickles. In species which inhabit mountain brooks as well as lakes,
the lake samples have the longer, sharper, and generally better-
developed spines. The armature is totally absent in some populations
of several mountain forms. Armature, like body prickles, may greatly
assist in the identification of species at any one locality, but is often
unreliable for relating or distinguishing allopatric forms.

In order to facilitate the identification of the species of Coitus in
southern Oregon and California (Pacific slope), we present the follow-
ing key. The data are based on studies of material from throughout the
known ranges of all the species.

KEY TO SPECIES OF COITUS OF THE PACIFIC SLOPE OF CALIFORNIA AND SOUTHERN OREGON =

1. Pelvic rays 3-3; palatine teeth absent; some pectoral rays branched: lateral
line incomplete 2

1. Pelvic rays 4-4 (if 3-3 then lateral line complete) ; palatine teeth present or

absent ; pectoral rays branched or not ; lateral line variable 3

2. Two or 3 (usually) preopercular spines, the uppermost relatively long and

pointed, the next lower typically recurved at tip ; lateral line more com-
plete, usually extending to or beyond base of last dorsal ray (see text
for variability) ; ventral surface of head and abdomen nearly immacu-
late G- tenuis

2. One or 2 (at most) preopercular spines, the uppermost comparatively

short and blunt, the next lower not recurved at tip and usually weak ;
lateral line less complete, not extending beyond base of third from last
dorsal ray ; ventral surface of head liberally sprinkled with melano-
phores C". asperrimus

3. Lateral line usually complete, with 28 or more pores (if fewer, palatine

teeth present) ; palatine teeth variable ; pectoral rays unbranched ; dorsal
spines 8-9 5

3. Lateral line incomplete (usually with 15-25 pores but higher in klam-

athensis below Klamath Falls); palatine teeth absent; some pectoral
rays branched; dorsal spines 5-7 (rarely 8) 4

4. Preoperculo-mandibular pores 10-1-10 ; infraorbital pores 7-7 ; preopercular

armature much reduced or absent ; dorsal soft rays usually 20-21 ; head
pores much enlarged (larger than nasal openings) C.princeps

4. Preoperculo-mandibular pores 10-10 ; infraorbital pores 8-8 ; upper pre-

opercular spine well developed (2 accessory spines often present below
it) ; dorsal soft rays usually 18-19; head pores small (smaller than nasal
openings) C klamathensis ^

5. Anal rays usually 17-18, pectoral rays modally 16 (often 17), dorsal rays

19-22 ; palatine teeth present and well developed C. asper

5. Anal rays 12-15, pectoral rays modally 15, dorsal rays 16-18 (if more, pala-

tine teeth absent) ; palatine teeth variable 6

6. Pelvic fins long (when depressed, reach anus) ; palatine teeth absent;

dorsal soft rays 17-20, total dorsal elements usually 27-29 ; caudal ver-
tebrae 24-27 - G- aleuticus *

6. Pelvic fins short (when depressed, usually fall far short of anus) ; palatine
teeth variable; dor.sal rays 16-18 (rarely 19), total dorsal elements usu-
ally 26 or fewer ; caudal vertebrae 21-24 C. gulosus

-C. beldingi has been reported only once from the Sacramento River basin (Rutter,
1908, p. 147), based on a collection taken prior to 1900. To our knowledge, the
species does not now occur in this drainage.

3 Including C. macrops, see text.

* Including C. protrus^is, see text.

s Including C. shasta, see text.

5

SCULPIN STUDIES 219

SYSTEMATIC ACCOUNT

Cotfus tenuis

(Tables 1 and 2)

Uranidea tenuis. Evermann and Meek, 1898, p. 83-84, fig. 6 (origi-
nal description ; type locality : lower end of Upper Klamath Lake, near
Klamath Falls, Oregon).

Cottus tenuis. Rutter, 1908, p. 145 (comparison with C asperrima
[sic]). Hubbs and Schultz, 1932, p. 50 (compared with tuhulatus).
Hubbs and Miller, 1948, p. 69 (relationship to asperrimus indicated).
Bailey and Dimick, 1949, p. 15 (comparison with C. Jiul)hsi).

The drawing of the type (Evermann and ]\Ieek, 1898, p. 83, fig. 6)
does not accurately depict the body form since the figure represents a
recently spawned and much emaciated individual. The undivided last
dorsal and anal ray, the profuse branching of the upper pectoral rays,
the long lateral line and the preopercular armature are correctly fig-
ured. The short spinous dorsal fin is contiguous with the soft dorsal but
the two are never broadly joined. In most specimens several of the dor-
sal and anal soft rays are branched. The pelvic rays are branched
distally in all but a few of our specimens, mostly young (see material
examined), a condition noted by us in only one other species (C asper-
rimus).

Frequency distributions of fin-raj- counts and numbers of lateral-line
pores are presented in Table 2.

The lateral line is never complete and generally terminates under the
last few rays of the second dorsal fin. Specimens from Sprague River
liave a less well developed lateral line than the other populations studied
(see Table 1). The head pores have seldom been used in studies on this
genus although they are useful in separating many species. In tenuis
there are 10-10 pores in the preoperculo-mandibular canals (61 speci-
mens), rarely 9-1-9 (6 specimens). The infraorbital canal has 8-8 pores
in 61 specimens, with very little variation (8-9 in one and 9-9 in two).

The body prickling is quite variable. Lake specimens usuallj^ have the
entire upper half of the body covered with fine prickles, while creek
specimens have a patch of prickles mesial to the pectoral fin or are
entirely naked. The dorsal surface of the head is never prickled. Simi-
larly the preopercular armature is best developed in lake examples.
Three sharp spines are noted in those from Upper Klamath Lake. Speci-
mens from Sprague River retain the three spines but they are shorter
and more blunt.

Vomerine teeth are present but palatine teeth are lacking in all
material examined by us. Caudal vertebrae (including hypural) number
23 (3 specimens), 24 (7), or 25 (1).

Compansons and Relationships

C. tenuis is most closely allied to C. asperrimus, which inhabits sev-
eral tributaries of the Pit River of the Sacramento River system. A
comparison of these forms is given under the treatment of asperrimus.
No very close relationship is noted with any other species. C. tenuis and
C. asperrimus are unique in having branched pelvic rays. C. princeps

220 CALIFORNIA FISH AND GAME

differs from tenuis in having- tlic dorsal fins fused, 4-4 pelvic rays, 7
dorsal spines, 20-21 dorsal soft rays, a less complete lateral line, mucli
enlar-icd head pores, 7-7 infraorbital ^lores. 1()-l-]0 preoperculo-nian-
dibvUar pores, and in numerous other ways. C. klamathensis shares with
tenuis an absence of palatine teeth and similar head-pore counts. It
differs in having- more numerous dorsal spines (usually 7) 4-4 pelvic
rays, a less complete lateral line (15-22 pores), and a broad union of the
dorsal fins (Gilbert, 1898, fig. on p. 10). Limited material of klamath-
ensis from below Klamath Falls shows a better-developed lateral line.

As discussed below, we treat ma crops and klamathensis as conspecific.
C. gulosus, an extremely variable form, has 4-4 pelvic rays, 8-9 dorsal
spines, unbranched rays in all fins and a more complete lateral line
(usually 32-36 pores). Evermann and Meek (1898, p. 83) ally tenuis
and C. marginatus, probably on the basis of two features which they
share: 3-3 pelvic rays and a lack of palatine teeth. The pelvic rays of
marginatus are not branched, however, and it has 10-1-10 preoperculo-
mandibular pores and 7-9 dorsal spines. In its general physiognomy
marginatus appears more closely allied to perplexus and Mamathensis
than to tenuis and asperrimus.

