California. Dept. of Fish and Game.

California fish and game (Volume 18, 1932) online

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liv the writer. For the purposr of thr i it wa* nerrswiry

to shoot Hevernl (pjail during the elo*M><l s* ..,-.,. . ... ij«ni?i' '.• - •■•»;••.• wu
to take sainph'.s hiTf atitl there from oumfrous eovi-yn vrvre

operating in settled communiti<>fi wc alway.s made it a point to explain
to the raiichriii'ii what we w«'re doiuj? and why we wrre doing it and




*Scn Jost

• V,c*9'.''k


SoM D-^f* _ _x — - -

to ask permission to take the necessary samples for study. Altogether
>12 (niail wore examined. These included all of the species and sub-
species of the jjenus Lophortyr which are to be found in California,
incluilina: Santa Catalina Island. Of all of the quail examined, 45
iier cent were uarasitized. Parasitism in different lots varies from


to 100 per cent. No species or subspecies of the genus Lophortyx
■was found to be free from the parasites. The highest per cent of
infection, 100 per cent, was found in Gambel Quail at a private aviary.
The accompanying map shows the locations where collections were

It will be observed that only a few places did not yield parasitized
birds, and it is quite probable that more extensive collections made
there would have sho-\vn that these localities are not exceptions to the
general rule.


Since the Sergent's work (1907)^ in northern Africa had proved
experimentally that Haemoproteus columhae Celli and Sanfelice is
transmitted by the bite of the louse fly Lynchia maura, it seemed prob-
able that some species of louse fly would be found to be responsi?jle for
transmitting this newly discovered parasite of the valley quail in Cali-
fornia. Accordingly^, a survey was made of entomological collections
in California which might include such flies. From this study it was
ascertained that two species of such flies had been reported from quail
in the state. They were namely, Lynchia hirsuta Ferris and Stilhome-
topa impressa Bigot.

The former species had only recently been described by Ferris
(1927)^, and was represented by only three specimens in the collections
of Professor G. F. Ferris, of Stanford University. The latter was
represented by two specimens in the University of California collection.
With this information as a starting point, field search was instituted.
Guided by the knowledge that the louse fly of the pigeon deposits its
pupae about cracks and crevices and around nests in pigeon lofts,
examination was made of the quail pens at the State Game Farm in
the hopes of finding the flies or their pupae. Repeated examinations
made during May and June proved to be fruitless. The next step was
to drive the quail into cloth-screened pens at the game farm in the
hope of finding the flies on the birds. This likewise yielded negative
results. Operations were then extended to the field. Penned quail
were taken out into natural environments of the wild quail, left for
a few hours and then placed in cloth bags to be examined for any
flies that may have alighted on them. Wild quail were trapped and
similarly examined, but with entirely negative results. Jn the mean-
time, examinations of the roosting places and haunts of wild quail
were continued but no hypoboscid flies were found.

Following upon the failure to obtain flies or thr-ir pupae by any of
these methods, it became necessary to shoot quail and place them imme-
diately in gauze bags. These bags were made up into conical shape,
about a foot long. Field practice indicated that unless a quail could
be shot in an open clearing and placed in the gauze bag within a few
.seconds after it was killed, any flies present among the featliers of the
quail would make their escape. After a quail had been placed in a
bag, the bottom of the bag was closed with a rubber band. It was

oi ^o^^^ol!}' -^.^ ^/'^r, ^>^ Etudes sur les hematozoalres d' olseaux; Ann. Ist. Past.,
21, 2.J1-280, pl.s. 6-7, 5 flgs. in text.

» Ferris GF. Some American hippoboscidae (diptera pupipera) Canadian Ento-
mologist 59, 24G-2ol, 4 figs, in text.



tlipn hi'ld HU.Hpcndp*! by the aprx niul niiy fli**-^ •*■ ■• •* nt

ifiviiriahly cniwlf*! up into th«? t«p«rMnf part of t; r«j

Ik" the instinct of t\\v My to leave it.H hoxt iuimi*(liati>ly after the drath

of tin? lattrr. Afti-r the ttu's hml erawlni up iiitn tl

traiisffiriMl to L'L'inin. slu'il vials which wfrr i-an-f':"

baK aiitl tiiovcti upwards until thr t1i**.s hatl l»i><-ii it

Fio. 50. Lirnchia hirauta Ferris. Female, wings removed. After Ferris.

About 3S X.



fly taken by this method was captured on July 7, 1928, at the Whitman
Ranch, near Mount Diablo. By November this method had yielded 32
flies. The greatest number taken on any one quail was four. Many
had none at all and several only one or two. The only species taken
throughout the investigations was Lynchia hirsuta Ferris.

