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Canyon mnule deer imerd was believed! to imave commtained! 200 immd!i- viduals. (Vieira et al., 1994) .... Grand. Island,. New. York,. USA). Samples were centrifuged at. 340. X G, at. -20. C for 15 mm. The ..... imm Yellowstone. National. Park.
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j(flifl1il

INFECTIOUS

KERATOCONJUNCTIVITIS

MULE

(ODOCOILEUS

DEER

ZION

NATIONAL

Sharon

K. Taylor,14

Victor

Sharon L. Fedorchak,2 Ralph E. Moore2 National

Park

P.O. Box 2 Resource

Service,

G. Vieira,2

and

pp

:326:35) 1996

ASMkiatu))

FROM

UTAH

Kenneth

Wildlife

37127 (WASO Management

I)96.

IN FREE-RANGING

HEMIONUS)

PARK,

:32(2) l)isras,

lhsaws, © \\iI(IliI(

Elizabeth

S. Williams,3

W. Mills,3Jackie

Vegetation

Pilkington,2 M. Boerger-Fields,3

Rupert

L. Cavender,3

Amy

and

Division,

490), Washington, D.C. 20013, USA Division, Zion National Park, Spnngdale,

Utah

Department of Veterinary Sciences, University of Wyoming, 1174 Snowy Range Road, Laramie, Wyoming 82070, USA Current Address: Department of Veterinary Sciences, University 1174 Snowy Range Road, Laramie, Wyoming 82070, USA

84767,

USA

of Wyoming,

Aim epizootic of in fectiomms keratocoumjumrmctivitis (1K) was stumdied opportmmmmisticahly iii mimimle (leer (Odocoileus /me;nionu.s) frommm Zion Natiommal Park, Utalm (USA), frommm Novemumber 1992 to Marclm 1994. Moraxella sp. and Chiamydia sp. were isolated! frommi tIme conjuimmctiva of two of sevemm deer. 1mm additiomm, Thelazia calzfornien.si.s occurred oum time commjummctivas of six of deer. Basedl omm field observations, adumlts appeared to he affected climmically at a Imighier imicidemmce dhmrmmmg botlm years as opposed to jumvemmiles. Conical opacity was time mmmost apparent chimmical sigmm from 1992 to 1993. However, iii time following year, blepimarospasmn ammdl epipimora were mmoted mimore oftemm. \Ve were also able to documment time chimmical recovery of tlmree affected deer. 1mm additiomm, Moraxella sp. was recovered from time eyes of a climmicahly unaffected (leer 1 A I3STRA(

T:

free-rammgiumg

year

after

Ke,, calzfi)nz

the

epizootic

words: h’nsis,

occurred.

Infectious

pinkeye, Zion

keratocommjummctivitis,

mmmi lie deer,

Odocoilen.s

hemionu.s,

INTRODUCTION

as

mule

Park,

deer,

n ian us), I mmfectious kmmowum

keratoconjunctivitis

as

pinkeye

1)eemm reported

imas

mmmost

1mm cattle, aumd

tagmouls

imm time

imave

1979).

etiological

Nei.s’seria si)., mimycoplasmmmas,

5k).,

sp.),

(Thelazia Few are

sp.) of

1K

docummimemmted.

tected!

because

by predators. dividumals

or

in

reports

smmmahl groups

much

i)een

viduals

ammd

mmemnatodes

appeared!

wildlife

involve animals,

of

easily

ranging

insuch

326

is

be

of

sustain

1994)

We

kin

from cammyon.

mnule

deer

tional

Park

1993

an!

deer

quantity

during 1993

describe

time to

and

poptilations

(Cuinaim epizootic in

hmerd

witimin wimmters

1994.

immd!i-

vegetation

keratoconjunctivitis mule

Time imerd

200

and!

is

wide, a road

time

adequate those

1992).

infectious

0.5

commtained!

al.,

ocwhichm

seeim

of

imave

et

epizootic

and

be

time

(USA)

Canyon

long

can

iii

Utaim

Time Zion

lengtim

to

to to

located!

Canyon

(Vieira

ningimanm,

unde-

only

area time

believed!

big-

populttion

of

Park’s

Zion

qulahity

takemm

the

wintering was

are

Park

37#{176}16’W).

runs

immor-

a large

1992).