There is presented in Table 1 a color comparison of prince ps, tenuis,
asperrimus, klamathensis, and "macrops", drawn by C. L. Hubbs from
freshly collected specimens ; that for the first three species is based on
material taken in Upper Klamath Lake (Pelican Bay) on August 14,
1934 (field sta. M34-130), and the notes on asperrimus and "macrops"
are based on specimens from the Fall River system near Dana, Califor-
nia, taken on August 17, 1934 (field sta. M34-135). Statements in
parentheses under asperrimus are our observations where no specific
reference to these items appears in Dr. Hubbs' notebook.

Geographic Variation

Populations from Upper Klamath Lake, Agency Lake-Crooked Creek,
and Sprague River — all in the Klamath basin — exhibit some degree of
differentiation. We do not feel that these differences are sufficient to
merit nomenclatural recognition. Specimens from Agency Lake and
Crooked Creek are most similar to the Klamath Lake population but
differ in having somewhat fewer dorsal soft rays and pectoral rays.
The Sprague River material has a distinctly lower pectoral-ray count
and somewhat fewer pores in the lateral line.

Habifs

An examination of the abundant type material of tenuis, collected on
November 3, 1896, reveals many females with enlarged eggs and others
which have obviously spawned. This evidence, coupled with Evermann
and Meek's (1898, p. 84) reference to a ripe female, definitely shows
that tenuis is a fall spawner in Klamath Lake.

Geographic Range

C. tenuis is restricted to Upper Klamath Lake and its tributaries,
Oregon.

SCULPIN STUDIES

221

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SCULPIX STUDIES 223

Cottus asperrimus
(Tables 1 and 2)

Cottus asperrima. Rutter, 1908, p. 14-i-145, fig. 3 (original descrip-
tion; type locality: Fall River at Dana, California). Snyder, 1917, p. 82
(synonym of gulosus) .

Cottus asperrimus. Hubbs and Schultz, 1932, p. 5 (compared to
tuhulatus). Hubbs and Miller, 1918, p. 69 (relationship with tenuis).
Bailey and Dimick, 1949, p. 16 (removed from synonymy of C. gulosus).

Frequency distributions of fin-ray counts and numbers of lateral-line
pores are given in Table 2. C. asperrimus is so similar to tenuis that
only the differences between them need to be noted. This similarity must
have been suspected by Rutter (1908, p. 145) since he presented a
tabular comparison between these two species, but the characters listed
by him are slight and do not hold wlien comparable specimens are exam-
ined. We note several diagnostic features. Typically, asperrimus has
only one well-developed preopercular spine and a blunt knob below it.
often absent, whereas tenuis has three preopercular spines. The lateral
line is less well developed in asperrimus, generally extending to the base
of the second to fifth ray from the last, but reaching to or beyond the
last dorsal ray in tenuis. The latter species spawns in the fall, whereas
asperrimus evidently spawns earlier (see Habits, below.) The life colora-
tion (Table 1) appears to be distinctive and the pigmentation also
differs, asperrimus showing profuse chromatophores on the venter,
whereas tenuis is nearly immaculate. Although the pelvic, soft dorsal,
and anal rays of asperrimus are branched, as in tenuis, the degree of
branching seems to be weaker, particularly on the smaller topotypes
(UMMZ 130650), than it is in tenuis. The number of preoperculo-
mandibular pores is variable. In 19 specimens (including the holotype)
the formula is 9-1-9, whereas in 41 specimens (including 2 paratopo-
types, SXHM 9968) it is 10-10, as in tenuis. The two forms perhaps
warrant only subspecific status, but our material is insufficient to decide
this point with assurance.

An extensive patch of prickles is present mesial to the pectoral fin,
and in most specimens the prickling extends caudad along the body
above the lateral line to a point under the middle of the second dorsal
fin. Few specimens exhibit such extensive prickling as the holotype
(Rutter, 1908, p. 144, fig. 3). The figure of the type errs in showing
unbranched pectoral rays and a peculiar head profile. Actually, the
ventral contour is flat to the tip of the chin, as in other sculpins, and
the dorsal profile turns abruptly downward in front of the eyes.

Comparisons

C. asperrimus differs from other sculpins in most of the characters
which separate them from C. tenuis (see above). The 3 soft rays of
the pelvic fins readily distinguish it from klamathensis, which occurs
with asperrimus and has 4 pelvic rays.

The origin of asperrimus is considered in the zoogeographical dis-
cussion.

224 CALIFORNIA FISH AND GAME

Geographic Variafion

Available material of this species is inadequate to determine the
rauo'e of variability but does show that the number of pectoral rays in
samples from above the Pit River falls is significantly greater than it is
in the one sample from below these falls (15 versus 14, Table 2).

Habits

Specimens of asperrimns from Hat Creek (below Pit River falls)
taken on May 6, 1949, include females with large eggs, suggesting that
this species, in contrast with tenuis, is a spring spawner.

Geographic Range

C. asperrimus is probably restricted to the Pit River portion of the
Sacramento River system, for a short distance above and below the falls
at Fall River Mills, California.

Cottus princeps
(Tables 1 and 2)

Cottns princeps Gilbert . 1898, p. 12-18, fig. on p. 12 (origmal descrip-
tion; type locality: Upper Klamath Lake, Oregon). Evermann and
Meek, 1898, p. 83 (specimens from Pelican Bay, Upper Klamath Lake).
Hubbs and Schultz, 1932, p. 4-5 (comparison with tuhulatus). Hubbs
and Miller, 1948, p. 68 (adaptation to lake life). Bailey and Dimick,
1949, p. 15-16 (compared with huhhsi; characters).

Cottus evermanni Gilbert, 1898, p. 11-12, fig. on p. 11 (original de-
scription ; type locality : Lost River near Klamath Falls, Oregon) . Hubbs
and Schultz, 1932, p. 4-5 (comparison with tuhulatus). Bailey and
Dimick, 1949, p. 15-16 (synonymized with princeps, rule of first
reviser).

C. princeps is one of the most distinctive species in the genus. There
are generally 7 dorsal spines, 20-21 soft dorsal rays, 16 anal rays,
4 pelvic rays, 15-16 pectoral rays, and 15-25 lateral-line pores (Table
2). The lateral line is incomplete and ends under the posterior half of
the second dorsal fin. The dorsal fins are entirely fused. A few speci-
mens show some branched dorsal and anal rays, but this is unusual. The
pelvic rays are never branched, whereas the upper pectoral rays are
usually, but not always, branched.

The pelvic fins are long, reaching the anus when depressed. The body
is extensively prickled. Below the lateral line the prickling usually
extends caudad to a point about under the center of the second dorsal
fin. Above the lateral line the entire body is prickled, except on the
caudal peduncle and head.

The figures of the types of pjrinceps and evermanni accurately depict
the body form and color pattern. The anterior nasal tubes are very-
well developed. The head pores are much enlarged ; those of the infra-
orbital and preo])erculo-mandibular canals are larger than the nasal
openings. The canals are very prominent and cause cavernous openings
in the bones through which they pass. All 34 specimens examined have
10-1-10 preoperculo-mandibular pores. Infraorbital pores number 7-7
(12 specimens) or 6-7 (2 specimens).