Collections of flies were made from several places in the vicinity of
Mount Diablo and at a few localities in the Berkeley Hills between
Berkeley and Orinda. When a fly was transferred to a vial, it was
customary to put the vial in one's shirt pocket in order to keep it
alive and warm until such time as it could be used in transmission

Fig. 51. Screen cage, used at Yountville Game Farm in a series of transmission

experiments with quail.

During the course of collecting flies, a few facts were gathered
which may be of assistance at some time in the future to anyone work-
ing out the life history of the fly, which is unknown. No flies were ever
found apart from their hosts. A pair of flies were observed in copula
October 5. Three different flies were observed to oviposit a pupa
immediately after being taken from quail and placed in vials. These
ovipositions took place July 7, August 2, and October 5. All of the
pupae were placed in a constant temperature box maintained at 35.5
degrees Centigrade. The one obtained August 2 gave rise to a fly in
thirty-one days. The other two failed to hatch. The longest time that
a fly lived in captivity was five days. They must live considerably
longer than this as will be apparent when consideration is given to
the development of Haemoproieus lophortyx in this fly.


I llt\N lire, liii\M-\ i|-, ii'ii'iit"" aii'i tti'ii" .' ' 'n

llii' wflltirr of tlifir luwl from wlnrh ili«-\ hu n

whirh they ohtaiii warmth. An will Im* m't-n in th<* ^'

Miif, thi'ir piii('h<*r-lik«? fret mablc them to hold fawt to the : m

thi'ir hosts ainl thrir Hat iMxIioji t-nahlr thdii to Klidi* una<T ibe

itlii-i's t'asil\'


I lirou^h till' cooperation of August Ha<l«-, Miiprrinti'iitlnit of thr
State (iam«' Farm, at Youiitvilii*. Uti youiik' quail were plar.<i with ih'ir
liroiMl.f ht'ii^i in sfn'fiKMJ ca^ri'.s iinmi'diatcly aft<-r hati-hint; <»n *' ''»,
l'.*L'>. At tilt' l)«'i;inninj; oi" tin* rxpt'rinicnl.s onr catfi* containtni ... ..e
little «inail and tin* otlirr fourteen. The most riu'id tart* was rxercijMHi
\>\ A \V. KIder and K. D. IMatt at the ^»ame farm to keep the.H«? cagt's'it Irif diiiinj; the time that thi* quail wcrf heini; kept for ujie io
the expfriniiiils. Sotue of the qujiil were renioveil from time to time
iiid transferred to smaller eajfes for the fly-hitin^r experiments. The of the ({uail were not removed from the sereenetl pens until Decem-

\'<T 7.

.\ lon^ series of transmission experiments was planned ainl earri«'«l

it. These be^'an by merely placing' an infeettnl and uninfectetl quail

I lom the screen caj;e in a small laboratory sereeneci caj^e about 6xlL'xl2

•lies tojjether with flies taken from t|uail shot in the Held. Scarcity

; Hies necessitated improving this proec«lure and eventually the fly

t'ltiii^ method adopted was to place the i\y in a sunill biting cup about

inni. in diameter and made from bobbinct ami gummed pa|>er tape

Nsliicli was scaled into position tlirectly ajrainst the skin of the bird.

The usual practice was to contiiie the tly over niijlit on th.* ImhIv of a

mail. When it became apjiarcnt that the scarcity of Hies iaij;ht pre-

nt a tlioroujih test by this method of the hyp<tlhesis that Lytuhia

rsutii Ferris was a transmittinjj best of the parasite, still more

luratc experiments were planned Wlu-n Hie.s were taken on para-

ti/cd qujiil, tiicy were subjiclcd to micro.scopic analysis to determine

wlicther the infective staj?e (sporozoite stajre) of the was

present in the in.sect. Three such infected Hies were found. One of

ihese taken on October ;"). 11)*JS. was heavily infei-ted, i' ' ' • is

lu'iiijr iroi-jrcd witli tlic .sporo/.oitcs of 1 1 lit inu pru( tU^ ^. ,.. . i.C

ilivary jrlands of this Hy when clu)pj)cd uj) in few cubic millimeters

nnniial saline solution and injected intravenously into the wing

\'in of a tpiail produced a positive case of transmis.sion of the | '•',

the infection ilcvelopinjr in about thirty ilays. The control tp. it

were kept in the orii;inal screened capes at the game farm r- .1

tree from infection with the parasite.