5 kin

of

Chbamydia

of

corner the

alces)

amm indirect

canadensis)

al.,

virgi-

tibocapra

(Alces

decline

National

which

si).,

i)e of

et

in

(Moraxebla

often

may

(Ovis

(0.

keratoconjummcas

the

approximately is

have

free-ranging

animals Most

(Brown

1979).

Cases

simeep

curred

(Baptista,

bacteria

viruses,

horn

(1 12#{176}59’S,

time

prevalence

agents

(Baptista,

cases

summligimt,

cattle

Listeria

in

T/u’lazia

(An

in oose

imiiplicated

southwestermm

transfer

1mm cattle,

inchmding

iulil)licated,

dust,

between

was factor

Zion

deer

Infectious

tality (Meagimer

Predispos-

in yoummmg ammi mnals

Various

con-

tailed

spp.,

ammtehope

and 1982).

tivitis

during

1968).

mnechmanically

1972).

hmighmest

in

commmmon

mmmost

immcluded

americana),

(Wilcox,

is hmigimly

are

iumvolved

Adkins,

usumally

disease

(\Vilcox,

wimichm

organismmms and

time

white

pronghmormm

(Timorne,

livestock

world

mumommthms, especially

imummimid weatimer immg factors

time

epizootics

warmer

ammd! flies

of

also

keratitis

imm domestic

coummtries

1968).

(1K), vascular

amid!

spp., Moraxella Utaim.

Chiamydia

National

Our

a free-

Zion of

1992

objectives

Nato

TAYLOR

were

to

identify

characterize likely

ET AL-INFECTIOUS

the etiological

MATERIALS

epizootiology

KERATOCONJUNCTIVITIS

and

of

agents. AND

were

examined

for

lesions.

and

centrifuged

supernatant

was

Blood,

major organs (lung, liver, spleen, kidney, intestine, heart, skeletal muscle), and the head were collected and sent overnight mail to the University of Wyoming, Wyoming State Veterinary Laboratory (Laramie, Wyoming, USA). Upon arrival, further gross examinations were conducted. Tissues (lung, liver, spleen, kidney, intestine, heart, skeletal muscle) for histologic evalumation were fixed in 10% buffered formahin, embedded in paraffin, sectioned at 5 to 7 j.m, and stained with hematoxylin and eosin. We tested for caprine arthritis-encephalitis/ovine progressive pneumonia antibodies with an immunodiffusion test (Veterinary Diagnostic Technology, Wheat Ridge, Colorado, USA). For bacterial isolations, conjunctival swabs were streaked on Columbia blood agar and MacConkey agar (Becton Dickinson Microbiology Systems, Cockeysvihle, Maryland, USA) and incubated in 5% CO2 at 35 C for 48 hr. The identification techniques of Carter and Cole (1990) were used to identify the bacteria. Chla;nydia spp. isolations were attempted using minced cornea and conjunctiva placed in Bovarnick’s medium (Bovarnick et al., 1950) containing 10 p.g/ml gentamicin (Gibco Laboratories, Life Technologies, Inc., Grand Island, New York, USA). Samples were centrifuged at 340 X G, at -20 C for 15 mm. The supernatant

well

plate

MULE

with

mixed

and

attempted isolation. C and inoculated

METHODS

gross

decanted medium

suiting

The first mule deer having vision difficulty in Zion National Park was observed on 20 November 1992. Organized field observations of the deer for clinical signs began on 14 Decemmiher 1992 and continued through March 1994 with a break during the summer when deer dispersed out of time canyon. The observation process entailed two resource managers traveling up and down the 5 km long Zion Canyon road and using binoculars to observe the deer for clinical signs. Over 2,600 individual deer observational reports were made. Observations for each deer included: date, sex, age (adult or juvenile), location, eye lesions (normal, epiphora, blepharospasm, cornea1 opacity), eye affected (unilateral or bilateral) and, when possible, individual animal identification (antler points and configuration, and torn ears). Documenting the course of the clinical signs in the freeranging mule deer resulted in many of the deer being counted on consecutive days. Seven deer (one aduilt female, three adult males, one juvenile female, and two juvenile males) in poor body condition were shot. Carcasses

was

IN FREE-RANGING

an as

eqimal

before.

stored

at

327

volume Thme

-70

Samples were onto dumphicate

(Corning,

DEER

C

reumntil

thawed at 37 wells of a 24

Cambridge,

Massachu-

USA) containing CRL 1696 McCoy cells (American Type Culture Collection, Rockvihle, Maryland) with 12 mm cover glasses. Wells containing samples were centrifuged at 2126 X G, 25 C, for 60 mm, followed by incumbation with minimnummim essential medium witim Earles salts and nonessential amino acids (Gibco Labsetts,

oratories)

containing

(Hyclone

Laboratories,

37

C

in

5%

examined

10%

CO2 daily

fetal Inc.,

7 days.