SCULPIK STUDIES 225

The palatine teeth are absent. The preopercular armature is re-
duced to one blunt spine. The lower jaw usually projects slightly
beyond the upper jaw. We note 24 caudal vertebrae in two specimens
and Baile}^ and Dimick (1949, p. 16) recorded 26.

Comparisons and Relationships

The combination of characters listed aboA'e imparts to princeps a dis-
tinctiveness that renders it unmistakable. C. Mamathensis has the dorsal
fins partly or completely fused, lacks palatine teeth, and has 4-4 pelvic
rays. It differs trenchantly in its lower ray counts (second dorsal
rays usually 18-19 instead of 20-21 ; anal rays 13-14 instead of 15-17), its
higher number of infraorbital pores (8-8 instead of 7-7), its separate pre-
operculo-mandibular canals (pores number 10-10 instead of 10-1-10),
its more robust body form, and in many other ways. Distinguishable
features for tenuis and aperrimus have already been exi^lained. C. gulo-
sus has separate dorsal fins, 13-15 anal rays, 8-9 dorsal spines, 16-18
soft dorsal rays, and a much more complete lateral line (usually 32-36
l)ores).

The relationships of princeps are uncertain. The higli number of
caudal vertebrae (24-26) places it with aleuticus and asper. It shares
with those species a prcopei-culo-niandibular pore count of 10-1-10. G
aleuticus apparently never develops palatine teeth, while asper has well
developed palatine tooth patches. Like princeps, asper has a long sec-
ond dorsal (19-22 rays) and anal fin (15-19 rays). C. aleuticus similarly
has a long dorsal fin but has fewer anal rays (12-15, usually 13-14).
Both generally have comi)lete lateral lines, although aleidicus is more
variable in this respect. The color pattern of asper is suggestive of
princeps. The long pelvic fins of aleuticus reach the anus when de-
])ressed, a feature seldom noted in othei- American species of Cottus but
characteristic of princeps and several Asiatic species. C. asper is more
prone to move inland than is aleuticus, a form common in coastal
streams. If princeps is derived from either an asper or aleuticus type,
the two have long been differentiated. Its relationships to other Amer-
ican species appear to be even more distant.

Hahih

All females in a series of s]iecimens collected on November 3-5, 1896,
by Meek and Alexander (USN]\I 124952), have the ovaries bulging with
mature eggs.

Geographic Range

C. princeps is known only from Upper Klamath Lake and from Lost
River (near Klamath Falls), Oregon.

MATERIAL EXAMINED

Cottus princeps. USNM 48227 (1. 41) holotype ; USNM 48228 (1,
47), holotvpe of C. evermanni. The following are all from Upper Kla-
math Lake. Ore.: USNM 124952 (18, 30-68); UMMZ 130627 (14, 30-
44) ; UMMZ 138574 (7, 47-64) ; SNHM 38029 (29, 45-60) ; and SNHM
40744 (2,42-63).

226 CALIFORNIA FISH AXD GAME

Cottus tenuis. USXM 48229 (1, 74), holotvpe. All others are para-
topotypes: USXM 48280 (38. 25-62); UtSNM 124958 (38, 23-70);
USNM 48780 (13, 22-71) ; and SXIIM 5705 (20, 20-44).

Additional material examined from Upper Klamath Lake, Ore. :
UMMZ 130603 (2. 38-51); V^niZ 130626 (6, 32-49); UMMZ 172293
(4, 26-38) ; and SXHM 9333 (1, 86).

Other material: Klamath County, Ore.: UMMZ 130610 (149, 24-80)
Crooked Creek; UMMZ 130581 ?9, 20-56) Sprapue River; UMMZ
] 30617 (27, 21-44) Agency Lake, month of Wood River; and SXHM
24185 (11, 36-48) Spring Creek, Shasta Connty, Calif.

Cottns asperrimus. USXM 58500 (1, 50 j. holotvpe; USXM 126463
(1, 48), paratopotype ; SXHM 9968 (2, 41-49), paratopotvpes ; UMMZ
130647 (1, 39) Rush Creek, Modoc County, Calif.; UM:\fz 130650 (7,
14-64) Fall River system. Shasta County, Calif.; and UMMZ 158416
(28, 41-89) reservoir on Hat Creek, about 4 miles above its junction
with Pit River, Shasta County, Calif.

NOTES ON THE CLASSIFICATION OF OTHER SPECIES
Coifus klamathens'is

Cottus macrops, described from Fall River, California (Rutter, 1908,
p. 146-147, fig. 4), has generally been maintained as a valid species
(Hubbs and Miller, 1948, p. 69 ;" Bailey and Dimick, 1949, p. 15; and
others), although these -writers noted its similarity to Mamathensis.
Snyder (1917, p. 82) failed to observe the intimate relationship of
macrops and klamathensis and combined both macrops and asperrimus
with gulosKS.

A careful comparison of the type material of klamathensis and
macrops and of fresh collections f)f the two nominal forms fails to reveal
differences that we .judge to be of specific value. Rutter recognized the
very close relationship between the two but indicated that macrops had
a heavier build, a larger eye, the head less pointed, and the dorsal fin
somewhat more posterior in position. Our material fails to confirm
some of these distinctions (e.g., the larger eye), and we regard other
features to be too variable or indicative of ranking below the species
level. Like klamathensis, the body prickles in macrops occur on smaller
individuals only (up to about 55 mm. standard length), the large adults
having the skin entirely smooth. Generally, macrops has only one well
developed preopercular spine, with the two accessory spines reduced
to blunt kjiobs or absent. In many samples of klamathensis there are
individuals that have all three spines developed, others that have only
two, but populations from beloAv Klamath Falls are somewhat inter-
mediate between klamathensis (above the Falls) and macrops in this
feature, having one or two developed spines. Life colors (Table 1) of
the two nominal forms indicate no marked differences. The most con-
sistent difference we have noted between the two involves the width
of the mouth. In macrops, the snout is blunter and the mouth wider
(its greatest width enters the anal base 1.2-1.6 times). In klamathensis
(above the Falls) the snout is more pointed and the mouth narrower
(its greatest width enters the anal base 1.6-2.0 times; in one collection
(15 specimens) below the falls, the ratios vary from 1.3-1.8 times).
The reliability of this feature is uncertain, and we feel it can be used

SCULPIN STUDIES 227

for subspecific separation at most. The two share a preoperculo-mandib-
nlar pore count of 10-10, lack palatine teeth, have broadly connected
dorsal fins, have the vertical fins similarly marbled and with a black
blotch on the posterior membranes of the first dorsal, and have similar
fin-ray counts, which mav be summarized as follows : dorsal spines, 6
(1), 7 (25), 8 (6); dorsal soft rays, 18 (10), 19 (21), 20 (1); anal
rays, 13 (6), 14 (25), 15 (1) ; pelvic rays, 4-4 (26 fins) ; pectoral rays,
14 (9 fins), 15 (49 fins), 16 (3 fins). The lateral-line pores range from
14-22. These data are based on the following material: USNM 58499
(14, 23-57) syntypes of macrops; USNM 126464 (4, 22-51) svntvpes
of macrops; SXHM 24186 (4, 31-57) svntvpes of »mcrops/ USNM
88464 (2, 40-44) Fall River, Calif.; USNM 48266 (1, 118) tvpe of
klamathensis ; and CU 21612 (7, 24-76) Shasta Eiver, Calif.