Ilaemoproteu!^ lophortifx is n sporozonn hnvin;: u cni:!,;! aied life
oydo similar to that of the causal organisms of human malaria which
is transmitted by the bites of mosquitoes. There are two distinct phases
in its life cycle, the sexual and the asexual phase. Trai. n is

biolo^rical. the jiarasitc jroinu: throutrh the .sexual part of its cycic ;n the
bodv of the ilv. A convoniont tlioiijjh perhaps not logical starting


point for discussing the life cycle is the gametocyte stage in the red
blood cells of the quail. When freshly drawn blood is placed between
a slide and a cover slip and examined under the microscope, it will be
noticed that the mature male and female gametocytes are at rest in the
red blood cells. Within a few minutes after the drop of blood is drawn
the male gametocytes escape from the blood cells and round up pre-
paratory to gamete formation. By a process of successive nuclear
divisions, the nucleus of the gametocyte forms the nuclei of from four
to six male gametes. In addition, to the nucleus, the male gamete is
provided with a small amount of cytoplasm. By rupture of the mem-
brane of the gametocyte these, gametes are liberated. They are ex-
tremely active vermicular organisms.

In the meantime, a single large macrogamete or female gamete is
being formed from each female gametocyte through the throwing off
of a polar cell. This process is a little slower than that of gamete
formation in the male gametocytes. After the gametes are formed,
fertilization takes place by the complete entrance of the male gamete
into the substance of the female gamete. The process requires but
fifteen seconds. As a result of this process a zygote is formed. About
twenty minutes later this zygote begins to elongate into a motile
ookinete. When this process occurs normally in the stomach of the
louse fly Lynchia hirsuta Ferris, the ookinete soon burrows through
the wall of the stomach and comes to rest on its outer surface, where
it develops into an oocyst.

Within the oocj^st, repeated nuclear divisions give rise to from 500
to 5000 sporozoites. The mature oocyst then ruptures liberating the
minute sporozoites into the haemacoele of the insect. These sporozoites
then enter the walls of the salivary glands and come to rest in the
central cavities of the tutular glands. When a fly thus infected bites
a quail some of these sporozoites are discharged and enter the blood
stream of the quail. From here they enter endothelial cells of the
blood capillaries of such organs as the lungs, liver and spleen. Through
repeated divisions numerous merozoites are produced. These in turn
produce more generations of merozoites in the tissues of the organs
mentioned above. Finally some of these merozoites give rise to game-
tocytes Avhich enter red blood cells and grow to maturity. When such
mature males and female gametocytes are present in the blood, the
bird becomes infective to another bird.


Numerous investigators have studied the pathological conditions
which are revealed by autopsies performed on infected birds. Few,
however, have paid any attention to the appearance and behavior of
infected birds and to the possibility that disease produced by Haem-
oproteus may be serious and even fatal. During the progress of this
work special attention has been paid to this phase of the studies. As
mentioned early in this article over 300 quail have been actually
handled. Hundreds of others have been observed in the field. As a
result of the observations on living quail, and of autopsies performed
on birds used in the investigations, four types or stages of Haem-
oproteus disease may be recognized. These are :



Tlio followiiii; (lia(;rain illtiNtrat«>H the tiiiw* .t'n.i-t.i invulved If '•>••
sexual (•yclc.

FlO. &2. I>ltiKra>n shuwInK the time i-lcmont Iru
fvrtilixaUoii aiul ot>klii«-tt< fonuatloii on the i:
111. nml» Kin"''tiicyte in blootl cell; lb, feiii
2b, gaini-tiK-yles out of blood cell anil r« •
tfaniftes foriHoil anil liberated ; 3b, Kanii-t
fertilization: 6, fortllliutlon lutinbrane r
elunKAtlns c>6kliiete ; 11-12. motile ookleicte.




1. Milil. chronii'. Tliis is by far the most I'oinmon, tlu* <iiiail exhib-
iting no outward symptoms of tlisoase and the presence of i- -^
in the blood lu-iiii; tlio only indii'Ution that the binl is j.i \.
This typo of infoi'tion is oxtremoly persistent, extending thr _ iit
the year. Birds thus infectetl are undoubtedly carriers although they
may manifest no disease symptoms themselves.

2. Mild, acute. This type apparently results from an initial infec-
tion tho m;isxnitude of which is sufTicient to cause the bird some dis-
comfort. It is restless and "otT feed" for from two to four days,
whereupon the infection is either tbrowu off or settles down to the
chronic tyix\




3. Moderate, clironic. This type is commonly observed under field
conditions. The bird is thin and anemic and more or less weakened.
Such an underlying condition might well account for losses of birds in
the field due to secondary causes, such as exposure or exhaustion fol-
lowing the nesting season.