for for

bovine

Utahm)

McCoy

cytopatimic

serumni

Logan,

cells

effect

and

at

were stained

at 7 days postinoculation withm a direct flumorescent antibody stain (Cultureset Chilanmydia Identification Reagent, Ortho Diagnostic Systems, Raritan, New Jersey, USA) to (letect Chiamydia spp. Infectious bovine riminotrachitis (IBR) isolations were attemmipted using minced cornea and conjunctiva placed in Bovarnicks mediuumm (Bovarnick et al., 1950). Samimples were centrifumged at 340 X C, at -20 C for 15 mimin. Sumpernatant

was

equal before.

volume of Resulting

-70

C

until

decanted

and

medium and supernatant

attempted

isolation.

mixed

with

an

centrifuged was stored

as at

Samples

were

thawed at 37 C and Mediumm 199 with Earles salts (Gibco Laboratories) containing 2% fetal bovine serumim (Sigma Chemical Co, St. Loumis, Missouri, USA) to control viruses. We added! BHV (bovine herpes virus) type I (National Veterinary Service Laboratory (NVSL), Ammies, Iowa, USA) to confluent monolayers of fetal bovine cells (Monfort Biologicals, Greehy, Colorado) in a 48 well plate (Corning). Cells inoculated with the samimples then were incubated at 37 C, 5% CO2 for 2 days at which time thmey were passed into tissue cumltumres. Thmey then were observed daily for 7 days with an inverted light

mimicroscope.

If

cytopathic

effects

were

noted, the cells were stained withm IBR flumorescein isothiocyanate conjugated antiserum (FITC) (NVSL) for the presence of IBR antigen. Direct flourscent antibody staining of conjunctival

and

conical

sections

for

IBR

and

Chiamydia sp. was conducted by placing 1 mm2 of tissume on to a cryostat chuck withm OTC embeddment compound (Miles Inc., Elkhart, Indiana, USA). The chuck then was placed in the cryostat at -10 C. Tissues were cut immto 4 pmii sections, placed on slides, fixed in acetone for 5 mm and then allowed to air dry. We added IBR FITC conjugated antiserum (NVSL) to half of these sections, and Chlainydia psittaci FTIC conjumgated antiserumm (NVSL) to time otim-

328

JOURNAL

TABI.E

1.

winters

of

OF WILDLIFE

DISEASES,

()bservatiomms

of

to 1993

and

1992

Number

deer

mule 1993

to

in Zion

National

1996

Park

affected

with

of

(leer

Number

observed

deer

Percent

of

affected

of all

deer

keratoconjuimmctivitis,

Percent

Percent of all juveniles

of all

adults

affected

affected

affected

1993

1)ecember

13

314

9

3

January

19

525

66

8 8

184

i3

218

9

13 7 4

Febniarv March Total

1993

infectious

1994.

Number

days

to

32, NO. 2, APRIL

of

observation

1992

VOL.

48

11 31

0 1 3 i

35

23

1,241

1994

to

November

6

I)eceinber

11 9

289 393 531

29 49 62

10 13 12

25 32 36

5 5

5

242

24

10

24

2

January February Total

37

C

in

applied

All sections 5%

and

icaily

1.455

31

er sectioums.

for

CO2

then

for

sections

positive

30

were

mm.

were

incubated

at

Coverslips

observed

were

microscop-

particles

(C.

from four of and examined microscopy

Hearne,

pers.

the by for

comm.).

tive

by

specimmmens

tional USA)

Parasite tinder

Kofoid

et

were

al.

(1937).

deposited

Collection accession

time

Clinically, years

a Imigimer (Table

adults

U.S.

Na-

(Beltsville, Maryland, number USNPC 85460.

were

prevalence 1).