"We synonymize macrops with Idamaihensis but note that study of
additional features may show them worthy of subspecific rank. Com-
jjarison of populations from above and below Klamath Falls shows that
those from the lower river (excluding one sample from Shasta River)
have a more complete lateral line (this was also noted by Gilbert, 1898,
p. 11), and these may represent a third form worthy of naming. Gilbert
believed Mamaihcnsis to be most nearly related to perplexus, a view
which we share. C. klamathensis affords another example of fishes
shared by the upper Klamath and Pit rivers (see discussion of zoogeog-
raphy).

Cottus aleuticus

This species is widespread in coastal streams from central California
(northern San Luis Obispo County) to Alaska. As noted by Ilubbs
and Schultz (1932, p. 6, table 1) it belongs to those species of Cottns
which have many (24-27) caudal vertebrae. In other features it is dis-
tinguished by having 10-1-10 preoperculo-mandibular pores, 8-9 dorsal
spines, 17-20 dorsal rays (usually 18-19), 12-14 anal rays, 4 pelvic
rays, 14-16 pectoral rays, long pelvic fins, no palatine teeth, and few
if any prickles. Schultz and Spoor (1933) described C. protrusns from
Unalaska Island. They noted its similarity to aleuticus in several fea-
tures but contrasted protrusus only with chomherlaini, a very differ-
ent form doubtfully distinct from C. cognatus. The holotype (USNM
104496) and a paratopotype (UIVEVIZ 64234) of protrusus were com-
pared with the syntypes of aleuticus (SNH^M 2610) from Uiuliuk,
Unalaska, and with other specimens from Alaska to California (vari-
ously labelled as protrusus, aleuticus, or Uranidea microstoma). No
differences signifying other than individual variation were noted.
Specimens which exhibit protruding eyes (the basis of protrusus)
are known from widely scattered localities and appear even in the
syntypic series of aleuticus. Whether this is a response to temperature
(improbable) or one induced by disease or parasitism is not known.
We regard the condition as an anomaly and, in the absence of any other
difference, consider protrusus synonymous with aleuticus. Uranidea
microstoma Lockington is preoccupied in Cottus, a fact long known, and
was placed in the synonymy of aleuticus when that species was de-
scribed. In the course of this study we re-examined the holotype
(USNM 26922) and take this opportunity to confirm its identity with
aleuticus.

228 CALIFORNIA FISH AND GAME

In snnimariziiig the distribution of this species in California, Ever-
niann and Clark (1981, p. 57) overlooked the southernmost record given
by Ilubbs (1921, p. 7-8). The presence of aleuticus in northern San Luis
Obispo County is confirmed by a recent collection (UMMZ 141194) of
74 specimens from San Carpoforo Jja.iioou, just sonth of the Monterey
County line.

Shapovalov and Taft (1954, p. 26) record this species from Waddell
Creek and (in litt.) note that their extensive collections are deposited
in the California Academy of Sciences.

Cotfus gulosus

It is unfortunate that this variable species, known widely in the
Sacramento-San Joaquin drainage and several lesser coastal streams of
California, has been considered a subspecies in the Ijairdi complex by
recent writers (Hubbs and Wallis, 1948, p. 141-142; Shapovalov and
Dill, 1950, p. 387, as C. h. shasta). There is little positive evidence to
support this relationship and many reasons for doubting it. We feel that
this subspecific allocation has caused considerable confusion and, pend-
ing more thorough study, consider gnlosiis a valid species very closely
allied to Jmh'bsi. For the present, we feel that the only western forms of
Coitus that should be aligned specifically with hairdi are C. &. semisca-
her and C. h. iDunctuJatus of the upper Columbia and Colorado rivers,
respectively.

We agree with Synder (1905, p. 337) that C. shasta and C. gulosus
are synonymous. A re-examination of the types (SNHM4193, 3 syntypes
of shasta; USNM 291, 2 syntypes^ of gulosus) and other material, in-
cluding topotypes, from the range of both forms, fails to reveal any
basis for specific separation. The preoperculo-mandibular pore count is
quite variable (10-1-10, 10-10, 11-11) but shows no geographic con-
sistency. A study of fin-ray couuts and numbers of lateral-line pores
indicates no differences between shasta and gulosus but suggests tliat
populations in coastal streams from Monterey Bay northward have a
much less complete lateral line.

TJranidea scmiscahra ccntroplpura Eigcnmann and Eigenmann has
never been adequately discussed. Jordan and Evermann (1898, p. 1945)
and Evermann and Clark (1931, p. 57) identified it with gulosus. Rutter
(1908, p. 145) placed it in the synonymy of asper. Re-examination of
the type (USXM 41919, 63 mm. standard length) shows conclusively
that this form belongs to the gulosus series. The two dorsals are separate
but contiguous. Fin-ray counts are as follows : 8 dorsal spines, 17 dorsal
soft rays, 13 anal rays, 13 rays in each pectoral, and 4 rays in each
pelvic fin. There are 35 lateral-line pores and 10-1-10 preoperculo-man-
dibular pores. Palatine teeth are present but not strongly developed.
These features are all characteristic of gulosus except the few pectoral
rays ; should this low count be typical then subspecific recognition might
be warranted for centropleurus.

Most of the records of gulosus from Washington and northern Oregon
are based on C. perplexus Gilbert and Evermann, a species synonymized
with gulosus by Schultz (1930, p. 14). None of these records represents
gulosus as we define that species. Counts of the holotype of perplexus,

« Jordan and Evermann (1898, p. 1945) erred in listing USNM 290 as types.

SCULPIN STUDIES 229

USNM 43387 (1, 71), and of 14 of the paratopotypes, USNM 45388
(24, 18-55), are summarized as follows: dorsal spines, 7 (13 speci-
mens)^ 8 (2) ; dorsal soft rays, 18 (4), 19 (5), 20 (2)^; anal rays, 13
(2), 14 (4), 15 (6)^ 16 (3) ; pelvic rays (both fins), 4 (30)^ pectoral
rays, 14 (3), 15 (18), 16 (9)^ lateral-line pores, 20 (2) ; 21 (1), 22
(1)^ 23 (1), 24 (3), 26 (1)^; preoperculo-mandibular pores, 10-1-10
(8)'^, 11-11 (1). Two other specimens in the series of paratopotypes are
C. rhotheus. In its robust body form, strongly marbled vertical fins,
and the joined dorsal fins, C. perplexus seems more closely allied to
klamatJie7isis then to gnlosus. The type material lacks palatine teeth
but in 39 subtopotypes (UMMZ 93437) these teeth are present or
absent.

Coifus asper

Jordan and Evermann (1898, p. 1945) and Jordan, Evermann and
Clark (1930, p. 383) listed Cottopsis parvus Girard in the synonymy of
(julosus. Of the four series of specimens recorded by Girard in his sum-
mary account (Girard, 1858, p. 54-55), we have located one of the 12
from Petaluma (UMMZ 171133) and three of the four from Fort
Reading (USNM 295; 3, 88-103). Since Girard considered only the ma-
terial from Presidio in his original description (1854, p. 144-145), the
specimens listed above are not types. Nevertheless, study of this mate-
rial and of Girard 's description leaves no doubt in our minds that
Cottopsis parvus is identical with Cottus asper. Particularly diagnostic
are the numerous dorsal and anal rays and the body prickles. A varia-
tional study of this widespread species is much needed. Its role as an
important predator on juvenile trout and salmon is discussed by Sha-
povalov and Taft (1954, p. 293). Their abundant material is included in
the collections of the California Academy of Sciences.