4. Heavy, acute. This type occurs in late spring or early summer,
whether as the result of new infective fly bites or the lapse of an old
infection is not known. The percentage of red blood cells containing
gametocytes that have reached the pigment-producing stage may be
as high as 10 per cent and even more. The bird loses flesh rapidly, is
unable to fly, refuses food, becomes droopy and may die. Only four
fatal cases have actually been seen by the writer, and they are believed
to be rare. It must be remembered, however, that under natural field
conditions, the struggle for existence might eliminate birds before they







Fig. 53. Graph showing the relative percentages of quail harboring the
various classes of infections observed. Chart represents 100 birds,
sample population.

reach such a stage of helplessness. Several cases have been reported
from widely separated localities of quail that were unable to fly and
which could be caught with the hands. Four such cases, personally
investigated by the writer, showed the birds to be heavily infected
with Haemoproteus lophortyx. Two of these cases ended fatally and
post-mortem examinations disclosed no lesions other than those due to the
Haemoproteus infection. No acute fatal cases have ever been observed
in young birds less than six weeks old, but it is conceivable that such
cases do occur. If, for instance, the brooding birds should be heavily
infected and if they should harbor a large number of flies which trans-
mit the parasites, conditions for infecting the young would be ideal.


Hill anyone faniilinr with ll»«' hahjU of \ ' * - • -• "he

rluiiicis of llii(iinu' tin in wluTf liu-y may ; ^

HurroiHHlin^'s an* almost nt •^,'ll^fil)l^^ Thn fort thftl ; -d

at liiiuw farms and private aviaripM arc not found to In- '-d u

probably dm- to tin- cinMnnstanrrs undrr v.' > .^ - -,,j

raisrd apart from ihrir pannt.s, thus not I" . _ to

any lli«'.s wliii-li nuKlit be livinj? on tho adult birdM.

Ah to iliaunosiH of the disra.m* and postinorti-m
Mill* in tlif rrd blooil ci-lls of baltrrsliapid «-> i

black piymiMit d<*po*'itJi is «liarai-trri>tif. Tin- ni. m

l>anLsiti/t'd niay run from a fraction of om* por riMit to mon- than ten
per cent. An ahnost constant lesion is an enlarfjed bl -n

contrasted with tlie mucb sniailer pink spleen of the ikjiu. d.

Similar black pij^ment deposits may also be found in the In mJ

liver. Destruction of the red blood cells by tlie parasites brir it

an anemic condition of the bird. Hypertrophy and loss of ty of

the blood cells due to the |)reseiice of the parasites tenil.i In brinjf
about con^'estion in the smaller capillaries of t)i.- limgs.

In a case of a suc«-cs,sful experimental tra: on wherein a por-

tion of a luntr of an infect etl ipiail was transplanted to the Innly cavity
of a susceptible (piail. the artificially infected bird was at its 1« bb

just before the di'velopin^' j;amctocytes be«'ame visible in ii- ;.. . J.
This period of depression of the bird was marked by .severe anemia.
which threatened to eiul fatally before the bird rallied and
recovered. It is conceivable that such a heavy, acute infection could
be produced in a natural way by insect transmission and terminate
fatallv without anv LMinetocvtos bein«? discernible in the red blood

I'Vom reference to the life cycle of the p ' on pre-

vious pafjes, it is apparent that there is con-i.i. i..;,.. .»;.... of

luiiir and liver tissue with the multiplication of the n .■>'s

of tlie parasite in the endothelial and epithelial cells of these organs.
This, to};ether with any toxic etT«'cts that may be produce.: " of

the presence of the parasites, may account for the severity (^ .... ....i;al

infection of ^reat intensity.


As previously noted, pap* 23(5. infections nuiy vary fri»m extremely
lij^'ht to fatally acute. variations are probably due to several
causes, such as the number of parasitized Hies, variations in the viru-
lence of the parasites, and state of natural resistance of the host.

Transmission would seem to be fa«'ilitated by the observed fact that
ganu^toeytes may be found in the blootl of the quail throuphout the
year, and that flies have been reported from nearly all months of the
year. The ijrrej,Mrious habits of quail would also contribute to the ease
of transmi.ssion. Tendinj.? to limit the extent of the infection among
quail could be the small number of flies observed, the low percentage
of infections amonj; them and the licrhtness of such infections. It is
also possible that the flies are limited to certain portions of the quail
range, a subject upon which more data are needed.