No

significant

affected

juveniles difference

both

deer

Corneal

125

with

eyes

and

epiphora

the

following

more

opacity

was

in

also

able

to

of three

In from

recovered

izootic in

the

the

164

affected,

in 82, in

of

of

epiphora

36.

the

clinical

that

had!

in

We

were

recovery identifiable

Moraxebba

the deer

eye

surface

1 yr

after

sp. of the

a

ep-

occurred.

six

of

moderate date

and the

keratoconjunctivitis seven

deer

amounts

were focal

hyperemia

at

vascularization

the

noted Mild

to

mucopurulent (Thebazia

on

areas

was necropsied.

of

nematodes present

affected

with and

53 in

signs and

addition,

unaffected

in in

became

clinically

deer

fre-

However,

clinical

document

was

most

occurring

eyes.

be

the deer

observers

the

characteristics. clinically

30

opacity

affected

of

noted

occurred conical

we and

year

time

changed to

and 1993.

affected sign

as

trend

observed and

1994),

blephraspasm

blepharospasm 67,

to first

194

with

this

eyes

time

year,

familiar

disease

to

unilaterally

clinical

eyes

47

1992

(1993

found

reported

were

in

affected

During we

quently

signs

from

winter deer

bilaterally.

eyes.

of observed

following

epizootic,

of affected

Clinical unilaterally

50

70

number

females.

occur

in

observed

sis)

timan

to

The

the

or

Bilateral

RESULTS

at

between

males

bilaterally

Representain

noted

adult

94

Tissimes were ground with a mumortar and pestle, diluted with distilled water and centrifuiged at 3,024 X C for 20 mm. The supernatant was timen cemmtrifimged for 1 hr at 39,191 X C. The pellet was resumspended in 10 drops of distilled water. One drop of the suspension was added to a mixture of the following ingredients: 20 drops of distilled water, four drops of 4% phospimotimngstic acid (Electron Microscopy Scieimces, Ft. Washington, Pennsylvania, USA) and drop of 0.1% bovine senmmmi albumin (GibCo Laboratories). The mixture was nebuhized omm to collodion-coated 300 mmmesh copper grids. Time immaterial on the grids was then examimined witim a Philips 410 LS transmission electron mmiicroscope (Philips Electronic Instnmments, Mahwaim, New Jersey) for time presence of virus-hike particles at 60 KV Ocuilar parasites were mmmounted in Hoyer’s mmmediimmn (Pritchard and Knmse, 1982) and identified based on morphological characteristics described

was

observed

flumorescence.

and conjunctivas deer were randomly selected mmegative contrast stain electron Cormmeas

virus-like

3

the

Corneas

of

opaqueness were

calfornien-

conjunctiva

deer.

limbus.

exuof

were Focal present.

and

six

cloudy slight

ulcerations

TAYLOR

2. Laboratory keratoconjunctivitis.

TABLE

tious

ET AL-INFECTIOUS

results

seven

from

Age

Zion

National

Electron micros-

IBR Sex

KERATOCONJUNCTIVITIS

Park

IN FREE-RANGING

mule

deer

affected

MULE

DEER

withm clinical

sigims

Bacterial isolate

FM

copy

isolate

Adult

Negb

Neg

Pos

Proteus

Male

Adult

Neg

ND

Neg

Pseudomonas

Ocular parasites

Thelazia

calfirnien.si.s

Thelazia

calfirniensis

sp.

sp.

Staphylococcus

sp.

Streptococcus

sp.

Male

Adult

Sus

Neg

Neg

Proteus

sp

Tijelazia

calfirnien.sis

Male

Adult

Sus

Neg

Neg

Proteus

sp.

T/zelazia

calfin-nien.si.u

Thelazia

calfirnien.sis

Escherichia Female

Juvenile

Neg

ND

Neg

Moraxella

Male

Juvenile

Neg

ND

Pos

No

Male

Juvenile

Neg

Neg

Neg

Proteus

Fluorescent

b Neg,

antibody

negative;

Pos,

test

for infections

positive;

Stis.

Microscopically, sentially

with

cuffed

were

by

was

phocyte

cells

and

epithehium.

ed

by

phils, and

iris,

normal.

juvenile

stroma

and in two

of to

female

the

ocular

Moraxelba

both

eyes

rescent

sp.

of

one

of

deer,

were

However,

negative

adult

of

two

female.

fluorescent

Two

antibody however

from

any

of

IBR the

nega-

agents.

2).

isolated

a juvenile deer

male

were

we

viruses seven

IBR were

deer.

No

negative.

male

on anti-

the

mule Forest,

1944).