ZOOGEOGRAPHICAL IMPLICATIONS

The mutual occurrence of Cottus klamathensis and the allopatric
forms C. tennis and C. asperrimus in the upper Pit and Klamath rivers
poses an interesting problem in distribution and origin. A consideration
of the fish faunas of the Klamath and Sacramento basins as a whole
is necessary to an understanding of the hypothesis proposed below to
explain the distribution of the species of Cottus.

Despite its geographic position between the Sacramento and Columbia
drainages, the fauna of the Klamath basin surprisingly lacks the char-
acteristic endemic genera of either of those great river systems (Miller,
1946, table 1). Even Ptycliocheihis, ubiquitous in both the Sacramento
and Columbia and with a representative species in streams between the
Klamath and Columbia rivers, is absent in the Klamath system. Yet the
latter basin has a comparatively rich, highly endemic fauna that is sug-
gestive of long isolation (Hubbs and Miller, 1948, p. 68-69, and refer-
ences cited). Close relationships are shown, however, between certain
fishes of the Klamath Lakes and the Great Basin; for example, the
peculiar suckers of the genus Chasmistes and the distinctive chubs as-
signed to the genus Siphateles.

^ Holotype.

230 CALIFORNIA FISH AND GAME

In addition to the common bond between the upper Pit and Klamath
rivers exemplified by the species of Coitus, there is a dwarf non-para-
sitic lamprey (genus Entosphenus) known only from these two regions,
and the stream representatives of Siphateles bicolor also show intimate
relationship.

These similarities call for some kind of past connection between the
Klamath system above Klamath Falls and what is now the upper seg-
ment of Pit River, including Goose Lake. It is reasonable to assume
that the connection was by way of interconnecting streams through a
chain of lakes (see Cope, 1883, map opposite p. 166) on the northern
part of the Modoc Plateau and that the evidence for the waterway lies
buried beneath the extensive Modoc lava flows (Anderson, 1941). That
none of the endemic Sacramento forms reported from the upper Pit
drainage (e.g., Hesperoleucus, Myloplmrodon, PtycJiocheilus, and Hys-
terocarpus) occurs in the Klamath suggests that the connection between
these two basins was severed prior to the invasion of the upper Pit
region (through headward erosion by Pit Eiver) by these Sacramento
types.

With this picture in mind, we may envision the origin and differenti-
ation of two species of Cottus in the upper Pit as follows. After pene-
trating the Klamath basin prior to the formation of Klamath Falls,
the ancestors of C. klamafhensis and C. tenuis subsequently crossed
the ]\Iodoc Plateau to reach what is now the upper Pit basin and thus
attained a wide distribution in these presently disrupted waters.
When the connection between the Klamath and Pit rivers was severed,
the isolated stocks underwent differentiation, weaker for C. Mamatli-
ensis than for C. tenuis, resulting in the evolution of the allpatric
form C. asperrimus. With the development of the Pit River falls, the
Hat Creek population of asperrimus below the barrier further dif-
ferentiated. Although by this hypothesis we regard the Hat Creek
population as a downstream derivative of asperrimus, an alternate
idea may be considered. This is to regard tenids and the Hat Creek
population as peripheral remnants of a single form which has some-
what diverged and that the form occupying the old center area has
diverged most into what is now called asperrimus. Perhaps future work
will show that there are three subspecies of tenuis, one in the upper
Klamath, one in the upper Pit, and one in Pit river below the falls.
Our material, however, does not permit us to adopt this arrangement
but it makes clear the intimate relationship between asperrimus and
tenuis.

The clear indication of close relationship between the fishes of these
two regions with those in the adjacent Great Basin finds support
amongst other vertebrate groups, as in birds. The northeastern corner
of California, including the upper part of Pit River (mostly above the
falls), is ascribed wholly or largely to the Modoc faunal district (of
the Great Basin faunal division) by Grinnell (1915, p. 12. pi. Ill) and
by Miller (1951, p. 591, fig. 4), a district which obviously extends
into the Klamath Lake region.

SCULPIN STUDIES 231

SUMMARY

The characters, distribution and geoo-raphic variation of C. tenuis,
nsperrimiis, and princcpa are treated and nomeiu-latnral notes are given
for klamathensis, aleuiicus, gulosus, and asper.

Emphasis in this paper has been directed toward working out rela-
tionships at the species level. Detailed studies need to be made in order
to determine differentiation of subspecific value.

C. asperrimus, of Pit River, California, is regarded to be a very close
relative of tenuis, limited to the upper Klamath drainage of Oregon
and California; further studies may show them to be conspecific.

The placement of evermanni in the synonymy of princeps is con-
firmed and the possible relationship of this species to aleuiicus and
asper is discussed.

C. macrops is considered to be conspecific with klamathensis, pro-
Irusus is placed in the s^'nonymy of aleuiicus, shasfa is regarded as
identical with gulosus, and parvus is synonj-mized with asper.

It is urged that the name hairdi not be used for any western forms
other than punctulatus, and semiscaher. The application of this name
to gulosus has been particularly confusing and does not seem war-
ranted in the light of our present knowledge.

Uranidea sentiscahra centropleura is aligned with gulosus and may
prove to be a valid subspecies.

C. perplexus is removed from the synonymy of gulosus and is con-
sidered a valid species, but its relationships with klamathensis need
to be clarified.

The relationships of the fish fauna of the upper Klamath and Pit
rivers is emphasized in a section on zoogeography, with a discussion
of the origin and dispersal of the species of Coitus common to these
two regions.

A key to the species of Cottus in the Sacramento and Klamath drain-
ages is presented.

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232 CALIFORNIA FISH AND GAME

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Rutter, Cloudsley

1908. The fishes of the Sacramento-San .loaquiu basin, with a study of their
distribution and variation. U. S. Bur. Fish., Bull., vol. 27 (1907), p.
103-152, 4 figs., pi. 6.

Schultz, Leonard P.

1930. Notes on the species of Cottus in western Washington. Copeia, 1930, no.
1, p. 14-15.
Schultz, I^onard P., and William A. Spoor

1933. Cottus protrusus, a new sculpin from Unalaska Island. Copeia, 1933, no. 3,
p. 142-145.

SCULPIN STUDIES 233

Shapovjilov, Leo, and William A. Dill

I^.jO. a check list of the fresh-water and auadromoiis fishes of California. Calif.
Fish and Game, vol. 36, uo. 4, p. 382-301.

Shapovalov, Leo, and Alan C. Taft

1954. The life histories of the steelhead rainbow trout {Sahno gahdneri gaird-
iicri} and silver salmon (Oncorht/nchiis kisutch) with special reference
to Waddell Creek, California, and recommendations regarding their man-
agement. Calif. Dept. Fish and Game, Fish Bull. 98, p^ 1-375, 32 figs.
Snyder, John Otterbein

1905. Notes on the fishes of the streams flowing into San Francisco Bay. U. S.
Bur. Fish., Rept., 1904. p. 327-338.

1917. The fishes of the Lahontan system of Nevada and northeastern California.
U. S. Bur. Fish.. Bull., vol. 35 (1915-16), p. 31-86, 9 figs., pis. 3-5.

NOTE

THE REPORTED EMERGENCE OF FISHES FROM CALIFORNIA'S
SUBTERRANEAN WATERS

On the basis of core samples and general stratigraphy, geologists
believe that in most parts of California there is small possibility that
organisms could emerge alive from underground sources, either through
artesian Avells or deep well pumps. If organisms were to emerge alive,
they would theoretically have to come through solution channels, which
means limestone formations. There is very little of this type of material
in California's geological makeup.

AYhen fishes emerge from a newlv-di'illed well, one fact usuallv over-
looked is that considerable water may be pumped into the shaft during
drilling operations. The water acts as a lubricant and is circulated under
a pressure head up to 50 pounds. Although we have not tested fishes
under pressure, we have found that sticklebacks (Gasterosteiis a.
acuhatus) in the laboratory will live for several hours in a heavy mud
slurry. Eowley (1955) found that eight-inch rainbow trout {Salmo
gairdneri) suffered no ill effects from the application of 200 pounds
pressure for 48 seconds. Needless to say, any fish emerging with the
drilling water or immediately thereafter must be suspected of having
entered the well at the time the water was pumped in.

There are three early California records, all questionable. Saxe ^ de-
scribed the artesian wells at San Jose, pointing out that fish occasion-
ally came from the wells and that sometimes sawdust flowed out with
the water. He was satisfied that the fish did not come originally from
a subterranean source but that they found their way there from open
streams or beds of water through underground channels. Chase (1874a)
told of a 143-foot artesian well drilled five feet above high tide line
on the beach at Point Hueneme. Pressure forced the w^ater 30 feet into
the air, along with thousands of two-inch "trout" (water temperature
64° F.). Smith (1884) reported a l>^-inch stickleback (Gasterosteus
acnJeafus wiUiamsoni) coming from the pipe of a 191-foot artesian w^ell
at San Bernardino, California. The locations of seven California fish
emergence records are shown in Figure 1.

Data on four recent fish emergences are given in Table 1. Of these,
the Dowd and Cholame ranches have been studied in detail by the
writer. For the latter there is ample evidence that the sticklebacks
actually did come from the well. Two individuals, Mr. Howard Jack
and Mrs. Palmateer Gibson, have described to me their observations of
the sticklebacks swimming within the well shaft. The velocity of the
artesian flow from the well would have prevented any fishes swimming
up to the well. This contradicts the viewpoint of Noble (1952), who

1 Remarks made in the report of the regular meeting of July 18, 1870, of the California
Academv of Sciences, reported in the Proceedings, California Academv of Sciences,
vol. 4, p. 130.

(235)

236

CALIFORNIA I'ISIl AM) (iA.Mi:

//Wheatland 1946

Unidentified Fish

' Lincoln 1950

Socramenfo Squawfisli

>San Jose 1870

\ Unidentified Fish
\
\
\
\
\
\
\
\

Shandon 1952

Sfic/(lebocks

\
\

Oxnard 1953
Stic/<tebac/<s .

Point Hueneme 1874

"Trour

San^prnardino 1884

'fict(teback

FIGURE 1. Locations of several subterranean fish emergence records in California.

NOTE

237

TABLE 1
Recent Californian Subterranean Fish Emergences

Ranch name

James Clack

Delbert Dowd

Cholame Rancho

Daryl Arnold

Location

3J^ miles north-
west of Wheat-
land

3 miles northwest
of Lincoln

North of Shandon

Near Oxnard

Geological forma-
tion

lone; Eocene clay, sand, shale

Quaternarj' alu-
vium conglom-
erate

Quaternary alu-
V i u m heavy
adobe

Kind of fish

"li4 inches, long
and silvery"

5 Sacramento
squawfish, Pty-
chocheilus gran-

About 50 .stickle-
backs

5 sticklebacks

Gasterosteus aculeatus microcephelus

Date of emergence

Spring, 1946

October 10, 1950

February 5, 1952

January 6, 1953

Location of speci-
mens

Unknown

California Acad-
emy of Sciences
20,504

California Acad-
emy of Sciences
20,693

Unknown

Depth of well

465 feet

543 feet

600 feet

1,300 feet

Diameter of shaft

14 inches

Unknown

Unknown

16 inches

Gallons of water
used in drilling

Unknown

500

36,000

Unknown

Source of drilling
water

Unknown

Main ranch well

Roadside pools and
main well

Old well and ad-
jacent new well.

Rate of effluent
flow or pump
size

10-inch pum))

8-inch pump

1 ,200 gallon per
minute artesian

1,150 gallons per
minute

Nearest permanent
water

3 miles

Coon Creek 2
miles north

Several miles

Calleguas Creek 3
miles south

Source of data

Correspondence

Several interviews

Many interviews

Correspondence

believed that the fish liad swum to the well by means of surface puddles.
However, we cannot be certain that some of the sticklebacks were not
inadvertently placed in the well with the drilling- water from the road-
side ditches.

It is hoped that careful observations of future emergences of fishes
from deep wells will provide sufficient data to provide answers to some
of the questions raised herein.

REFERENCES
Chase. A. W.

1874;i. Note on subterranean fishes in California. Amer. Jour, ^^ei.. .ser. 3, voL

7. p. 74.
lS74b. Note on subterranean fishes in California. Forest and Stream, vol. 2, p. 70.
Noble, Glenn A.

1952. The case of the artesian sticl<lebacks. Calif. Fish and Game, vol. .38. no.
3, p. 439.
Rowley, William E., Jr.

195.J. Hydrostatic pressure tests on rainbow trout. Calif. Fish and Game, vol. 41,
no. 3, p. 243-244.
Smith. Rosa

1884. Note on the occurrence of Gtisterosteiis uiUiainsoni Girard. in an artesian
well at San Bernardino, Calif. U. S. Nat. Mus., Proc, (1883) , vol. 6, p. 217.
Earl 8. Herald. Sfeinharf Aquarium, California Academy of Sciences, San Francisco
IS, California, January. 1057.

REVIEWS

Tomorrow's Birthrighf

By Barrow Lyons ; Funk & Wagnalls Company, New York, 1955 ; viii + 424 p. $5.

In Tomorrow's Birthright, a history of the political battles which have been
fought over our natural resources, the reader will find a wealth of reliable informa-
tion. Anyone who desires to bring himself up to date on the whole problem of con-
servation of natural resources will do well to put this book on his desk and keep
it there until he has digested every chapter. Here is no dry compilation of statistics
but a carefully written, interesting book which discusses our past, present, and
future conservation problems. It is "a political and economic interpretation of our
natural resources".

The author, Barrow Lyons, divides his book into three parts. Part I gives a
complete discussion of our problems relating to soil, forest, and water resources.
Part II — The Politics of Power — debates the methods most suitable for a per-
manent control of our land and water resources. Part III — Resources for Sur-
vival—draws a bead on our future prospects for survival and includes discussions
of oil, atomic power, solar energy, and basic metals.

At no point does Mr. Lyons lose sight of his theme : the political problems con-
nected with planning our economic future. He believes that our experience has
shown that whenever the American people are fully and correctly informed and
have been alerted to the dangers involved, they can be relied upon to make the
proper choice among courses of action. He proposes, in his book, to place the
necessary information before the voters. This book differs from previous books
on conservation in that it presents this full account of the interaction of politics
and our natural resources.

For every voter, and every member of Congress or a state legislature. Tomorrow's
Birthright will lay the groundwork for intelligent understanding of this very im-
portant problem. It is clearly written and the type is excellent. Every conservation-
ist should be pleased to add this book to his library.

"It is not suggested here that we have become so weak we may not survive. That
could happen. But plenty of good land remains, plenty of forest that can be used
and made to yield lumber for thousands of years, plenty of grassland upon which
cattle may graze. And much of what is lost can be regained. But it has become
necessary for all of the people operating individually or through the government
to promote a great effort in conservation and wise use ; and there are other meas-
ures which must be adopted if we are to stop the destruction of our natural re-
sources soon enough to rescue them so that they will provide for the future." —
Willis A. Evaus, California Department of Fish and Game.

Cowry Shells of World Seas

By Joyce Allan ; Georgian House, Melbourne, Australia, 1956 ; x + 170 p., with 15
plates and text figures. 63 shillings, Australian.

Although cowries are found in most tropical and subtropical seas of the world
and have always been highly prized for their superb beauty, there have been few
complete treatises such as this available to the modern conchologist or amateur
shell collector. The present volume, which undoubtedly represents many months
of intensive work on the part of the author, includes an introductory chapter
containing a considerable amount of general information on the life history, habits,
and classification of cowries. This section is followed in turn by chapters detailing
the various genera and species of true cowries and allied cowry gi-oups. For each
species the naming authority and date are given, as well as descriptive information
on shell color, size, and characters. The known geographical range is stated and
occasionally synonyms and other pertinent data.

Six of the 15 plates illustrating the species discussed are in color and add much
to the attractiveness of the book.

(239)

240 CALIFORNIA FISH AND GAME

The ainhor orii-ny uiscu^ses a common undesirable practice known among taxon-
<«niis:T* as "STtlitting". She then unfortimatelv (or nnwisely) proceeds to use a
system of classific-ation that is sadly lacking in logic. Using this system of classi-
fication <tesed only upton shell characters! one would have to give a different
generic name to opposite sexes of one of the near relatives of the cowries found
in Californian waters. The same awkward situation is unquestionably true in other
ktcalities. Only when cx^wry devotees look beneath the shells, and perform critical
studies on tie animals that live within, will this problem be resolved.

The index is arranged alphabetically so it is not difficult to locate a page refer-
ence for any species of cxiwry regardless of the genus to which it is assigned. All
in all this volume is a worth-while addition to the library of any shell collector,
whether amateur or pixjfessional. — John E. Fii<-h, CaVifornla Department of Fish
o«'<f Gusae.

Duck Shooting Along the Atlantic Tidewater

Edited by EugeJie T. Connett : William Morrow and Co., Xew York, 1947 ; xii-f-
o(»S p., illustrated with photographs and drawings, and color plates by Dr.
Edgar Burke and Lynn Bogue Hunt. -$ir>.

This c<«mprehensive work is the pntduct of 16 prominent sportsmen who wrote
more or less indei»endently. Each chapter has to do with a particular section of the
Atlantic seaboard from Xew Brunswick to Florid:^ and the Louisiana marshes.

The pturpose of the bc»ok may best be descril>ed by quoting from the preface:
"^^Teral years ago L»r. Edgar Burke asked me if I would join him in writing a
IxKtk on i>uck Shooting, saying before many more years elapsed few of the men
who actually witnessed those memorable old days of market gunning, and the years
of sufterb shooting which followed them would be with us. He felt that the time had
come to gather as much authentic information of those bygone days as possible and
make a wrmanent recxtrd of them for future generations of duck shooters."

This bttctk is written expressly for the layman, contains a minimum of scientific
terms, and is easy to read. It is printed in the large 8- by 11-inch size, which make
the iUusIratioiiis more appealing to the eye. The 12 reproductions in full color of
the paintings by Lynn Bogue Hunt and Dr. Burke are superb. It also includes 137
black-and-white plates, falling in two categories: (1) photographs on waterfowl
contributed by the Fish and Wildlife Service and ( 2 i photographs on the sh(>oting
aspect, hunter success, decoys, blinds, etc. The ithotographs by the Fish and Wild-
life Service are excellent in showing waterfowl in their native habitat, but are
just added to the Ixtok for interest and do not follow along with the continuity of
the story.

The first chapter. "Future of Waterfowl", by Fredrick C. Lincoln, and the last
chapter. "Waterfowl", by Ludlow Griscom, add greatly to this edition because both
of them stress conservation and give even the average reader an inkling there is
J, - " ,vl than just shooting them. The ixdnt is brought out time and time

fj _ are other influenc-es on waterfowl populations, such as : disease,

t: . - habits, migration, etc.

^ , , ,. ;,. „ -: ccdorful and descriptive chapter is the one by Dr. Burke on
"Making and Painting L^ecoys". Dr. Burke believes that today's factory made out-
T- " " ' 'n quality. If the quality is acceptable, the price

i^ , ;: -- on is to make one's own decoys. A very good

3iess<tm in dec<:<y making l^tiiMW:^.

T> :, '.■K'k should be of interest to the gener.-il miblic. — Jnrl- R, Slogson, California
I' »i of Fiih aitd Game.

A.r.;;- ••e;-;- Bird Guide

By Richard H. Pough : 1 y & Company. Inc., New York. 1907: xxxvi-|-

3H) p« 32 c-'-"' ■'■ iT-^^ Ef-kelberrjr, line drawings liv T^rry M. Shortt.

The area covered in this rolame approximates some 2i million square miles in
^est- ^' -*h America, inclading the bordering w-eans. It includes the area west
of a wn from the Bering Strait and the Arctic Ocean south to the Mexican

t«(»rder. fiicH^«mpassing portions of the Dakotas. Nebraska. Kansas. Oklahoma, and
Texas. Alth<iUgh 614 kinds of birds are known to oc-cur within this area, only 203
at* eidusiTely western and it is these 203 to which the bulk of attention is given.
The other 411 species, migrants and water birds, were described in two previous
works by the author and only their names and ranges are included here.

REVIEWS 241

The birds are listed in tL _ ^ _ . . :_ ;

the American Orintholosists L

For the 203 species dis^" - _-,.:.-, ,_ „,i--- >_ ; ,.

amount of up-to-date infi>r:

been presented. These birds ar-r .rXa-" -

tions and black-and-whi*e ---r f i":r-i

Of general interest .
other large geographica - . ^..j-- -.:_. .

index appears to be co:

This volume, as well ai tat liv.j yi^

Guide and Audubon Water Bird Gu: .. \„:

Society. — John E. Fitch, California Department of Fish and Game.

The Book of Flowering Trees and Shrubs

By Stanley B. Whitehead : Fre ' ^arne & Co., Ltd., Xe\r York, 1956 ; 244

p., 63 color plates by Joan Ll._ -53.50.

Gardeners in Calif'Tni-i and ^^IsAwher^ have ac-cess to descriptive litera"
ing a wide variety es. However, there alwaj -

big question, what . . . . .. .. color? The Book of Flov, . .

and Shrubs answers this important question. A total of 248 varieties of ■ ver-

ing plants is oorre- -' - ■ - -

soil and exp^-sure :

the garden. ; . _

Western _>.. . » . .....„- .......... ..... .: ... ^. ......_._ „_-_ :. .-

able to draw upon the vast experience of gardeners in other parts of the world- we
shnuld take ' " ' . -^^ • 7 ■ ' " , is a st- " - ira-

tion to prv- . - to c» ; - H.

Toung, California Department of Fi^h and Game.

The Ornithologisfs' Guide

Edited by Major ' ■ ' H. P. W. Hutson : Philosophical Library. Inc.. New York,

1956: XX — 27 ■. figs. SIO.

This publication is a compilation of 67 shnrr articles, t'ar-h ■written by riue of 46
authors and e<i' V ■ .

professional or: . - , _ -

in England or her 1 - - ns.

The introduction .... u.^ editor -.- '^ -' - •-'•■; 1 -^ ;. v 1 . — .3 .

encourage the study of birds by :
tunities for ~ ' " . _

Special *-: - ~ ~

tional bird watchers. Many good - . _ ns are given on note taking. -

niques. selection -' • -■ - ...,•;, - _• studies, bir' - ^- -

of photography. ns. nest o< .

vices, bird study w.

This book, cover - . -

study, is necessarily g> its over-a^.

ten and concise, but btv.i.-i;^■ ..-f their bre.- ,■ ■^■- ...-. _.
tion useful to the scientific worker.

One section will ■ • . • ■

of information friji ~ . -

referenc-es, omithologicai societie> a. locations of extensive eoUeciions. and

some of the leadin? ••'■' "^ -^logists •._ -' - ...^.---..^-c

The only regret • viewer is t. nforma-

tiou and references ui. oird wat' ' _ :.ei^u which

would have been of some aid to f . - . ing on this

continent.

Technicians in this country may disagree w -:..: _.-^;- on theories

and techniques, but on the whole ""The Or; . ie" is a book recom-

m^-nded to the amateur ornithologist. — "" ' ■ ' - Ifornia Department of

Fi-gh and Game.

242 CALIFORNIA FISH AND GAME

Fishes: A Guide to Familiar American Species

J>y Herbert >S. Ziiu and Hurst H. Shoemaker; Simoii aud Schuster, New York,
1956 ; IGO p., 278 color drawings. Limp bound $1, de luxe cloth $1.95.

Only four inches wide and six inches high, this little book is another addition
to the familiar Golden Nature Guide series. Between the covers (be sure to buy
the de luxe cloth edition — you'll want this booklet on hand for a long time) in
realistic color and often amid native surroundings are 278 species representing most
of the families of American bony fishes. Notes on sharks and rays, lampreys and
hagfishes, and animals mistaken for fishes are also presented. There is, refreshingly
enough, no added emphasis on game fishes. Some of the remaining short, but direct
and informative, sections include Parts of Fishes, Adaptations, Scales and Tails,
Origin and Development of Fishes, and Amateur Activities for Fish Study. The
subject is neatly wrapped up in the last few pages by a very adequate index, an
index to scientific names, a list of the larger public aquaria and biological stations,
and a bibliography of seven nontechnical books about fishes for further reading.

This guide answers the need for an inexpensive, general guide to fishes, both
marine and freshwater, written in popular style. Much of the information on the
habits of fishes is submerged in dry, scientific documents. One has to pluck the
interesting tidbits from among chi-squares, holotypes, and angler-hours; few
readers have the time or patience. Such labors even tax the fidelity of some fishery
scientists.

The numerous books on angling assist but little, since they are concerned mainly
with knots and gadgets and commonly insert, at best, only scattered, inaccurate
notes on fish life history. Semipopular works such as "The Life Story of the Fish"
by Brian Curtis and "A Study of Fish" by Chapman Pincher, while more than
worthwhile for those who can afford them, lack the full-color drawings found in
this guide.

Skin divers and aquarists will furnish the main audience for this guide. Sales
should continue strong, since both these groups are growing and numerically are
probably equal to or greater than the several million bird watchers in the country,
and bird guides do sell. Relatively few anglers will be interested. Science teachers
and students and members and employees of conservation organizations both public
and private will provide another large outlet. — Almo J. Cordone, California Depart-
ment of Fish and Game.

Animal Agents and Vectors of Human Disease

By Ernest Carroll Faust; Lea and Febiger, Philadelphia, 1955; 660 p., 216 text
figures, and 9 plates, including one in color. $9.75.

It is quite evident that the distinguished Doctor Faust is a professor of para-
sitology, in that his book devotes 428 pages to the endoparasites and 63 pages to
the ectoparasites. There are eight sections, with titles as follows: General Infor-
mation and Orientation; Protozoan Agents of Disease; Roundworms (Nematodes) ;
Tapeworms and Flukes ; Other Helminths Parasitizing Man ; Arthropods as Agents
and Vectors; Other Invertebrate and Vertebrate Agents of Disease; and Technical
Aids. The book contains many useful tables, including ones on the natural history
of the more important endoparasites ; pathology and clinical manifestations pro-
duced l)y animal agents of disease; diagnostic examination procedures; chemo-
therapeutic procedures ; differential characteristics of protozoan parasites ; arthro-
pods as vectors of human pathogens; and snakes harmful to man. The excellent
illustrations and plates that have appeared in the author's other publications are
liberally used thoughout.

To this reviewer it appears that Doctor Faust has summarized the most im-
portant contributions of his authoritative books on clinical parasitology and hiiman
helminthology, and combined them in this book with a small addendum of miscel-
lany. In the condensation he has sacrificed some of the morphological characteristics
of the endoparasites, and as a result the facility of identification is somewhat im-
paired. Some of the uncommon helminths have been omitted.

The entomological section is not exhaustive, but is rather adequately covered in
a narrative sense.

The miscellany includes a chapter on the urticating marine invertebrates, and a
chapter on veniferous reptiles. A short chapter of three pages deals with the role
of fishes in the discomfitures of mankind. Specifically, there are discussions of food
poisoning through spoilage of fish, poisoning through the ingestion of toxins present

REVIEWS 243

ill the reproductive organs or blood of certain piscine species, and mechanical and
poisonous injury from forcible contact with ectodermal spines. Finally, there
is a chapter on certain mammals harmful to man. Mentioned in passing are
the mechanical injuries that might be inflicted by dogs, cats, bulls, elephants,
etc. ; however, descriptions are given of the duck-billed platypus and its venom
apparatus, and the vampire bat's proclivity for human blood, with resultant anemia
in the victim.

The last section of the book involves descriptive material on laboratory diagnosis,
culture technics, seriological procedures for diagnosis, and the preservation of
arthropods, mollu.sks, and vertebrates.

The limitations of the book may be illustrated by the following, taken from the
identification of specimens. "The time has long since passed when it is sufficient
to identify an arthropod as a tick, mite, fly, or mosquito." As an alternative, the
author suggests addressing a letter to the Curator of Arthropods, U. S. National
Museum, Washington, D. C, for instructions on how to ship the specimen, and
then proceeding to follow instructions received. In the preface the author stated
that the contents would be useful and stimulating to the zoologist, laboratory diag-
nostician, and public health worker, and others.

The viral, richettsial, and bacterial diseases are considered in 22 pages, with some
rather important omissions. The vampire bat is mentioned as a possible transmitter
of rabies, luit the insectivorous bat, dog, fox, skunk, and others are ignored. The
book discusses many invertebrates and vertebrates, but other than a brief mention
of birds in connection with the encephalitides, the members of the class Aves are
ignored, along with their role as vectors of disease, e.g., ornithosis. In the preface
it was written that the book would be useful to the epidemiologist, yet the entire
field of the zoonoses is minimized or absent.

The undergraduate student and some cliniticians will find the book bandy for
reference, particularly in using the excellent tables contained therein. — Merion N.
Rosen, California Department of Fish and Game.

printed in California state printing oftice
54385 4-57 5,200