For the most part the experimental work involved in this paper has
been done under controlled conditions where the various factors were
known and could be evaluated. In a state of nature, it is not always
possible to arrive at such precise values. For instance, one hears
reports that quail in certain partly protected areas do not increase, and
in fact become reduced in numbers. It would be of interest to make
most searching investigations into the actual field situation which gives
rise to the reports. Does full protection at first result in overcrowding
and may such a condition be followed by flare-ups of parasitism and
disease which defeat the purpose of protection? What is the situa-
tion with regard to parasitism of quail by Haemoproteus lophortyx
in this respect"? We have data showing that in some cases in private
aviaries the quail were 100 per cent infected. The highest percentage
of infected birds found in any one lot in a state of nature was about
65 per cent. Does the presence of heavily infected, weakened birds
react unfavorably upon the rest of the covey? Would such birds
under normal conditions be eliminated by predatory animals and cer-
tain species of raptorial birds? These are practical questions which
can not be answered except by the most painstaking comparative study
of field areas over a relatively long period of time.


In the foregoing pages an account has been given of the incidence
and pathogenicity of Haemoproteus lophortyx, a new species of blood
parasite of the California valley quail, Gamble quail and Catalina Island
quail. It is widespread in the state, having been found in numerous
localities. Fortunately, by far the greater number of cases seem to
be light, benign infections, showing that at least partial immunity is
well established. Fatal cases do occur, however, and it is possible that
Haemoproteus lophortyx may be the indirect cause of losses commonly
attributed to other causes. The parasite is transmitted from quail to
quail \)j the ectoparasitic louse fly Lynchia hirsuta Ferris.

Plates I and II in this paper are identical with those appearing in
a more technical account ^ published by the University of California.
This publication contains a complete bibliography.


Cxrateful acknowledgment is made to the many captains and depu-
ties whose assistance in the field was invaluable. Special credit is due to
Professor C. A. Kofoid of the University of California under whose
supervision the experimental and laboratory work was done.

To Mr. Eugene D. Bennet, Dr. H. C. Bryant, Mr. E. E. Ludlum,
Mr. D. D. McLean, Dr. H. Van Roekel, Dr. K. F. Meyer, Professor G. F.
Ferris, Professor AV. B. Herms and Professor E.*^ C. Van Dyke the
writer is indebted for cordial cooperation and assistance.

The painstaking care given the experimental birds at the State
Game Farm at Yountville by Superintendent August Bade and his
capable assistants as well as many other courtesies extended by Mr.
Bade have been greatly appreciated.

' O'Roke, Earl C. The morphology, transmission and life history of Haemoproteus
lophortyx O'Rokfe, a blood parasite of the California valley quail. Univ. of Calif Pub.
in Zool., Vol. 36, No. 1, 1930.





By JoHM flnwcsD

IT HAS |'.i;i:.\ SAID that lifr ana li"
IlH'lll. ( 'i)M.H« rVHlioll woikrrs will ; !»|f

that i(l<>a as it is uppamit at all tinu-M in < ii work and

the perpetuation of the wild life i.s eontiiiu'ent iipuii uit A-

ineiit. Civili/atinM is foiiHlantly rUfroai-liiii ' '' ly

orcu|)i('(i liy tish aixi uaiiie in uii(listiiri»Ml ( . h

to cxpeet tliat \\v may au'aiii have a eon«liti<)n an ri'lat«'«| ' -*,

when (s'l'cat hiM'ils of ilrrr were seen and "hiniN in Kueh it < thnt

their fli^'ht ohseurrtl thf smi." with " iiiit;ratinif fixh ho ' 'he

streams that prrsons walki'il from shore to -~li'>i<- ..if)
Thesi' storifs of ahnndanee may he sliu'htly «'\.

oner as eompart'tl to prc.m-nt eonditioAs may not Im* drnied i-ven by the
ski'ptieal. Thf many rstahlislicd fai-tors wliirh have • ' 'il

are not so ^'rin'rally known. Intelliirfut eonsrrvatifjn li-

/ance of these factors and makrs ffVort tc» anticipate r; li-

tions so that appropriate measures may he atTeeted and thu.H minimize
or postpone the ill etTects that otherwise would aecnie or ponible
destruc'tion of the species.

I'rohahly the first advei-se factor afVectinif fish life in thi.H State
was the atlvcut of the t'old miner in 1H4I>, who very shortly be^an
pollutinj? the streams, makini: them undesirahle to fish life and con-
structiiiir dams for the diversion of water with no means provided to
enahle miLrratim; fish to these «lams to the spawning' LTi-uri'ls ahove.
This continued without restriction for many years.

In ISTO, twenty years aft«'r the State was admitti'd to tf: n.

n Hoard nf I'^jsh ( 'oinniissjimers was ereated i-hariTed hy law wr.n the

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