Affected eyes,

to

signs.

mule

deer

Platte

River

cular

keratitis

isolat-

however

virus-

ported.

no

drainage, annually diagnostic

the

agent

and in

Ummfor-

nearby

young

not

idlelitiwere as

re-

severe

Wimmter

(1956)

antlered

male

upper

Nortim

the Wyoming from

pu-

condition was

but

Honess ranging

to

and

blindness.

similar, that

time Beathm,

reddish

with

located

show

reported

had

etiological

imm Bow

during

complete

cattle

epizo-

Medicine

ulceration,

exudate

Many

time

and

deer

to no

the

was

a chimmi-

reported!

(Rosenfeld conical

ocular

also

after

Wyoming,

1943

progressing

sp. of

was in

of

etiolog-

surface

1 yr

litwere

recovery

identify

Moraxella eye

deer

opaque

clinical

and

addition, deer

of

sp.

to

time we

clinical

Blindness

spring

fled.

that

able

smaller

in

exception the

In

withm

reported

were

unaffected

ported

from

time

occurred.

all

suspect for

all

consistent

document

tunately,

sec-

from

Chbamydia was

Fluo-

frozen

were

from

rulent frommm

conjunctiva

testing

no of

ical

10

Escherichia

(Table

sp. deer,

and

or eye

recovered

for

Chiamydia

eyes

and

was

and

to

deer

otic

gross

(coagulase

Se-

aimtibodies pro-

were

studies

able

time

Pseudomonas

staining

cornea

results with

cally

mule

Only

sp. juvenile

antibody

tions

male

sp.

for

DISCUSSION

recovered

con-

from

sp.,

electromm

tested.

arthritis-encephalitis/ovine

National

Proteus

Streptococcus

coli.

and

obtained

Staphylococcus

tive),

body

significant

deer

tests

pneumonia

erature

lesions.

isolates

included:

caprine

Our

essentially

adult

no

to

by

four

immunodiffusion

observational

invad-

cihiary

resolving.

observed time

lym-

melanocytes.

were

were from

gressive

neutro-

corneas

be

had

Bacterial swabs

lens

particles

by

by

a few

the

the

of

like

beneath

nerve,

calfirnien.sis

Thelazia

microscopy

conical

was

infiltrated and

retina,

histologic

ed

neutrophmils

optic

appeared

gens;

infiltrated

the

of

deer

an

areas

and

Lesions

junctivas

the

with

cells,

of

The

a few

Neg

sp.

rum

of

intact

and

plasma

Sections

sp.,

thickening limbus.

capillaries

es-

Corneas

Limbal

sp.

growthm

done.

vessels

the

aggregation,

plasma the

by

along

epithelium

blood

lymphocytes.

characterized

epithelium

not

were

some

coli

rhinotrachites.

ND,

conjunctivas

normal

lightly

bovine

suspect;

infec-

of

Clila-

mydia

Female

a

329

imad 1943

informiiatiomm

to

vas1955;

was

re-

JOURNAL

330

OF WILDLIFE

Thebazia (!eer

calzf()rnien.sis

inimabiting

tiommal

Parks,

immals

severe

hmad

they as

time

Park

mmmule deer

frommm

Ziomm

dhspersed!

Canyon,

5pretdl immto time imormm shmeep (0. had

beemm

of curred

to

canadcn.s’i.s’

1982,

Mouimtain

imm Yellowstone

National

dlirect

was

of

mortality

time

ever,

hmerd as

h)igimorn been

July in

observed

imas

Park

in

mmmonly occur izootic

clinical

signs

have

in domestic

1K nor

the

thirough

Time

cooler

Marcim,

No

livestock Utah’s or

epizootics

imm the

Ziomm

Departnment

Departmiment

of

of of

area

I K

fly a de-

reported Resources

We tlmammk time resource mamiagers amid rangers at Ziomi National Park for extemmsive docummmientatiomm of field observations. We also thank C. Ilearime, C. Lynn, C. Stitim, ammd V. Welcim frommi time Wyomimmg State Veterinary Laboratory for teclmmmical expertise.

P

J.

jmmnctivitis-a 135:

225-242.

1979. review.

Infectious British

ix)vimme Veterimmary

1990.

Press,

Di-

bacteriology

Samm Diego,

Cal-

pp.

36-39. ANI)

A

in

N. C. VEALE. mmemnatode eye

man, witim a review of the Thelanimals. University of California 47:

Zoology

J.

W.

Park.

QUINN,

225-233. L.

AND

STACKHOUSE.

infectious

bigimorn sheep Journal of Wildlife

of

keratocon-

Yellowstone

Diseases

Na28:

171-

176. M.

C.

II., AND

collection

and

Ummiversity

of Nebraska

p.

0.

W.

preservation

KRUSE.

1982.

of animal Press,

Time

parasites.

Lincolmm,

Nebraska,

125.

ROSENFELD,

I.,

ANI)

panoplmthalmitis Managemnemmt E.

T.

1)iseases

0.

A. BEATH.

in deer.

The

1944.

Contagiouis

Jouirmmal of Wildlife

8: 247-250. 1982. of

Immfectious wildlife

in

keratocommjumctivitis. Wyomimmg,

E.

T. Timor-

ime, N. Kimigstoim, W. R. Jolley, and R. C. Bergstrom (eds.). Wyoming Came and Fish Department, Cheyenmme, Wyoming, pp. 81-84. VIEIRA, V., R. MORE, J. BURNS, S. FEDORCHAK, D. SIDLES, AND L. P. NAYL0R. 1994. Condition of Zion’s natuiral resoumrces. Zion National Park Resource Mammagement Plan. Zion National Park, Springdale. Utah, pp. 94. WIEA:ox,

keratocon-

in Research

(editors).

Chmlamydial-catmsed

C.

juinctivmtis:

l3iuprmsTA,

COLE

cal/riiiensis,

in

M.,

PRITCHARD,

In:

CITED

of Veterimmary

WILLIAMS,

of dog and

TII0RNE,

ACKNOWLEDGEMENTS

L.

of domestic

tional

Livestock.

LITERATURE

auturnnalis

imm veterinary

Wyoming,

jummctivitis

doimmestic

were

Wildllife

Relationship

of keratoconjunctivitis

J. R.

AND

Thelazia

1992.

mimonthis

in

1972. (Musca

fly

Journal

A., 0.

MEAGHER,

ep-

wimen

face

procedures

Publications

corn-

populttiomms were not active amid (luring 1)eriod whmerm imours of sunhigimt were creased!.

R.,

C.

azia

National that

livestock.

(luring

C.

worm

Zion

of

2551-2555.

Cheyemmne.

keratocon-

at

activity

acids,

of rickettsiae.

59: 509-522.

to production

1937.

time trends

stability

T. H. ADKIN5.

American

KoFomD,

occurred.

mmmuhe deer

occurred

diesert

of

infectious

follow

In60%

Park

omm the

SNYDER.

amino

B. 1992. Diet and disease of Zion Mule Deer, Zion National Park, Utah. Park Service/Cooperative Natiommal Park Study Unit, Contribution number CPSU/UNLV 048/02. Ummiversity of Nevada, Las Vegas, Nevada, 104 pp. LIONESS, R. F., AND K. B. WINTER. 1956. Vascular keratitis, pammophthahinitis. In Diseases of wildlife in Wyoming, R. F. Hones and K. B. Winter (eds.). Wyomiming Canme an(l Fisim Department,

How-

time

feeding

salts,

Canyon National Resources

oc-

be

1992). of

Natiommal

of

Novemnber

of

al.,

immortality

(lid! not

to

AND

J. C.

ANt)

CUNNINGhAM,

sheep

(USA).

none

Zion

epizootic

jummctivitis

et

with

significammt

Time

Park

1994,

sheep

bigimorn

estimmiated

(Meagimer

of

1K

MILLER,

ofcertain

and rmmycology. Academic ifornia, pp. 165-176.

winter

of

F.,

agnostic

bigwimicim

time

epizootic

J.

33:

might

neb.s’onz)

ammd proteins

CARTER,

spring

desert

Dimring

an

Rocky

iii

of

sugars,

calves.

timat

immfection

imerd

TIme influence

DeGeer)

3 wk.

imm time

time

nearby

reimmtroduced.

1981

2 to

1950.

BROWN,

but

concenmed

was

R., J. C.

M.

BOVARNICK,

of

blindness within

staff

1996

Jouirmmal of Bacteriology

ani-

immflamnmation,

recovered!

Zion

1934

Timese

temporary

chirmicahly Time

Na-

during

1944).

ocular

produced

in

Yosemite

(USA)

Beathm,

32, NO. 2, APRIL

reported

and!

Cahiformmia amid

VOL.

was

Sequoia

(Rosemmfeld whmichm

DISEASES,

E.

1968.

Infectious

A review.

The

bovine Veterinary

349-360.

Jourmmal Received

fir

publication

5 July

1995.

kerato-con-

Bulletin

38: