Ticks of the Genus Haemaphysalis in the Western United States

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Theses and Dissertations

1967-05-02

Ticks of the genus Haemaphysalis in the western United States

Joe M. Merino Brigham Young University - Provo

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TICKS OF THE GENUS HAEMAPBYSALISIN THE

WESTERNUNITED STATES

A Thesis

Presented to the

Department of Zoology and Entomology

Brigham Young University

In Partial Fulfillment

of the Requirements for the Degree

Master of Science

Joe M. Merino

May 1967 This thesis by Joe M. Merino is accepted in its present form by the Department of Zoology and Entomology of Brigham

Young University as satisfying the thesis requirement for the degree Master of Science.

Typed by Berna B. Allred ACKNOWLEDGMENTS

I express my deepest appreciation to Dr. D Elden Beck, chairman of my advisory committee, for his kind assistance, his friendship and guidance throughout this study, and to other members of my committee, Dr. Dorald M, Allred and Dr.

Joseph R. Murdock.

I am also endebted to the Rocky Mountain Laboratory, and especially Dr. Glen M. Kohls for loan of material and data, and for kind assistance and suggestions. Grateful acknowledgment is made to Dr. Betty S. Davis, University of

California Museum of Vertebrate Zoology, D. Eo Johnson 9 Dug- way Proving Ground, Dr. Carl 0. Mohr, University of California

Agricultural Experiment Station, and Dr. C. M. Senger, Western

Washington State College, for loan of material. The Depart- ment of Zoology and Entomology, Brigham Young University, supplied laboratory space, equipment, and supplies.

To my wife, Julie, I am thankful, for her moral support and strength throughout my graduate work. TABLE OF CONTENTS

Page

ACKNOWLEDGMENTS iii

LIST OF TABLES . V

LIST OF ILLUSTRATIONS vi

INTRODUCTION..... 1

REVIEWOF THE LITERATURE 4

FIELD METHODS 13

LABORATORYMETHODS 15

TAXONOMICLISTING 20

Haemaphysalis chordeilis 20 Haemaphysalis leporispalustris 21

RESULTS 42

Haemaphysalis leporispalustris 42

Geographic Distribution 42 Seasonal Occurrence 43 Host-Parasite Association 45 Anatomy and Morphology, 47

Haemaphysalis chordeilis 48

Geographic Distribution 48 Seasonal Occurrence 48 Host-Parasite Association 49 Anatomy and Morphology, 49 DISCUSSION, so CONCLUSIONS 100

LITERATURECITED . 104 LIST OF TABLES

Table Page

L Collection data for Haemaphysalis leporispalustris (Packard) 1869 in the Western

United States , , o ••••• 23

2. Collection data for Haemaphysalis chordeilis (Packard), 1869 in thE Western United S ta t es , , , . , . . , , , , , 41

3. The number of times that Haemaphysalis leporispalustris was collected from a given host species in California . , , . , . 76

4. The number of times that Haemaphysalis leporispalustris was collected from a given host species in Utah ... 77

5. Haemaphysalis leporispalustris adults taken from hosts other than leporids in the Western United States 78

6, Numbers of Haemaphysalis leporispalustris in all developmental stages taken from Lepus spp, and Sylvilagus spp. in Utah 79

7 . Immature Haemaphysalis leporispalustris taken from hosts other than leporids in the Western United States 81

8 0 Hosts of Haemaphysalis chordeilis in the Western United States .. 86 LIST OF FIGURES

Figure Page

1. Geographic distribution of g. leporispalustris taken from L. americanus and L. californicus. The lined area represents the distribution of L. americanus and the rough area that of L. californicus (After Hall and Kelson,-1959) 53

2. Geographic distribution of g. leporispalustris taken from Lepus spp., and ~- townsendii, and non-leporid hosts. Dotted area represents the distribution of L. townsendii (After Hall and Kelson, 1959) . . . 54

3. Geographic distribution of H. leporispalustris taken from Sylvilagus spp. and S. auduboni. Lined area represents the ~istribution of~- auduboni (Afcer Hall and Kelson, 1959) . . . .. , 55

4. Geographic distribution of~- leporispalustris taken from~- bachmani, ~- floridanus, and~- idahoensis. The fine- stippled area represents the distribution of~- bachmani, the rough area that of S. floridanus, and the stippled area thai of S. idahoensis (After Hall and Kelson, 1959) ...... 56

5 0 Geographic distribution of H. leporispalustris taken from S. nuttallii (After Hall and Kelso~, 1959) ... 57

6. Geographic di.stribution of H. leporispa 1 us tris .i.n Utah taken from Lepus spp. , Sylvilagus spp., both Sylvilagus and and Lepus, and from non-leporid hosts 59 vii

Figure Page

7 0 Total numbers of H. leporispalustris occurring by month in Arizona (1924, 1938, and 1948) ...... 63

8 0 Total numbers of H. leporispalustris occurring by month in Colorado between 1925 and 1943 ...... 63

9 0 Total numbers of H. leporispalustris occurring by month in Idaho between 1939 and 1961 . , ...... , . 63

10. Total numbers of H. leporispalustris occurring by month in California between 1920 and 1964 .. , . , . 65

11. Total numbers of H. leporispalustris occurring by month in San Mateo County, California in 1961 ...... , 65

12 0 Total numbers of g. leporispalustris occurring by month in Montana between 1937 and 1959 ...... 68

13. Total numbers of H. leporispalustris occurring by month in New Mexico (1924, 1940, and 1963) ...... 68

14. Total numbers of H. leporispalustris occurring by month in Nevada (1941 and 1961) ...... , .. . 68

15. Total numbers of H. leporispalustris occurring by month in Oregon between 1922 and 1961 ...... 69

16. Total numbers of H. leporispalustris occurring by month in Washington between 1922 and 1959 . , .... 69 viii

Figure Page

17. Total numbers of go leporispalustris occurring by month in Wyoming between 1922 and 1943 ... , . o o o , 69

18. Total numbers of H. leporispalustris occurring by month in Utah between 1949 and 1953 . , o • o •• , , 72

19. Total numbers of H. leporispalustris occur-

ring by month in Utah (1961) ... o •• 72

20. Total numbers of H. leporispalustris occurring by month in Utah from 1960 to 1964, excluding 1961 ... " . 73

2L Geographic distribution of H. chordeilis in the Western United States .. , . , 83

22. Seasonal occurrence of H. chordeilis by month in the Western United States , 85

Dorsal and ventral views 9 larval H. chordeilis (upper) and larval H. leporispalustris (lower) ... -. 93

24. Dorsal and ventral views, nymphal H.

chordeilis .. , . , ... , o • 94

25. Dorsal and ventral views 9 female H. chordeilis . . . . , , . . . , . . , , . . . 9 S

26. Dorsal and ventral views 9 male H" chordeilis ...... 96

27. Dorsal and ventral views, nymphal H. 1 e po r is pa 1us tr is . . . , . . . . , . . . . . 9 7

28. Dorsal and ventral views 9 female H. leporispalustris . o , • o ••••••• , • 98

29 0 Dorsal and ventral views 1 male H. leporispalustris ... , ... 99 INTRODUCTION

In 1893 Smith and Kilbourne made the first reference to ticks being vectors of disease. This has been frequently proven over the years (Ricketts, 1906; Clarks 1918; Todd,

1919; Parker, 1923; Badger, 1932; Bishopp, 1933; Davis 9 1941, to mention a few). Since this initial discovery was made, interest and research regarding ticks has constantly increased,

In the last decade the Department of Zoology and Ento- mology at Brigham Young University has conducted studies on geographic and ecological distribution as well as host-parasite relationship for ticks of the western United States (Beck,

1955a, 1955b; Allred, Beck and White 9 1960; Beck, Allred and

Brinton, 1963; Brinton and Beck, 1963a, 1963b; Brinton and

Kohls, 1963; Mohr, Beck and Brinton, 1964; Brinton, Beck and

Allred, 1965; and Herrin and Beck, 1965). These studies are being conducted in order to contribute further to the understanding of arthropods as vectors of disease agents.

Thousands of specimens collected over the years are in the immature stages of development as larvae and nymphs.

Most of the early keys which are still available for classi- 2 fication of tick genera are inadequate to make accurate identification of tick larvae. The most recent keys for identification of ticks in various stages of development were com-

posed by Brinton and Beck (1963a 9 1963b) and Brinton 9 Beck and Allred (1965). These apply only to the ticks of the western United States.

This paper is a continuing part of these tick studies which centers attention on species of a single genus, One such study was completed for species of the genus Otobius

(Herrin and Beck, 1965). The present study of the genus

Haemaphysalis was to achieve the following objectives:

(1) describe geographic distribution and seasonal occurrence 9

(2) show host-parasite associations for all stages of development, and (3) devise keys for specific identification of all stages of development. As a part of the data for taxonomic identification, detailed anatomical and morphological features were illustrated for all species at all stages of development.

There are only two species of Haemaphysalis in the western United States--Haemaphysalis chordeilis (Packard) and g. leporispalustris (Packard). Published information on

H, leporispalustris is extensive, The Brigham Young University 3 collections of this species for the western United States

are abundant" On the other hand 9 Ho chordeilis collections are limited due to its restricted distribution in the western states. 4

REVIEW OF THE LITERATURE

In 1827 Audouin described Haemaphysalis leachii and placed it in the genus Ixodes. It was not until 1844 that

Koch established the genus Haemaphysalis with H. concinna as the type species. Neumann (1897) published a key to the known species of this genus, His paper was considered the only original work since the 1844 publication of Koch (Nut- tal, Warburton and Robinsonj 1915). Salmon and Stiles (1901) followed with a publication in which they included a "short

generic diagnosis, synonyny 9 bibliography and translation of

Neumann's key" (Nuttall et al. 1 1915). Other authors followed

with separate works giving generic descriptions~ keys 9 diag-

nostic characters, habits of oviposition 9 and other notes on

biology (Guy 9 1906; Hunter and Hooker 9 1907; Banks 9 1908;

Nuttall, 1908; Hunter 9 1909; Neumann, 1911; Hooker 9 Bishopp and Wood, 1912). In 1915 Nuttal et al. published a monograph of the genus thus bringing up to date all existing information. The taxonomy of the New World haemaphysalids was up-dated by Cooley (1946) who added the new species H. juxtakochi from Panama. 5

During the last ten years renewed interest in ticks has been evident on a world-wide basis. This has been

prompted by the need to know more about all stages of tick development concerning host-parasite association, disease

transmission potential, and related aspects of tick biology.

Haemaphysalids have received specific att.e.ntiorL Le:adir:g in

the recent investigations of ixodid ticks are such workers as Serdyukova (1955), Gregson (1956), Cerny <1957), Clifford

(1958), Hoogstraal and Kohls (1959), Kohls (1960). Hoogstraal 1

Trapido and Rabelo (1963), Hoogstraal and Trapido (1963) 9

Hoogstraal (1964), Trapido, Hoogstraal and Vanna (1964L and

Hoogstraal, Trapido and Kohls (1965). In practically all

instances careful attention wa.-3 giver\ Lo Lllustrations and accompanying descriptions for all staasss of dE:velopmerx of haemaphysalid ticks. How~ver. comparatively little attention has been given to the well knm..m. H .. leporispalustris since

the work of Nuttall et al. (1915), and still much less atten-

tion to H. chordeilis.

In 1869 9 A. S. Packard described the species "leporis-

palustris911 but included it in the genus lxodes. Neuman

(1897) removed leporispalustris from lxodes and assigned it

to Haemaphysalis. MorE:over 1 he dropped the hyphenated name 6 and listed the species as g. leporis. In 1907, Hunter and

Hooker used the name H. leporis-palustris. The present cor- rect use of the specific name is not hyphenated and is g. leporispalustris (International Code of Zoological Nomencla- ture, 1961),

Early workers considered H. leporispalustris to be more widely distributed than their collection data revealed

(Hunter and Hooker, 1907). Hooker (1909) mentioned that this

species had been collected in Arizona, California, Colorado 9 and Nevada. Workers since 1945 have reported a wide distri-

bution throughout the United States (Bishopp and Trembley 9

1945; Bequaert, 1946; Cooley, 1946; and Kohls, 1960). Gregson

(1956) observed that this species occurs from British Columbia to Newfoundland in Canada, Its distribution extends from

Alaska to Argentina according to Cooley (1946).

Haemaphysalis lepori.spalustris is present throughout. the year in all developmental stages in nature according to

Hooker, Bishopp and Wood (1912). Nuttall et al. (1915) mentioned that "under ideal conditions two life cycles per year could be observed." Recent workt 0 rs have noted that climate affects tick activity (Bequaert, 1946; Yeatter and Thompson,

1952; Beck, 1955b; Bacon, Drake and Miller, 1959, and Camin, 1963). 7

Over eighty hosts for H. leporispalustris in the

United States were mentioned by Bishopp and Trembley (1945).

Host-parasite relationships were further updated by Cooley

(1946) and Kohls (1960) for the New World. These studies indicated that leporids were the p~eferred hosts of this tick. Early workers also reported that leporids are the pre= ferred hosts of this tick in the adult stage. They also stated that the immature stages are widely found on birds.

However, Beck (1955), Beck, Allred and Brinton (1963) and

Mohr, Beck and Brinton (1964) mentioned that the immatures are not commonly found on birds in their studies ir.cluding parts of the western Uniced States.

The life cycle of H. leporispalustris was studied by

Hooker (1908) 5 Hooker, Bishopp and Wood (1912) 9 Nuttall et al. (1915), Miura (manuscript in preparation). Gregson

(1956) and Arthur (1962) also have summarizf:d life cycle data for this species.

Haemaphysa 1 is lepor i~a Lus tris has long DeE:n implica- ted as a capable vector of Rocky Mountain spott~d fever.

Parker (1923) first noted the probabl2 significance of this tick in maintaining pathogenic spotted fever rickettsiae in nature. The geographical distribution of spotted fever and 8 of Sylvilagus nuttallii was compiled by Jellison and Parker

(1945). Serological studies conducted by Parker et al. (1951) revealed rickettsiae recovered from H. leporispalustris found on cottontails. Coffey (1954), Beck (1955b) and Gregson

(1956) stated that this tick is important in maintaining the disease in nature.

Parker (1924) observed the transmission of tularemia experimentally by H. leporispalustris in the laboratory. He stated that this tick was a probable vector of the disease among rabbi ts and "ground-frequenting birds"" Franc is ( 1937) also mentioned the role of this tick in transmitting tularemia

in nature. In Alaska 9 Philip (1933 9 1954) rE:'.ported the re- covery of Bacterium tularense from naturally infected H" leporispa lus tris taken from hares. That this tick i.s a natural carrier of tularemia in nature has been reported by

Brown (1944a) 9 Jellison and Parker (1945L Cooley (1946),

Yeatter and Thompson (1952) 9 and Gregson (1956).

As with!:!- leporispalustri.s 1 thf: "bird tick" H- chor~ deilis has also undergone chang(;;S in r.omE:nclat.ure. PacK.ard originally described the species chordeilis ln 1869 and placed it in the genus lxodE:s" Neumann ( 1905) later published a work in which the tick was rightly placed with 9

Haernaphysalis but as punctata var, cinnabarinao Later Nuttall et al. (1915) stated that H. cinnabarina Koch and H. chordeilis

Packard were synonymous, Bi~hopp and Trembley (1945) reported that the H, cinnabarina female described by Nuttall tt al,

(1915) was identical to the type specimen of chordeilis,. and implied that all cinnabarina in North America wE.rE: chord,,U1s,

Cooley (1946) listed this tick as H. chord0ilis and did not include H, cinnabarina as a valid species in the knericas.

In 1960 Kohls recognized only three specibs of haemaphysalids

in the New World--H, chordeilis 1 Ho juxt_akochi. 9 and H, lepo,E_- ispa lustris.

Extensive data for go chordeilis w~re lacking when

Hunter and Hooker (1907) noted that it was found Hl the eastern

United States. Later 1 Hooker 1 Bishopp ari.d Wood (1912) r,2-

ported that i.ts distribution was probably' wide 9 but due to lack of extensive collection data} not extensively known,

Hewitt (1915) mentioned that this tick occurs in Canada and

Nuttall et al. (1915) stated that Ho chordeilis appeared to be widespread in North America. Two decades ago Bishopp and

Trembley (1945) reportEd that the "bi.rd t;ick" was distri.buted in the eastern and southern lJnited States~ and in eastern

Canadao Bequaert (1946) also observed this tick in Oregon 10 and North Dakota. Cooley (1946) reported H. chordeilis in

Montana 1 and Gregson (1956) noted that it occurs commonly on upland game birds in Canada except in the Pacific Coast wet belt. One collection from California was reported by Car~ penter (1956). This tick has also been reported ~n Mexi.co by Hoffmann (1962).

A paucity of published information exists for seasonal activity of H. chordeilis in the western United States. ]n

the south 9 however 9 Bishopp and Trembley (1945) reported that all tick specimens were collected in the late fall~ winter

and spring, In the north, the ticks occurrE:d in spring 7 summer and fall, Hooker, Bishopp and Wood (1912) observed these ticks in April and November. Carpenter (1961) made hi.s lone collection in California during the month of January.

The type specimen of H. chordeilis was collected from a night hawk by Packard (1869). Othtr authors have reported

it from a variety of ground-inhabiting birds (Banks 9 1908;

Hadley, 1909; Hunteri 1909; Bishopp, 1911; Hooker, Bishopp and Wood, 1912; Nuttall et al., 1915). Later workers

(Bishopp and Trembley, 1945; Bequaert 9 1946; Cooley 9 lg46;

Gregson, 1956; Kohls 9 1960; Clifford~ Anastos and Elbl 9 1961;

and Hoffmann 1 1962) also indicated that H" chordeilis is a 11 common parasite of birds, It has also been collE:cted from

man, cattle 9 rabbits 9 horses 5 and sheep.

Bequaert (1946) reported that Parker, Philip and Davis

(1932) found H. chordeilis naturally lnfectE:d wlth tularen,ia"

Gregson (1956) stated that in Canada this tick is important

"as a parasite of turkeys and as a vector of d1.sEast:'s of wildlife."

Little is known concerLing thE lifE eye l.F. of lhe "bird

tick," Hooker 9 Bishopp and Wood (1912) obtainE,d some. information from ticks collected in the field and reared in tne laboratory. They found that engorged larvae gan to drop from the host after feeding for five days, Afttr tw~lve days feeding the last 1.arva had droppE:d from thr, host, Fourteen days after dropping engorged th~ first Larvae ~merged as

nymphs, Nymphs 9 thE:y obstcrV(cd 9 fully Engorgt=cd in fl\;'£, LO eight days after attachmenL After dropping from the host as an engorged nymph the first adult emerged in twenty-six days. One female fed for nineteen days and dr0pped fully engorged. Haemaphysalis choniE.:::ilis is n?portt::d Lo attach to the host mainly around the eyes and ears, tht n~ck, and the top of the heado

As with H. leporispalustris, the anatomy of H, chordellis 12 has not recently been treated, Nuttall et al, (1915) examined this problem briefly. Cooley (1946) presented drawings of all developmental stages of this species. Included in

Gregson's (1956) publication were line drawings of structures of taxonomic value. Lastly, Clifford et al, (1961) treated only the larval stage. 13

FIELD METHODS

A large part of the tick collections used in this study were supplied from the field surveys of the Zoology and Entomology Department, Brigham Young University. Lago- morphs were collected primarily with a shotgun using various gauges. Occasional specimens encountered dead on highways were collected and examined for parasites.

All representative stages of g. chordeilis were selected from preserved specimens supplied by the Rocky Mountain Lab- oratory, Hamilton, Montana.

Hosts were placed in large, heavy-weight paper bags which were carefully folded and stapled to prevent the escape of ticks or other ectoparasites. These bags were labeled to show locality, date, and host. All specimens were returned to the laboratory or field station as soon as possible after collecting. If return to the university laboratory in a few hours was planned, the specimens were stored under cold but not freezing conditions. Removal was done at convenience of schedule. When return to the laboratory was not anticipated

for several days 1 the host specimens were allowed to cool 14 overnight and the ectoparasites were removed the following

morning. Smaller animals, mainly rodents 1 were obtained by use of live traps of several kinds. Break-back snap traps were also employed. Trapped specimens were placed in paper bags following the same procedure as with the lagomorphs. 15

LABORATORYMETHODS

Hosts were examined in deep 1 whitE., enamel-ware pans o

A 75-watt lamp was placed over the pan containing the animaL

As the heat from the lamp warmed the body? ectoparasites usually crawled or jumped from the host and moved about on the pan where they were easily seen and collected" The host's body was carefully brushed? and the areas around the ears, face, and axillae examined for specimens which may have remained attachedo All ticks were placed in Oudemans preserva~ tive which was prepared as follows:

70% ethyl alcohol 0 0 0 0 G 0 87 parts

Glycerol . . 0 0 0 0 0 0 0 5 parts

Glacial acetic acid 0 0 0 8 parts

Ticks die in this preservative with their legs fully extended,

This is especially important for immature stages which in some instances require mounting on microslides and subsequent examination under high magnification to determine species"

Representative larval and nymphal g. leporispalustris were selected for slide mounting after first observing them under a Leitz dissecting microscope" Selected specimens were 16 then cleared and mounted by the following procedure.

Specimens were removed from the 70% ethyl alcohol preservative and placed in Nesbitt's solution where they were allowed to remain at room temperature, Nesbitt's solution is prepared as follows:

Chloral hydrate 40 go

Water o . 25 go

Hydrochloric acid 2o5 ml,

This solution is used to clear the specimens before mounting,

Checked hourly, an additional period of not more than three hours was allowed beyond the time when the specimens first appeared to be cleared. Overclearing is undesirable as it inhibits leg extension. Some engorged specimens readily cleared, and body puncturing was unnecessaryo

Cleared tick specimens were mounted directly from the

Nesbitt's solution to a drop of Hoyer's medium previously

placed on the microslide, Before the coverslip was applied 1 specimens were oriented in the medium to the desired position.

This facilitated later observation under high-power magnifi- cationo The mounted specimen was warmed over an alcohol lamp at brief intervals until the legs were uniformly extended by the warmed mediumo The prepared microslide was 17

then placed in a warming oven for forty-eight hours at a

temperature of approximately fifty degrees centigrade" This

resulted in further clearing and solidification of the mount-

ing medium. If at the end of twelve to twenty-four hours the

legs had moved from the extended position~ rewarming over a

flame usually caused them to extend again, Mounted specimens were ringed with Zut slide ringing compound.

Representative larval and nymphal H. chordeilis were

selected for mounting after observing them under a Leitz dissecting microscope. Selected specimens were then cleared by the following procedure.

The immature specimens removed from the 70% ethyl alcohol preservative were placed in 85% lactic acid. Speci- mens remained in lactic acid for 72 hours. This period was spent in a warming oven at approximately fifty degrees centigrade at which time the specimens were sufficiently cleared to mount on microslides. They were removed from the lactic acid and placed in a solution of 50% lactic acid and 50%

HoyeY's medium at room temperature. After remaining in this

solution for about thirty minutes the immatures were mounted on rnicroslides following the same procedure as with the H.

leporispalustris. 18

The live specimens of !!,o leporispalustris which had

been placed in 70% ethyl alcohol cleared much faster than

did specimens which were already dead at the time of preser-

vationo Freshly preserved specimens also cleared much faster

than did those which had been preserved for any length of

time o

The immature stages were drawn from a series of speci~

mens selected after examination with a Zeiss phase-contrast

microscope. The specimens were projected from a Leitz micro-

slide projector and the body outline was reproduced on quad-

rille paper ruled ten squares to the inch. A penciled out-

line of the specimen showing general anatomical structures

both dorsal and ventral was sketched on the quadrille paper.

Detailed anatomical and morphological structures were added

after careful examination of each specimen with the phase-

contrast microscope. Final drawings were done by comparing

the semi-completed sketch with unmounted and mounted specimens observed with a Leitz stereomicroscope at magnifica-

tions of 120 to 216 diameters. Two American Optical Uni-

versal Illuminators were used for illumination. These were

placed about four inches from the specimens.

The adult ticks were not mounted for study. They were 19 instead supported on molding clay to facilitate manipulation and observation of all body surfaces. The adults are easily manipulated with a pair of fine-pointed forcepso A pair of

American Optical Universal Illuminators provided good light- ing, A Leitz stereomicroscope with a magnification range of

120 to 216 diameters was used for adult drawings, A standard net reticule of 10 mm square ruled into 1,0 mm squares was employed, Inserted in the eye piece it facilitated comparative size illustrations made on similar paper as that used for the immature stages, 20

TAXONOMICLISTING

Synonomy generally follows that of Cooley (1946),

Haemaphysalis chordeilis (Packard) 9 1869

1869. Ixodes chordeilis Packard 9 p. 67,

1905. Haemaphysalis punctata var. cinnabarina Koch~ Neumann 9

p, 237,

1908, Haemaphysalis chordeilis Packard~ Banks 9 p, 34,

1911, Haemaphysalis punctata cinnabarina Koch: Neumann,

p, 108.

1912. Haemaphysalis chordeili.s (Packard): Hooker, et atv,

p O 9 7 0

1915. Haemaphysalis cinnabarina Koch: Nuttall and Warburton,,

p. 372,

1945. Haemaphysalis chordeilis (Packard): Bishopp and

Trembley 9 p. 25,

1946, Haemaphysalis chordeilis (Packard)~ Bequaert 9 p. 19L

1946. Haemaphysalis chordeilis (Packard): Cooley 9 p. 37,

1956. Haemaphysalis chordeilis (Packard)~ Gregson, p. 35. 21

Haemaphysalis leporispalustris (Packard), 1869

1869. Ixodes leporis-palustris Packard, p. 67.

1896. Rhipistoma leporis Osborn 9 P 0 26L 1886. Gonixodes rostralis Duges, p, 1290

18970 Haemaphysalis leporis (Packard): Neumann 9 p, 3430

1901. Haemaphysalis leporis (Packard): Neumann, p. 262,

1905. Haemaphysa lis leporis (Packard): Lahille, P 0 45, 1907. Haemaphysalis leporis-palustris (Packard): Hunter

and Hooker, p. 530

1908. Haemaphysalis leporis-palustris (Packard): Banks 1

p. 33.

1908. Haemaphysalis leporis-palustris (Packard): Hooker 1

p. 47.

1909. Haemaphysalis leporis-palustris (Packard)~ Hooker 9

p. 423.

1909. Haemaphysalis proxima Aragaoi p. 100-101.

1911. Haemaphysalis leporis-palustris (Packard)~ Hunter

and Bishopp, p. 228.

1911. Haemaphysalis leporis var. proxima Aragaoi po 167.

1912. Haemaphysalis leporis-palustris (Packard): Hooker

et al., p. 890 22

1913. Haemaphysalis leporis-palustris (Packard)~ Hadwen 9

p. 9 7.

1915. Haemaphysalis leporis-palustris (Packard)~ Nuttall

and Warburton 9 p. 387a

1938. Haemaphysalis leporis-palustris (Packard):: Hear le;

p O 100.

1941. Haemaphysalis proxima (Aragao): Osorno-Mesa~ p. 428.

1943. Haemaphysalis leporis-palustris (Packard)~ Fairchild 9

p. 586.

1944. Haemaphysalis leporis-palustris (Packard):

1945. Haemaphysalis leporis-palustris (Packard): Bishopp

and Trembley, p. 27.

1946. Haemaphysalis leporis~palustris (Packard)~ Bequaert,

p. 194.

1946. Haemaphysalis leporis-palustris (Packard)~ Cooley,

p. 3L

1950. Haemaphysalis leporis-palustris (Packard): Brown

and Kohls, pa 198.

1956. Haemaphysalis leporis-palustris (Packard)~ Gregson, p. 3Li

1960. Haemaphysalis leporis-palustris (Packard): Kohls, p. 355.

1961. Haemaphysalis leporispalustris (Packard): International

Code of Zoological Nomenclature, p. 27 and p. 35. Table 1. Collection data for Haemaehysalis leeorisealustris (Packard) 1869 in the Western United States,

No. Collected State Locality Date Host Animal Collection No. L N A

Arizona Kingman, Mohave County 25 June 1924 Sylvilagus sp. 1 R.M.L. 16896 Winslow, Navajo County 8 July 1924 auduboni 2 16910 Maricopa County 18 Apr 1938 sp. 2 16393 4May 1938 Citellus sp. 1 16388 Mohave County 11 May 1938 Sylvilagus sp. 2 16390 Haulpai Mts., Mohave County 3 Oct 1948 Passerella iliaca 7 10 28678 3 Oct 1948 Pieilo erythroehthalmus 50 1 28678 California Corralitos, Santa Cruz County 8 Aug 1920 Sylvilagus bachmani X X 15849 15 Aug 1920 Leeus californicus X 15842 20 Aug 1920 Sylvilagus auduboni X X 15848 20 Aug 1920 bachmani X X 2 15853 Balls Ferry, Shasta County 22 Aug 1922 auduboni X 16802 Lc>-pusealifomicus 1 2 16804 Soledad, Monterey C,Junty 30 Aug 1922 Sylvilagus bachmani X 16807 30 Aug 1922 Lepus californicus 2 3 16809 3 Sept 1922 2 16813 San Antonio Mission, Monterey 3 Sept 1922 X 15965 County 4 Sept 1922 Sylvilagus auduboni 1 2 15966 Morro, San Luis Obispo County 9 Sept 1922 bachmani 1 16821 San Andreas, Calaveras County 19 Sept 1922 X X 16824 Lemoore, Kings County 26 Sept 1922 Leeus californicus 1 15971 La Jolla, San Diego County 9 Oct 1922 Sylvilagus auduboni X X 16826 10 Oct 1922 X 16841 11 Oct 1922 1 16844 Barstow , San Bernardino 20 June 1924 sp. 3 15985 County Hastings Reservation, Monterrey 16 May 1934 auduboni 9 H.Res. 6381 County 22 June 1934 2 3 6325 29 July 1934 Lepus californicus 3 37 6344 N 30 July 1934 Sylvilagus auduboni 2 6382 w 14Aug 1934 3 6376 Table 1. {continued)

No. Collected State Locality Date Ha;t Animal Collection No. L N A

California Hastings Reservation, Monterey 14Aug 1934 Sylvilagus auduboni 1 H.Res. 6350 County 31 Mar 1935 2 6351 E Rosarito, Baja California 17 June 1938 bacbmani 4 12 R. M. L. 18806 Hastings Reservation, Monterey 18 June 1938 auduboni 2 21 H.Res. 6366 County 21 June 1938 bachmani 4 35 6322 IBesquintal, Baja California 23 June 1938 sp. 10 1 R. M. L. 18805 Hastings Reservation, Monterey 24 June 1938 auduboni 6 H.Res. 6361 County 4July 1938 3 18 6371 5 July 1938 II 2 5 12 6369 5 July 1938 3 6377 9 July 1938 2 9 6367 9 July 1938 1 8 6370 10 July 1938 12 6 6380 BJuly 1938 23 6383 18 July 1938 20 18 6384 14Aug 1938 11 4 6374 9 Apr 1939 bachrnani 25 6319 Monterey County 25 Apr 1939 1 2 R. M. L, 16059 Hastings Reservation, Monterey 25 Apr 1939 3 H.Res. 6320 County 3 May 1939 auduboni 1 6345 Monterey County 16May 1939 5 6 R. M. L. 16046 Hastings Reservation, Monterey 12 June 1939 4 H.Res. 6349 County 13 June 1939 6 6348 14June 1939 7 6343 Monterey County 18 June 1939 X 5 R. M. L. 16047 22 June 1939 5 2 16048 N Hastings Reservation, Monterey 22 June 1939 3 6 H.Res. 6363 ~ County Table 1. (continued)

No. Collected State Locality Date Hou Animal Collection No. L N A

California Hastings Reservation, Monterey 22 June 1939 Syl vilagus auduboni 4 H.Res . 6138 County Monterey County 23 June 1939 5 R. M.L. 16049 3 July 1939 1 16052 Hastings Reservation, Monterey 3 July 1939 bachmani 1 8 H.Res . 6323 County 3 July 1939 auduboni 1 6368 3 July 1939 4 6373 Monterey County 13 July 1939 1 13 2 R. M. L. 16055 Hastings Reservation, Monterey 13 July 1939 5 H. Res. 6153 County 13 July 1939 2 8 6327 Monterey County 27 July 1939 1 R.M . L.16056 Hastings Reservation, Monterey 27 July 1939 1 H.Res . 6372 County Monterey County 29 July 1939 Lepus californicus X 1 R. M. L.16045 Hastings Reservation, Monterey 29 July 1939 11 H.Res. 6197 County 29 July 1939 Sylvilagus auduboni 4 6379 14Aug 1939 3 R. M. L, 16280 2 Mar 1940 3 H.Res. 6342 9Apr 1940 bachmani 7 14 6317 13 Apr 1940 5 6154 15 Apr 1940 18 6318 7 July 1940 -:luduboni 6 6347 15 July 1940 1 6155 2 Aug 1940 3 6156 2 Aug 1940 4 6326 San Francisco County 18 Sept 1940 bachmani 100 R. M. L, 18436 Calaveras Dam, Alameda 23 May 1941 10 30 30 18880 County Hastings Reservation, Monterey 1941 auduboni 2 N 8 July H.Res. 6233 lJ1 County Table 1. (continued)

No . Collected Collection No. State Locality Date Host Animal L N A

California Hastings Reservation, Monterey 8 July 1941 Sylvilagus auduboni 1 H.Res. 6324 County Amedee, Lassen County 31 May 1942 Lepus califomicus 1 R.M .L. 19964 Hastings Reservation, Monterey 10 June 1942 Sylvilagus auduboni 13 H.Res . 6362 County 20 June 1942 1 6378 1 July 1942 4 6375 " W Portal Area, Fon Bakex, 13 Oct 1942 Thryomanes bewicki 29 R. M. L. 20143 Marin County Hastings Reservation, Monterey 16 Jan 1943 Sylvilagus bachmani 2 H.Res. 6157 County Ft. Barry, Marin CoUDty June 1944 Sp. 5 25 R.M.L. 21319 26 Aug 1944 X X X 21334 2 Sept 1944 12 8 3 21378 20 Jan 1945 Lepus califomicus 2 H 21508 Hastings Reservation, Monterey 7 Mar 1945 Sylvilagus auduboni 6 H.Res. 6357 County 2 Apr 1945 2 28 6354 2 Apr 1945 1 25 6355 10 Apr 1945 8 6353 4June 1945 4 6352 4June 1945 8 6356 4June 1945 9 6358 4June 1945 6 6359 13 June 1945 3 6360 Los Angeles, Los Angeles County 25 July 1945 bac hmani 5 R.M.L . 21583 Hastings Reservation, Monterey 13 Apr 1945 6 H.Res. 6321 County 26 June 1946 auduboni 4 8 6365 14Aug 1948 4 4 6364 Jessups Ranch. Modoc County 9May 1949 sp. 26 1 1 R.M . L. 26598 N Hastings Reservation, Monterey 14 July 1950 auduboni 2 H.R es. 6346 (j'I County Table 1. (continued)

No. Collected Collection No. State Locality Date Host Animal L N A

H. Res. 6341 California Hastings Reservation, Monterey 15 July 1950 Sylvilagus auduboni 2 County 26 Apr 1951 1 6336 23 May 1951 7 15 6338 23 May 1951 8 6339 23 May 1951 3 9 6333 24May 1951 31 17 6335 Sutro Forest, San Francisco 26 June 1951 bachmani 1 4 R.M . L. 31705 County Hastings Reservation, Monterey 21 July 1951 auduboni 1 H.Res. 6334 County 23 July 1951 6 13 6332 Dana Point, Orange County 8 Dec 1952 sp. 1 4 R.M.L. 32034 11 Dec 1952 1 2 32035 Hastings Reservation, Monterey 28 June 1953 auduboni 5 3 1 H.Res. 6340 County 22 July 1953 Lepus californicus 1 6330 22 July 1953 7 6331 1 Aug 1953 1 6329 3 Aug 1953 3 6316 4Aug 1953 Sylvilagus auduboni 3 4 6337 18 Aug 1953 21 6315 23 June 1954 Lepus califomicus 16 26 6328 San Juan Capistrano, Orange 30 Sept 1957 Sylvilagus sp. 2 18 R.M . L. 32033 County Napa, Napa County 26Aug 1959 Peromyscus maniculatus 1 B.Y . U. 4991 Spring Valley, San Diego 28Dec 1960 Lepus califomicus 2 27 3 5122 County 28Dec 1960 1 I• 4983 San Mateo County 25 May 1961 Sylvilagus bachmani 2 2 17 5497 6 June 1961 Pipilo fuscus 1 5405 5339 N 8 June 1961 Sylvilagus bachmani 1 1 -....J 8 June 1961 9 23 5473 Table 1. (continued)

No. Collected Date Host Animal Collection No. State Locality L N A

California San Mateo County 23 June 1961 Lepus sp. 2 52 48 B. Y. U. 5348 27 June 1961 Sylvilagus bachmani 42 28 5569 6 July 1961 16 14 5492 7 July 1961 l l 5411 7 July 1961 Unknown 59 7 5313 7 July 1961 Sylvilagus bachmani 3 5457 12 July 1961 27 9 5501 12 July 1961 3 5459 14 July 1961 l 5452 14 July 1961 l 13 1 5562 19 July 1961 Unknown 2 l 5134 19 July 1961 Sylvilagus bachmani 3 6065 20 July 1961 20 9 5509 4Aug 1961 ~ uy..th.topthalm.us 1 5400 .. 4Aug 19£1 2 5355 4 Aug 1961 2 5402 17 Aug 1961 Unknown 10 5352 17 Aug 1961 Thryomanes bewicki 2 5406 22 Aug 1961 Sylvilagus bachm.ani 11 6 2 5500 22 Aug 1961 Pipilo erythropthalmus 65 4 5451 Dana Point, Orange County --- Citellus Beecheyi 1 Ryckman et. al. 1955 San Juan Capistrano, Orange --- 1 1955 County Camp Pendleton, San Diego --- 1 1955 County Loma Linda, San Bernardino -- 1 1955 County Reche Canyon, San Bernardino --- Lynx rufus 1 1955 County Colorado Blanca, Costilla County 17 Aug 1925 Sylvilagus audoboni 1 R.M.L. 16888 6 mi So, Boulder County 22 June 1939 sp. 1 15684 N 9 mi No, Boulder County 24Apr 1940 X X 00 27 May 1940 X X 16629A Table 1. (continued)

No. Collected Collection No. State Locality Date Host Animal L N A

20277 Colorado Chalk Bluffs, Weld County 4 July 1943 Syl vilagus floridanus 2 R.M . L. 5 July 1943 Lepus sp. 5 11 20282 10 July 1943 Sylvilagus sp. 7 10 5 20283 Rock Port, Weld County 10 July 1943 Lepus sp. 1 5 20284 Chalk Bluffs, Weld County 22 Aug 1943 Sylvilagus sp. X 1 20461 16748 Idaho 16 mi W Grandview, Owhyee 16 June 1939 Lepus californicus 2 1 County Rupert, Minidoka County 16 June 1939 Sylvilagus sp. 2 2 16750 Gem County 27 July 1940 nuttalli 1 18414 Payette County 29 July 1940 Lepus californicus 1 18415 Gem County 1 Aug 1940 Speotyto cunicularia 1 18417 2 Aug 1940 Lepus californicus 1 18418 Payette County 9 Aug 1940 Peromyscus maniculatus 1 18422 10 Aug 1940 Lepus californicus 4 5 1 18423 Shoshone Falls, Jerome County 30 June 1941 Sylvilagus sp. 1 18887 7 mi E. Shoshone Falls, Jerome 13 July 1941 Neotoma cinereus 2 6 18924 County Clifton, Franklin County 27 Mar .1942 Lepus townsendii 4 24962 2 mi S E. Arco, Butte County 17 June 1947 " californicus 2 23810 1 mi W of Naf, Cassia County 19 May 1948 9 25585 Ruby Mountains, Owhyee 2 June 1961 Sylvilagus nuttalli 51 54 35 B.Y . U. 4932 County 2 June 1961 Lepus californicus 16 4 23 4931 3 June 1961 Sylvilagus nuttalli 1 2 1 4935 3 June 1961 Lepus californicus 11 3 2 4937 NE Ruby Mountains, Owhyee 4June 1961 20 34 4947 County Ruby Mountains, Owhyee County 4June 1961 Eutamias minimus 11 4945 Blacks Ranch, Bruneau, Owhyee 9Aug 1961 Lepus californicus 2 5030 County Blacks Ranch, Shoofly Creek, 6 Aug 1961 Sylvilagus nuttalli 3 5031 N Owhyee County I.O \ Table 1. (continued)

No . Collected Host Animal Collec ti on No. State Locality Date L N A

5034 Idaho Nip and Tuck Creek, OWhyee 9 Aug 1961 Sylvilagus nuttalli 7 3 B.Y.U . County Arco Hiway, 15 mi W Idaho 14Aug 1961 Lepus californicus 20 7 2 5055 Falls, Bonneville County 14Aug 1961 Sylvilagus sp. 2 5056 15508 Montana Big Ditch, E Corvallis, Ravalli 22 Mar 1937 X X X R. M.L . County Rocky Mountain Lab, Ravalli 18 Mar 1940 Lepus americanus 1 18788 County Custer County 17 Apr 1940 1 18381 Powder River County 18 Apr 1940 Sylvilagus nuttalli 1 2 2 18383 18 Apr 1940 1 1 2 21 Aug 1940 sp. X X X 17069

6 Sept 1940 1 1 17187 Birch Creek, E Corvallis, 23 Aug 1941 40 19339 Ra villi County Willow Creek, Ravalli County 1 Aug 1944 5 40 6 21298 E of Corvallis, Ravalli County 8 Sept 1944 40 21061 N of Moiese, Lake County 6 Oct 1944 5 21405 Coat Pit, Ravalli County 19 Oct 1944 45 6 21410 Birch Creek, Ravalli County 27 Oct 1944 1 2 1 1• 21413 27 Oct 1944 1 21414 3 mi E of Corvallis, Ravalli 11 June 1945 5 35 21732 County S Fork of Birch Creek, Ravalli 6July 1945 2 9 21846 County 8 mi NE of Corvallis, Ravalli 11 July 1945 Pica pica 2 21872 County 11 July 1945 Sylvilagus sp. 2 21873 H Horn Ranch, Hamilton, Ravalli 3 Apr 1946 Lepus sp. 16 22258 w County 0 Girds Creek, Ravalli County 28 May 1946 Sylvilagus sp. 6 14 16 22378 Table 1. (continued)

No. Collected Date Host Animal Collection No. State Locality L N A

Montana Bench E of Corvallis, Ravalli 22 Apr 1947 Sylvilagus sp. 32 27 43 R. M . L, 23643 County Swanson Ranch, Big Ditch, 21 June 1949 15 70 90 26569 Ravalli County Hwy. 93, Goosebay, Flathead 7 May 1950 Lepus sp. 6 27578 Lake 10 mi E Winnett Petroleum 28 Mar 1958 Sylvilagus auduboni 1 6497 County Ptyor Mtns . , Carbon County 8 July 1958 1 1 6480 Bannock, Beaverhead County 21 May 1959 idahoensis 9 1 4 6487 21 May 1959 Lepus californicus 1 6498 21 May 1959 Sylvilagus nunallii 3 6499 21 May 1959 Lepus californicus 1 1 6500 Nevada Barrell Springs, Esmeralda 2 July 1941 Marmota flaviventris 1 I• 19005 County McGill, White Pine County 8 June 1981 Lepus californicus 1 5118 Pioche, Lincoln County 8 June 1961 2 5124 McGill, White Pine County 8June 1961 3 5099 Pioche, Lincoln County 8June 1961 2 5126 Wells, Elko County 10 Aug 1961 17 2 1 5039 Elko, Elko County 10 Aug 1961 Sylvilagus idahoensis 6 12 5047 10 Aug 1961 33 10 5052 10 Aug 1961 sp 2 5046 Wells, Elko County 11 Aug 1961 Lepus californicus 7 1 2 5051 11 Aug 1961 Sylvila.gus idahoensis 36 2 5043 11 Aug 1961 Lepus californicus 4 4 5048 11 Aug 1961 5 5050 11 Aug 1961 Sylvilagus sp. 12 5045 Mercury, Nye County llAug 1961 Lepus californicus 4 5041 Mercury, Code JM(G), Nye 15 Aug 1961 " 1 5158 County w Mercury, Code CE(G), Nye 1 Nov 1961 Sylvilagus audoboni 1 49 5996 1--' County Table 1. (continued)

No. Collected State Locality Date Host Animal Collection No. L N A

Nevada Mercury, Code IGG3C, Nye 6 Nov 1961 Thomom ys umbrinus 1 R.M.L. 5973 County Mercury, Code 12 E, Nye 21 Nov 1961 Sylvilagus nuttallii 1 5992 County Mercury, Code CE(G), Nye 22 Nov 1961 6 5977 County Mercury, Code TE(G), Nye 28 Nov 1961 auduboni 2 32 5995 County Mercury, Code CE(G), Nye 14Mar 1961 Melos piza lincolnii 1 5978 County New Mexico Quemado, Catron County 15 July 1924 Sylvilagus auduboni 1 16917 Otero County 3 May 1940 1 18428 20 mi N Truth or Consequences, 21 July 1963 sp. 2 3 B. Y. U. 6312 Sierra County Oregon Baker, Baker County 20 June 1922 Lepus californicus 7 R.M.L, 15960 20 June 1922 Sylvilagus nuttallii 1 16682 21 June 1922 idahoensis X X 6 16684 Upper Klamath Lake 12 June 1940 Lepus americanus 2 16240 12 June 1940 2 16241 12 June 1940 2 16244 12 June 1940 X X 2 16245 Hood River County May 1941 ~ sapiens 1 18891 .. Roxy Ann Hill, 3 mi E Medford, 9 July 1958 Lepus californicus 3 54 B. Y. U, 4997 Jaokson County Dairy, Klamath County 25 Mar 1961 Sylvilagus nuttallii 1 5114 Kinsley AFB, Klamath Falls, 26 Mar 1961 Lepus californicus 1 5003 Klamath County Utah Millard County 27 Apr 1940 Sylvilagus nuttallii 1 R. M.L. 18356 10. 7 mi E Thompson, Grand 1 Sept 1940 sp. X X 17142 County Diamond Valley, Washington 1 15 July 1943 Lepus californi cus 1 B, Y. U. 3588 w County l'v Table 1. {continued)

No. Collected State Locality Date Host Animal Collec tion No . L N A

Utah Millcreek Canyon, Salt Lake 21 July 1946 Lepus ca li fornicus 2 5 R. M.L. 24657 County 4 mi N Green River, Emery i2 June 1947 Sylvilagus sp. 3 24658 County Green River, Emery County 23 June 1947 6 24646 4 mi SE Valley City, Grand 23 June 1947 2 24643 County 1 mi N Lehi, Utah County 3 Apr 1948 12 40 25579 Clear Creek Camp, Tooele 20 May 1948 18 25580 County 38May 1948 Peromyscus maniculatus 1 2558 1 15 mi S Hatch, Garfield County 17 June 1948 Sylvilagus sp. 2 25582 Elberta, Utah County 3 July 1949 Lepus californicus l B. Y. U. 55 7 July 1949 3 2 5 59 Cedar Valley, Utah County 21 July 1949 1 66 21 July 1949 1 2 66 21 July 1949 1 4 66 Rush Valley, Utah County 16Aug 1949 1 81 West Mountain, Utah County 29 Aug 1949 1 1 94 Ce.dar Ft. , Utah County 14 Sept 1949 Sylvilagus nuttallii 2 5 84 14 Sept 1949 N 1 2 83 Heber, Wasatch County 30 Sept 1949 1 91 30 Sept 1949 1 91 30 Sept 1949 L""pus californicus 1 60 Enterprise, Wa~hington County 30 Oct 1949 SylvUag:usidahoeosi.s 27 52 128 Arches Natl Monument, Grand 10 May 1950 mmallii 2 2 154 County Moab, Grand County 9 Sept 1950 sp. 12 2 207 Beaver, Beaver County 3 Nov 1950 Lepu.~ californicus 6 269 St. George, Washington County 4Nov 1950 SyJvUagus audubo,,i 2 267 4 Nov 1950 2 272 w Zion Natl Park, Washingtan 17 Dec 1950 sp. 4 347 w County Table 1. {continued)

No. Collected Collection No. State Locality Date Host Animal L N A

347 Utah Zion National Park, Washing ton 17 Dec 1950 Sylvilagus sp. 1 B. Y. U. County St. George, Washington County 19Dec 1950 5 7 366 20 Dec 1950 7 6 372 Beaver ·Dam, Washington County 21 Dec 1950 " auduboni 8 56 315 Sand Dunes, Juab County 8 Apr 1951 Lepus californicus 3 2 5 59 Bluff, San Juan County 4May 1951 Sylvilagus sp. 10 633 " H 4May 1951 Thomomys bottae 16 633 4May l.951 2 12 633 Antelope Spring~. Delta, Millard 1 June 1951 Lepus californicus 6 12 771 County u 1 June 1951 X 2 771 Buckhorn Reservoir, Emery 9 June 1951 Sylvilagus sp. 3 9 847 County Tropic, Garfield Cpunty 12 June 1951 Lepus c.alifomicus 8 767 12 June 1951 30 3 767 Fisher Tower.s, G:i:andCounty 12 June 1951 Sylvilagu s sp. 1 769 2 mi S Tropic, Garfield County 12 June 1951 Lepus californicus 50 1 767 12 June 1951 1 1 767 Mercur, Tooele. County 26 July 1951 2 35 15 1074 11 mi E Met c ur, Utah County 27 July 1951 Sylvilagus nutta.llii 7 1 1080 27 July 1951 Lepus californicus 90 3 1076 27 July 1951 H 20 1 1077 27 July 1951 25 1 1081 1076 " Fairfield, Utah CoUIIty 27 J1.1ly 1951 X 10 6 Mercur, Tooele County 27 July 1951 Oberholseria chlorura 2 1094 27 July 1951 5Jlvilagus mmallil 2 55 8 1080 Fairfield, Utah County 27 July 1951 Citellus vaxiegatus 1 1082 11 mi E Mcrcur, Utah County 27 July 1951 L::Pus californicus 75 6 6 1077 27 July 1951 4 2 2 1081 Mt . Pleasant, San Pete County 2 Aug 1951 Sylvilagus nuttalll.i 1 12 1105 w Dinosaur Natl Monument, Uintah 14 Aug 1951 Lepw townsendii 2 1259 ~ County Table 1. ( continued)

No. Collected State Locality Date Host Animal Collection No. L N A

Utah Dinosaur Natl Monument, 14Aug 1951 Lepus townsendii 1 1 B. Y.U. 1258 Uintah County Echo Reservoir, Summit County 15 Aug 1951 Sylvilagus sp. 52 20 1290 16 Aug 1951 4 1 1291 Manila, Daggett County 16 Aug 1951 Lepus townsendii 1 1292 16 Aug 1951 Sylvilagus sp. 1 1287 16 Aug 1951 9 1288 Cedar City, Iron County 4 Sept 1951 Lepus californicus 50 1453 4 Sept 1951 1 1454 4 Sept 1951 9 1452 Navajo Wells, Kane County 7 Sept 1951 Sylvilagus sp. 1 1500 .i:t " 7 Sept 1951 Lepus califomicus 1 1506 45 mi E Kanab, Kane County 8 Sept 1951 Sylvilagus sp. 21 3 2 1511 Orderville, Kane County 8 Sept 1951 4 5 1512 Grafton, Washington County 5 Apr- 1952 12 8 5 1746 5 mi S Myton, Duchesne County 4June 1952 Neotoma cinerea 1 3366 Lucin, Box Elder County 18 June 1952 Sylvilagus sp. 2 2119 Kelton, Box Elder County 19 June 1952 Lepus californicus 1 2148 Lucio, Box Elder County 19 June 1952 Sylvilagus sp. 3 1 2140 Locomotive Springs, Box Elder 19 June 1952 1 3 4 2188 County Castleton, Grand County 2 July 1952 1 2243 15 m1 W Fillmore, Millard 11 July 1952 Lepus califomicus 6 2271 Couhty Panguitch, Garfield County 23 July 1952 6 2429 24 July 1952 1 2 3 2430 24 July 1952 Sylvilagus sp. 5 2428 Piute Reservoir, Piute County 26 July 1952 Lepus californicus 1 1 2452 Cedar Valley, Utah County 30 July 1952 1 2 1 2467 30 July 1952 3 2470 Paradise Valley, Sevier County ID Aug 1952 2 3 2610 w Cedar Valley, Utah County 31 Aug 1952 3 2717 VI 4 Oct 1952 1 3 2747 Table 1. ( continued)

No. Collected State Locality Date Host Animal Collect ion No. L N A

Utah Fish Springs, Juap County 28 Mar 1952 Lepus califomicus 1 B.Y . U. 3025 Cedar Valley, Utah County 22 Apr 1952 Sylvilagus nutta llii 150 8 8 .. 3138 Lucin, Box Elder County 23 May 1953 Oberholseria chlorura 9 3251 23 May 1953 Sylvilagus nuttallii 18 8 7 3274 23 May 1953 Lepus californicus 1 3279 Chimney Rock Pass, Utah 28 May 1953 1 3314 County 28 May 1953 2 3313 28 May 1953 3 3315 Red Creek, Daggett County 10 June 1953 Sylvilagus sp. 76 3 4 3389 Brown Parks, Daggett County 10 June 1953 au4'uboni 1 9 8 3393 Willow Creek, Daggett County 12 June 1953 2 3 3408 12 June 1953 Lepus townsendii 1 3406 Chimney Rock Pass, Utah 21 June 1953 " californicus 1 1 3485 County 4 mi N Aitzona-Utah line, 12 July 1953 1 3 3567 Hwy 59 Veyo, Washington County 16 July 1953 Sylvilagus auduboni 1 3597 Heiroglyphic Gap, Iron County 17 July 1953 10 1 3616 8 mi N Enoch, Iron County 17 July 1953 Lepus californicus 1 3614 Heiroglyphic Gap, Iron County 17 July 1953 3 3615 Airport at Crossroads June., 17 July 1953 H 7 6 3612 Iron County Enoch, Iron County 17 July 1953 6 3613 Spty, Garfield County 21 July 1953 Sylvilagus auduboni 1 3 4 36£4 Koosharem, Sevier County 23 July 1953 Lepus californicus 1 3683 23 July 1953 4 20 3681 Delta, Millard County 31 July 1953 1 1 2 2708 Dugway, Tooele County 1 Aug 1953 1 3709 Callao, Juab County 11 Aug 1953 5 3727 Flowell, Millard County 24Apr 1954 1 3 3826 Government Creek, Tooele 18Apr 1957 1 5371 w (}'\ County Table 1. (continued)

No. Collected State Locality Date Host Animal Collection No. L N A

Utah Pinrura, Washington County 26 Jan 1960 Sylvilagus sp. 7 B. Y. U. 4984 W Look out Pass, Tooele County 15 May 1960 Leeus califomicus 10 5469 Dewey , Grand County 6 June 1960 Syl vilagus auduboni 3 3 4750 Hanksville, Wayne County 9 June 1960 sp. 3 2 4701 10 June 1960 Lepus califomicus 6 4709 l0Jtme 1960 Sylvilagus sp. 1 5 4706 11 June 1960 100 13 17 4707 Huntington, Emery County 29 June 1960 6 6 4731 5 mi Ela Sal, San Juan County 7 July 1960 3 4756 Cottonwood Creek, near Bluff, 8 July 1960 Lepus califomicus 1 4759 San Juan County Skull Valley, Tooele County 2 Oct 1960 61 5 5468 3 mi E Indianola, San Pete 11 Feb 1961 6 3 4877 County Diamond Valley, Washington 26 Feb 1961 2 1 4896 County 11 mi NW St. George, Washing- 26 Feb 1961 Syl vilagus auduboni 1 4897 ton County 28 Feb 19€1 Lepus californicus 1 4890 2 mi NE Clover, Tooele County 9 Mar 1961 67 3 3 6253 4 mi W Rosette, Box Elder 17 Mar 1961 222 4900-1 County 17 Mar 1961 147 4900-2 Rosette, Box Elder County 17 Mar 1961 15 5113 3 mi W Rosette, Box Elder 17 Mar 1961 Syl vilagus idahoensis 382 20 26 49-01 County 17 Mar 1961 Lepus califomicus 56 18 42 4900 Rosette, Box Elder County 18 Mar 1961 3 4907 11 mi SW Rosette, Box Elder 18 Mar 1961 Sylvilagus nuttallii 3 4906 County Rosette, Box Elder County 18 Mar 1961 Lepus califomicus 1 4909 2 w 11 mi SW Rosette, Box Elder 18 Mar 1961 4 i 4904 -..J County Table 1. (continued)

No. Collected State Locality Date Host Animal Collec tion No. L N A

Utah Terry's Ranch, Beaver Darn, 7 Apr 1961 Sylvilagus auduboni 10 B. Y. U. 6426 Washington County West Tintic Mts., Juab Comty 21 Apr 1961 Lepus califomicus 30 54 49 4913 Sheeprock Mts., Juab County 21 Apr 1961 Syl vilagus auduboni 9 1 4916 21 Apr 1961 9 22 4916 3 mi W junction of Green and 22 Apr 1961 Lepus califomicus 19 17 7 4917 San Rafael Rivers, Emery County Buckhorn Reservoir, Emery 28 Apr 1961 Syl vilagus sp. 4 14 4919 County 28 Apr 1961 Lepus californicus 8 56 4918 Goblin Valley Area, Emery 29 Apr 1961 Sylvilagus sp. 9 5 4924 County Hanksville, Wayne County 30 Apr 1961 Lepus califomicus 3 3 4925 Hanksville, Green River Road, 30 Apr 1961 Sylvilagus sp. 4 14 4926 Wayne County Terry's Ranch, Beaver Dam, 6 May 1961 52 6425 Washington County Rosette, Box Elder County 30 May 1961 idahoensis 1 7 10 4927 30 May 1961 Lepus califomicus 9 16 42 4928 Jericho, Juab County 1 June 1961 2 50'22 St. George, Washington County 3Jwie 1961 120 5726 3 June 1961 1 4971 3June 1961 1 4979 Snows Canyon, Washington 3 June 1961 Sylvilagus sp. 2 5117 County St. George, Washington County 4 June 1961 auduboni 200 5885 4 J .me 1961 sp. 8 24 4993 4 June 1961 1 4963 4 June 1961 2 4 50'21 4 June 1961 Lepus califomicus 2 5140 Manila, Daggett County 27 June 1961 w Sylvilagus sp. 2 5186 00 Vernal, Uintah County 28 June 1961 1 5111 Table 1. (continued)

No. Collected Collection No. State Locality Date Host Animal L N A

Utah Vernal, Uintah County 28 June 1961 Sylvil agus sp. 1 B. Y. U. 5011 West Tintic Mts, Juab County 30 June 1961 Dipodomys ordii 1 5120 30 June 1961 Syl vilagus sp. 75 5 5116 30June 1961 2 4977 7 mi w Tintic Mts. , Juap 1 July 1961 Lepus califomicus 1 5090 County 14 July 1961 5 7 4969 Jericho, Juab County 14 July 1961 1 5009 Rosette, Box Elder County 7 Aug 1961 1 3 5026 Lynn Pass, Box Elder County 7 Aug 1961 1 1 5028 Rosette, Box Elder Cowity 7 Aug 1961 Sylvila~ sp. 1 1 "5147 Mercur, Tooele County 11 Aug 1961 Lepus c";'i:fomicus 4 1 1 504-1 Levan, Juab Cotmty 30 Aug 1961 131 12 5069 Diamond Valley, Washington 30 Aug 1961 21 2 5141 County Colwnbia Mines Area, Iron 19 Dec 1961 Sylvilagus sp. 2 13 3 5018 County 26 mi W Jericho, Juab County 20 Oct 1962 Lepus califomicus 17 2 6100 8 mi W Nephi, Juab County 2 Mar 1963 1 6246 2 Mar 1963 3 6249 2 mi NE Clover, Tooele County 9 Mar 1963 2 8 14 6254 8 mi W Nephi, Juab County 16 Mar 1963 1 1 6259 16 Mar 1963 Sylvilagus sp. 1 6256 5 Apr 1963 Lepus califomicus 1 3 3 6262 5 Apr 1963 116 1 6 6263 5 Apr 1963 18 6261 5 Apr 1963 1 1 6264 S Tintic Mts. , 25 mi S Eureka, 17 Oct 1964 3 7 6585 Juab County Washington Walla Walla, Walla Walla 31 May 1922 Sylvilagus nuttallii 1 R. M. L. 16675 County w Easton, Kittitas Collllty 22 July 1922 Lepus americanus 1 15962 '° Table 1. (continued)

No. Collected Date Host Animal Collection No. State Local ity L N A

1 15963 Washington Kent, Kings County 25 July 1922 Mustella satuvata R. M. L. Alpha, Lewis County 29 July 1922 Lepus americanus 23 2 16700 29 July 1922 X X 16784 11 mi N Pasco, Franklin 3June 1939 Sylvilagus sp. 5 3 16733 County Sunny~de, Yakima County 4June 1939 1 16734 4June 1939 Lepus califomicus 2 16735 Spokane County 5 Aug 1939 Sy:lvilagus sp. 4 2 16408" Lake Tapps 14 Aug 1939 Lepus americanus X X 16969 18411 Spokane County 19 July 1940 Syl vilagus n uttallii 2 3 22 July 1940 2 2 18412 Moses Lake, Grant County 29 Aug 1940 100 18745 29 Aug 1940 100 18761 8 mi W Odessa, Lincoln 9 June 1949 idahoensis 6 18 13 26679 County Grays Harbor County 24 Mar 1949 Lepus americanus 3 16 36561 H Columbia River Hwy, Skamania --- 2 25 33487 County 16848 Wyoming Rawlins, Carbon County 24 Oct 1922 Syl vilagus nuttallii X X 24 Oct 1922 1 16851 22 June 1923 Lepus americanus X 16854 16365 Sweetwater County 30 June 1939 Syl vilagus n uttallii 1 5 July 1939 2 16293 12 July 1939 4 16300 11 Aug 1939 Dipodomys ordii 1 16322 Sheridan, Sheridan County 3May 1940 ~a pica budsonica 1 18386 Oleyenne, Laramie County 24 July 1943 Syl vilagus floridanus 4 6 20297

+-- 0 Table 2. Collection Data for Har::maphysali.s cbordelilis in the Western United States.

Stare Localiry Date Host Animal No. Collected CoUection No. L N A

California Fort Barry, Ma~i.n Coumy 9 Jan 1952 Cloth drag Carpenter (1956) Montana Roy, Fergus County 23 Sept 1931 Sage hen 43 5 !3'7 R.M.L. 7941 Lew Lsron, Fer.gus County :06 Od 1%1 Grouse' 6 Tbomomys sp. Cooley (l.946) 22 Aug 1959 Blue grouse C,:; R. :tv!.L. 35404 2,J Sepl 1949 Pir; taU prar.ie chick.er 4 35555 Fergus 4 SP.pt 1960 Sharp=!ai.led grouse ~; 8.Y.U. 6501 ·,washi-1gto'l Fr.,1ltla:1d, Sk.ve'.':S ?,'7 Apr 1J10 CUih/ 1 Bishopp 1564 4 rr/. SW e.stpcn, c:uy'"' Ha~x>r '2.l J ,Ele 19t:9 Whl.te~,. r:cwnecJspa,:::ow 1 l R.M.L. 3'?847

Litti Co<;se. Car,yor;, S",,',:iJar; 18 Sepr 19e 1 Blue grou:s1:->. 3'Hi'l6

+' 1----' 42

RESlJLTS

Haemaphysalis leporispalustris

Geographical Distribution

Haemaphysa1is leporispalustr1s is distr1buud through~ out North and South America, Gregson (1956) reported extensive distribution from British Columbia to Newfoundland in Canada. It has also been collected as far north as CirclEi

Alaska by Philip et al. (1954), aLd as far south as Argentina by Aragao (1938). Kohls (1960) has brought up to date the information on geographic distributiun~ Huffmaf'.n (1962) updated all existing information on this tick and others in

Mexico.

Nuttall et al. (1915) stated that its dlstribution is extensive in the United States, Later a distribution map was presented by Bishopp and Trembley (1945) showing actual and probably distributio11o Addlt.iona] data w(,rc: published

by Cooley (1946) and Bequaert (1Y46) 1 who stated that H. leporispalustris occurs widely in the United States. Since

1946 new published data have not extPnded the known distrlbU1:ior; 43 of the species (Kohls, 1960).

Data on geographical distribution presented in this study are a result of years of field surveys by the Zoology and Entomology Department, Brigham Young University" Other institutions have also loar;,Ec0 d tl:Hair data and/or preserved specimens for this study.

Table 1 lists coll~ction loc.aLiLies in chronological order by state, location, date~ host, and numbers of ticks per host. From these data the distr1bution for g. leporispalustris has been designated. FigurEs 1 to 6 show places where specimens have been collected. Each dot represents a collection locality and does not indicate the number of timEs that ticks were taken from a sp~cific locality. Since col-

0 lection data for Utah an, abundant 0 Figure 6 shows localitH S for this state. No attempt was made to 'c,sLabl1sh a hypothe- tical distribution.

Seasonal Occurrence

In 1912, HookE:0r, Bishopp and Wo::;d stated that Ho

1 eporispa· ·1us t ris" " cou ld b e 1··ounr 1 at a. 11 seaso~s in all stages of development. in thE: United StatE::s,., Undt 0 r "Ick,al condi.- tions" Nuttall et aL (1915) runark.-,-'d thaL two lifE, cycle3 per year might be observedo Harkema (1936) noted tick activity the year round in North Carolina, and suggested that two life cycles per year existed.

Bequaert (1946) claimed that thLs tick hibernates away from its hosts during the winter in the northern part of its distribution, whereas in the southern extension of its distribution H" leporispalust:ris is actiV(c throughout. t.h<:, yEaL

A cessation of activity "during the colder parts of the year in the more northern states" was also reported by Bishopp Bnd

Trembley (1945)0

Beck (1955) noted an apparent modification of its life cycle in the north-south distribution in Utah. For eastern

and central Washington 9 Bacon, Drak.E, and M1 ller (1959) observed a period of inactivity during tbe winter months but the exact date of hibernat1.on was never di:0 terrnirwd, Yeatter and Thompson (1952) four!d that by delaytng th(' rabbit nuntl.:-1g season in Illinois until fr~e21n~ temperatures forced H. leporispalustris into hibercation, less cases of tularemia occurred among hunters. Later, Mohr and Lord (1960) concluded that hibernating stageB of adults and nymphs existed in northern Illinois. In Mirn1t2sota this same ti.ck was ob-

served throughout the year (Cr~en 9 Evans and Larson 9 1943). 45

Seasonal occurrence for all developmental stages of

H. leporispalustris is shown by month for each state in the western United States in Figures 7 to 20. Each figure rep~ resents total ticks per month during different years in all developmental stages. For the majority of thti western stau'.s the data are sparse. Collections have rot en made sys tern~ atically or continuously.

Host-Parasite Association

Although H. leporispalustris has been taken on a variety of hosts in the United States) workers generally agree that leporids are the preferred hosts, Green et al (1943) reported that immatures were abundant or. grc,use in Mi..rmesota.

Bishopp and Trembley (1945) list~d over eighty different hosts for this tick, but noted that leporids are the primary hosts. Bequaert (1946) and Cooley fl946) reported diverse hosts for this tick, but alsc mentioned that it is most

"common on lagomorphs." In t.hel...r Washington study 9 Bacon et al (1959) found this tick only on lagomorph::L Coffey (1954) reported that hosts of H. leporispalustris in Utah included, besides leporids, small ground-inhabiting birds and rodents.

Beck (1955b) also stated that in Utah tht:s leporids appeared 46 to be the preferred host of this tick,

In another study by Mohr, Beck and Brinton (1964) in

northern California 1 mention is made that "H. leporispalustris were commonly encountered on brush rabbits as larvae, nymphs and adults"" The immature stages were also collected

from towhees and wrens. Similarly 9 Hansen (1964) in Oregon found that of eleven mammalian hosLs this tick was taken only from a lagomorph, Sylvilagus nuttalliL

George (1963) postulated that the adult of the species may be highly host specific because of responses to behavior patterns of the leporidso Since the adult does no climbing at night and responds negatively co light, as demonstrated in the laboratory, Gamin (1963) suggested that the adult is mainly a nest-inhabiting parasite. This may partially ex- plain its host-specificityo In contrast, the larvae and nymph's behavior is that of actively seeking suitable hosts by climbing low shrubs and grasses and waiting for the host to pass.

In this study I found a variety of hosts for H. leporispalustris in the western United StatE:s. These hosts, in- cluding leporids, are shown in Tables 3, 4, 5 and 7, The number of times that a certain developmental stage was collected from each host is also indicated.

Anatomy and Morphology

A comprehensive treatment of this subject has not been attempted at the generic level since Nuttall et al"

(1915) published his findings. EVPTI here 9 complete drawings of all developmental stages of g,, ,leporispalustris are lack~ ing. Cooley (1946) made line draw1.ngs of some anatomical parts in the several stages of development. Gregson (1956) used line drawings of anatomical parts of taxonomic value.

Detailed drawings of external anatomical features observed in adults and nymphs of this species were published by Brin- ton and Beck (1963). Outline drawings of the larvae were also presented. The United S tatE'S Department of Agriculture published a "Manual of Livestock Ticks" (1965) which contain detailed illustrations of the male and female,

During this study all developmental stages of the rabbit tick were illustrated for purposes of comparison with

H. chordeilis, Only the larval g, chordeilis and H. leporispalustris were labeled to conserv8 space. 48

Haemaphysalis chordeilis

Geographical Distribution

No H. chordeilis specimens were col.lected during this study. A sma 11 number wc·re rece i vc0 d on loan from the Rocky

Mountain laboratory, and from Dr. C M" Senger of Washington

State College" From data obtained with the loan material

(See Table II) known distribution in thr:: western Uri.ited States was plotted on Figure 21.

Seasonal Occurrence

Bishopp and Trembley (1945) mentioned that. g. chordeills

col lee tions in t.he south were during "late fa 11 9 winter 9 a.nd early spring and those in the north wpre collected during the

0 spring 9 summer 9 and falL" Javidpour (unpubli.shed thl Sis) reported this tick coll.cctcd during Octc rand November in

Wisconsin.

Published and unpublished records from the Rocky

Mountain Laboratory wen: u::ed for this phase of study, and the results are graphed on Figure 22. 49

Host-Parasite Association

Workers have reported H. chordeilis as a parasite of a number of birds, especially ground-inhabiting species.

Hadley (1909) states that in Vermont turkeys were heavily parasitized. Bishopp and Trembley (1945) have mentioned that throughout the south and north ground-inhabiting birds are the main hosts. Included in their publication is also a table of common hosts. Bequaert (1946) stated that grouse seems to be the preferred hosts in the eastern part of its distribution, In Canada, Gregson (1956) has reported that this tick is a common parasite of upland game birds, Kohls

(1960) reported that this tick was also taken from a cow in Washington. The few records available for the western

United States indicat.es that birds are a conirnon host.

Anatomy and Morphology

As with H. leporispalustris, this tick has not received much attention with respect to anatomical features since Nuttall et al. (1915). Cooley (1946) and Gregson

(1956) both presented li.ne drawings with structures of taxonomic value. In this study all stages were illustrated and are presented to compare with H. leporispalustris. 50

DISCUSS ION

Bishopp and Trembley (1945) published a geographic distribution map for H, chord(,ilis an,J H. le:eorispalustris which included probable di.stributiotL ln this study I have extended such informati.on by bringing r~cords up to date,

But more irnportant 9 I have pinpointE:d actual collection localities,

Haemaphysalis leporispalustris appears to be wide= spread. But within this wide distribution there occur local areas where this tick is not collected, Also the tick may not occur one year and yE:t be observed a season later, Such a situation was encountered by Bacon et al, (1959). More than one season's investigatiot~s in an area are necessary to

cietermine the problems of distribution 1 and ecological factors influencing same,

An interesting problem concerns that of factors determining altitudinal distribution. In Utah although a few hosts of H, leporispalustris have been reported at montane elevations, this tick has not been collected at such elevations, Beck (1955) observed the following about distribution 51 of the rabbit tick in Utah: "It is more common at lower altitudes in basins, valleys and desert-like conditions whether flatland or canyon. This may be due Ln part to certain preferred hosts, members of the family leporidae being more abundantly distributed at lower elevations."

Although birds in the western United States are not

the normal host of this tick 9 they arE: sometimes found parasitized by the immature stages. A study of the migratory habits of these birds may also extend information on distribution.

Geographic Distribution

Arizona. With reference to collections of ticks in

general 9 there is a paucity of published information. I have examined the records at the Rocky Mountain Laboratory and those supplied by the Arizona Fi.sh and Game Commission.

According to available data H. leporispalustris is not widely distributed. A more comprehensive exami.nation of the parasitic fauna of lagomorphs in Arizona would matE,rially extend the knowledge on geographic. distribution of this tick. Avail- able records show H. leporis£alustris to be distributed in

Maricopa 9 Mohave 9 and Navajo Counties (See Table I and Figures 52

2, 3).

California. Haemaphysalis leporispalustris is distributed along the coast with a few records from the central valley (See Table I and Figures 1-4). The majority of records, however, are from Northern California, especially from the extensive collections of the Hastings Reservation, Monterey

County. It may be generally concluded that any concentrated study on parasite-host relationships in any locale will pro- vide a better understanding on distribution than the occurrence of occasional collections.

Colorado. The nine collection records supplied by the

Rocky Mountain Laboratory are all that were availablf: for this study. They are insu£ficient to make an accurate state- ment on the distribution of the rabbit tick Ln this state.

These ticks, taken from rabbits, were from Boulder, Costilla,

Moffat and Weld Counties (See Table I~ Figs. 2-4),

Montana. Thirty records throughout the state corres- pond quite well with the distributions of leporid hosts as outlined by Hall and Kelson (1959) (See Figs. 1-5). This tick has also been collected from L. californicus and S. 53

Geographic distribution oi' ll• le-oorisrio.lnstrijl tc.ken from 1• Fig. 1. arnericonus O , and L. californicus ... • The lined area represents the distribution of-b• americ~~llli_ and the rough area that of I:• californicug (After Hall e.nd Kelson, 1959). 54

Fig. 2. Geographic distribution of' g, J_eporis"alustri~ taken from Le"'-!§ spp, ■ , 1,. !,_o,msenctg O , and non-leporid hosts .A , Dotted andarea Kelson,represents 1959). the distribution of J:,, :t<,wnsencii1 (After Ball 55

~ ~ ··'" ·O ··11111111 !II ll!]i.ll.il'!_!lil 111111I l>r11( 1111111I ljj 11 lr i/ljil!Jl"tlj,~,~; ! ii,; :,11'1n,:ll1 I I 0 I J l !/ I l l - ~ I l!111 111111111111111,,r!,l 1j I ~ 'I I ;q ! l ! II l 1/ I I i [ I ,: /:; 11 ll 1 ·i! It! .. . ·---- ?

'-- 00 0 ~ 11:,

[11[11111 I //Ii/! I iil!j' 1 . I l I 'I 111!I 111/lllll-111/ I• I .t11, ! ' I 1li!i 1I • Ii 11 l I Ii Ill Ill ·,,i,,!I'/ !IIIIIIIIN,l.li/llillll 111111111111111/1111 I 1111111111111111 II•/' I /I ) l I 1 I 111/11,lll/11/l/1.J..i/ll/l/lllll// /lrlill/ii1/ I ~ I I I 'I I I 1 1111 jl' I I I 1/i II 1 II I . ~ ~11 ;11 ,1/ nil ·•11 0 !Ill '!'ii(I I111111111111111 ~ 111,11111

Goographic distribution of I.I• 6£_.".)_orisuu·i.ustris tclcen from :Nlvilti.;:;us Fig. J• spp. 0 , and §• 0.1.1£!.!'~'fliA . Lined area represents the ciistribu- tion of ~• audubonl (After Ball and Kelson., 1959). 56

,·. .i··

..• ,, C· •aO .•.J • • -;.:-

Fig. 4• Geographic distribution of g. l§:l..12.£!.k~us~r:,.istaken from~. 1 • .A C' i' ■ l"1\,, ~~~b .a , £• ~.2J'1'1 ac.nu:'l , ana., ~•s 19-..?~}!lli"' • . 0 • ...ue fine-stippled area represents the distribution oi' 2• bachn.anH, the rough arec. that of .§.. florid~, and. the sti:ppled area ~Ii that of~• idahoensj_s (After Hall and. Kelson, 1959). Geographic distribution of li• laporisp~~ri& taken from 2•

Fig. 5• nutta.llli A {After llall o.na.Kelson, 1959) r• 58

idahoensis which the above authors do not mention in Montana a

These records appear to represent a ~ood portion of the state

(See Table l)a

NE,vada O RE:cords from th.is s tatE i udica tE"': distribution in the eastern and southern. portions" Jt would be well to also survey the western part of NFvada. fhough the collection data are few, Fig11t~(~s 1-'> show that!:!, lE:porispalustris

has been taken wi..thin. tlw distribution of its lepori.d hosts a

New Mexico. Only three records were available from

0 this state, representing Catron, Otero 9 and SiE: rra Counties

(See Figure 3). Extensive investigatio~s are necessary to complete a picture of g raph1,al dLst.ributiono

Oregon a Thtc f(,W re._ onh aVELi.lable from this :state were supplied by the, Rocky Mounthl:, l abnratory and from members of field survey partifs s nt nur. Hrigharn Young University.

Figures 1 9 2 9 4 9 und ') 1ndiu1t, that th t1ck has been col-

lected at ditfereol localiti~s 111 the state corresponding to the distributions ot itE' lq)1)rld hr1st • StiU, mon, exten= sive collectior\:s will fill 1n 1lH: gaps in distribution. 59

0 ••

0 0 •

0 0 0

0 A.60 • •0 •• •• • • Fig. 6. Geograohic 6istribution 01' H. leporisnalustris in Utah t9.ken from kr_~ spp. 0 , 0ylvilc:);us-spp. £ -;- ooth Sylvilugus and 1.epus • , and f'roL, non-leporid hosts ■ . 60

Utah. Figure 6 shows that H. leporispalustris has

been collected extensively throughout the state 9 especially along routes of main highways. Parts of the state remain to be sampled. Collection localities are concentrated at valley and foothill elevations; none representing the montane elevations.

Washington. Haemaphysalis leporispalustris has not been extensively collected throughout the state, The most

thorough data have been supplied by Bacon 9 Drake, and Miller

(1959) from Adams and Grant Counties.

Wyoming. The few available records show that H. leporispalustris has been collected in the northern, southern and southeastern portions of the state (See Figs. 1-5), More data are needed to gain a better picture of distribution.

Seasonal Occurrence

An accurate determination of seasonal occurrence in the western United States for H. leporispalustris cannot be described in general terms. Two primary factors make such a determination difficult: (1) the highly variable ecological 61 environments which comprise the biogeography of the western

United States, and (2) the general seasonal observation of tick-host relationships on a twelve month schedule in the many variable ecological situations,

The majority of tick collections in California represent the spring and summer months, Ticks have been collected during all months in the state of Utah, although not systematically. Collection sites in Idaho and Montana are not numerous, and the remainder of the western states have only been sampled superficially on a seasonal basis.

From the observations made, tick activity varies in the north-south distribution in Utah. In coastal as well as inland California, all stages have been reported in the winter months while in Montana no ticks were collected from December to March, Data compiled during this study have added some-

what to the knowledge of seasonal activity in various states 9 but not regionally.

Mohr and Lord (1960) concluded in their northern

Illinois study that hibernating adults and nymphs existed in the H. leporispalustris population. They also determined that each developmental stage had a peak population which was in- fluenced by the activity and relative numbers in the entire 62

population"

Data from the western United States were analyzed to point out possible areas where tick activity may be observed

the year round, and where climate may influence hibernationo

Similarly, an attempt to recognize peak populations, if pos-

sible, was made, Although activity by state may suggest a

pattern as described by Mohr and Lord (1960), these may be misleading in that the data for the western states cover many years. Equally important, collections were not made

systematically nor do they represent one location for a

twelve month period. The figures on seasonal activity rep-

resent total numbers of ticks over a period of many yearso

Arizona. Haernaphysalis leporispalustris has been

taken in April, May, June 9 July 9 and October. Adults were

collected only in June and larvae only during October. Dur-

ing April, May 9 July and October nymphs were collectedo Dur-

ing no month were all three developmental stages taken to-

gether" Additional data are necessary to determine whether

or not this tick is active at other times of the year (See

Figure 7)"

Californiao Most records are from the northern 63 100 Adults - Nymphs _ Larvae :}:'?

J F M Fig. 7. Total munbers of H. leporispalustris occm·ing by month in Arizona (1924, 1938, and 1948).

100 .A..iults - Nymphs __ larvae ::::::::::::::

S O N D

Fig. 8. Tota1 numbers of !!• ~orispaJu.stris occ11!'ing by rno11'th in Colorado between 1925 and 1943. 100 Adults - ·.•, ·.•, Nymphs ,•.· Larvae

50 ·.·.· I1111: J F H A r,i ;j U N lJ

Fig. 9. Total nwnbers of g. Jgporispalustri,2 occu.r·ing in Idaho between 1939 and 1961. 64

portion of the state 9 mainly Hastings Reservation, Monterey

County and San Mateo County (See Table I), The months most . numerously sampled are from April through August, but gener-

ally speaking, g. leporispalustris has been collected in all

developmental stages in California except for February and

November.

Figure 10 shows the total numbers of ticks in all

developmental stages taken in California for which we have

records at Brigham Young University, Department of Zoologyo

Figure 11 represents the collections for San Mateo County 9

1961. This reveals that from May through August all devel-

opmental stages were taken each month. From the total num-

bers of each stage it seems that each has a definite peak

period: the adults peak first and decline, followed by the nymphs, and lastly the larvae,

Records for the entire state indicate activity dur~

ing the winter months in northern California. From Marin

County two adults were reported in January (1945), During

the same month (1943) two nymphs were collected in Monterrey

County. In southern California all three stages have been

observed during December by the author (BYU Field No. 5122)"

Coloradoo Records from this state represent four 65 350 Adults - Nymphs_ 300 Larvae ::::::::=:::::

200

100

Fig. 10. Total numbers of !.!• leporispalustris occuring by month in California between 1920 and 1964.

200 Adults - Nymphs_ Larvae :}}:::::

100

0 J F M A S O N D

Fig. 11. Total numbers of !!.: lepcrispalustris occuring by month in San Mateo County, California in 1961. 66 different years and show that ticks have been collected from

April through August. The April and May collections have not been graphed since numbers of ticks collected were not reported along with the collection data (See Figure 8). Ad- ults were observed during June, July~ and August, with greatest numbers in July. Nymphs were collected in July and

August, and larvae are represented only during July. More data are needed to complete the picture of seasonal occurrence.

Idaho. More records of H. leporispalustris are available for this state than for either Arizona or Colorado. But these data only represent the months of June, July and Aug- ust, making it impossible to comment on seasonal activity on a yearly basis (See Figure 9). All developmental stages were collected during June and August with only the nymphal and larval stages representing July.

Montana. Rocky Mountain Laboratory collection records in our files show that H, leporispalustris has not been collected in this state from November through February. Dr. Glen

M. Kohls (personal communication) of the Rocky Mountain

Laboratory stated that the rabbit tick has been collected during November; but not in the other winter months stated 67 above. Lagomorphs have been taken during these months though not specifically for tick parasites, He further states that this tick is not a winter-feeding species in the northern states.

Figure 12 represents a twenty year period of thirty collection localities mainly around Ravalli County, Montana.

All three stages have been collected in every month that the tick is active, September being the only exception.

Nevada. Tick activity has been observed during March 9

June, July 9 August, and November, All three stages were taken together only during August (See Figure 14). Sampling on a seasonal basis is necessary to extend the activity period of this tick in Nevada.

New Mexico. Three records for H, leporispalustris from this state were available (See Figure 13). All three stages were collected during July, and only the nymphal stage is represented in May.

Oregon. All three developmental stages were collected during January and June, but are not represented in Figure

15 because the data did not include numbers of immature stages, 68 125 .Adults - Nymphs _ 100 lt\rvae

0 J F N D Fig. 12.. Total numbers of !!• leporispalust1:i2 occuring by month in Montana r.e.tween 1937 and 1959.

50 Adults - Nymphs_ 25 Larvae ::::::/:::

0 J ]' M A A S O N D

Fig. lJ. Total nur,1bers cf g. lepori,e,Ealustris occuring by month in New Mexico (1924, 1940, and 1963).

125 Adults - Nymphs - 100 Larvae ::::::::::::::

·,•.· ·.·.·. ::::: 50

•••••

0 J F & 0 N D

Fig. 14. Total numbers of g. ~eporispalustris occuring in Nev&da (1941 and 1961). 69 75 .Ad.ults • Nymphs_ 50 larvae ://::

Fig. 15. Total numbers of g. leporispalustris occLU"ing by month in Oregon between 1922 and 1961.

125 Adults - Nymphs_ 100 larvae :;:::::::::::

0 J F S O N D

Fig. 16. Total numbers of g. leporisQalustris occuring by month in Washington between 1922 and 1959.

50 Adults - Nymphs _ 25 Larvae :}Ji

0 J F M A N D Fig. 17. Total numbers of H. leporispalustris occuring by month in Wyoming 1::etween 1922 and 1943. 70

Tick activity was observed during January; March 1 May, June, and July, Further sampling on a seasonal basis is necessary for this state.

Utah. The rabbit tick has been collected in some stage of its life cycle during every month of the year in

Utah. The Brigham Young University records represent fifteen years of observations, The majority of these records represent the years 1949-1953 and 1960-1963 (See Table I).

In the southwestern part of the state (Washington

County) all three stages are active during the winter months, while no such activity has been observed for Box Elder and

Utah Counties (northern Utah). The northern-most activity

during November was observed in Beaver 9 Beaver County (southern Utah); for December; Columbia Mines Area, Iron County

(southern Utah); for January; Pintura, Washington County; and for February, Indianola, San Pete County (central Utah).

During February and March (1964) I collected jackrabbits from

Box Elder and Utah Counties which were negative for ticks.

In November of 1964 all jackrabbits examined from Utah County by me were also negative for ticks, although fleas were found to be present. All stages have been reported from Box Elder 71

County during March,

Figures 18-20 repre,sent total numbers of ticks taken

from 1949-1953, 1961 3 and 1960-1963 excluding 1961, respec- tivelyo Comparing the first figure with the latter two indicates a shift in numbers of ticks per montho The former shows activity during the summer months while the latter i.n~ dicate spring and early summer activity" This probably re- flects hunting pressure at different times of year rather than shifting in activity of populations.

The 1961 collections (Figure 19) show a great deal of larval activity during Marcho These data represent the northern part of the state (Box Elder County) which are subject to freezing temperatures. It is interesting to note that Javidpour (unpublished thesis) and Mohr and Lord (1960) both reported larval activity in June and not soonero

Washingtono From the Rocky Mountain Laboratory records

ticks have been collected in March 1 May, June 1 July, and

August, All three stages were collected during Juneo The greatest number of ticks were observed during August (See

Figure 16) o

Bacon et al, (1959) indicated in their study that all 72 JOO Adults - Nymphs _ Larvae ::;:::::::::

200

100

0 J F Fig. 18. Total numbers of fl. le_E9ris2aJ.u~tris occuring by month in Utah bet~een 1949 and 1953•

450 :::;: ·,•,· Adults• ·.·.· Nymphs_ 400 Larvae :::::::::::: ·,·.· ::::: .·.·

·.·.· 300 ·.·.·

::;:: ·,·.· ·.•.· .-.·,l,'

200

100

0 1-.· J S O Ii D Fig. 19. Total numbers of tl• leporispalustris occuring by month ih Utah (1961;. 73

125 Adults - Nymphs _ 100 I Larvae ::::::::::::::

·.·.· ··••· fl

0 ...... _ J F J A S O .N D Fig. 20. Total numbers of g. leporispalustris occuring by month in Utah from 1960 to 1964 excluding 1961. 74 developmental stages were taken in February. From jackrabbits collected during December (1954) no ticks were observed. These authors indicated a cessation of activity though exact dates were not given.

Wyoming. An examination of the nine records available for this state show that ticks have been collected during May, June, July, August, and October. Larvae were

observed in October only 1 but this is not indicated on Figure

17 because the number collected is not known,

Host-Parasite Relationship

In the western United States lagomorphs are the preferred hosts of g. leporispalustris though occasionally smaller birds and rodents are parasitized, Observations of records for the western United States does not show that H, leporispalustris is a common parasite of birds though it has been reported for eastern and southern United States. Man and a lynx are two examples of unusual collections. In the laboratory immatures have been fed with some degree of suc-

cess on domestic rabbits by Javidpour (unpublished thesis 1

1964) and chicks (Camin, 1963). A colony has been maintained 75 for several years at the Rocky Mountain Laboratory by feeding the immatures on domestic rabbits and the adults on snowshoe hares (Kohls, unpublished data).

Tables have been made for the western United States,

California and Utah by determining the number of times H. leporispalustris was collected from a given host (See Tables

3-6), It must be remembered that these are not exactly host- preference indices as such since each host was not collected extensively, Most of the data were from chance collections.

These tables do not show the total numbers of ticks nor the developmental stages unless indicated,

Table 3 shows that H. leporispalustris has been collected from eleven hosts in California. These represent 154 collection records, all but nine of which are from leporids.

The non-leporid hosts are deer mouse, lynx 9 squirrel, and several species of birds. The results strongly indicate that

H. leporispalustris prefers leporids over other hosts.

The Utah data represent 197 collections (See Table

4) indicating the number of times that the rabbit tick was encountered on a particular host. From leporids !:!.• leporis - palustris was taken 189 times. The other eight collections include a bird and several rodents. These data indicate that 76

Table 3, The number of times that Haemaphysalis leporispalustris was collected from a given host species in California.

Host

Citellus beecheyi 4

Lepus californicus 18

Lepus species 1

Lynx rufus 1

Peromyscus maniculatus 1

Pipilo erythrophthalmus 4

Pipilo fuscus 1

Sylvilagus audubonii 76

Sylvilagus bachmani 34

Sylvilagus species 9

Thryomanes bewicki 2

Unknown species 3 77

Table 4. The number of times that Haemaphysalis leporispalustris was collected from a given host species in Utah.

Host

Citellus variegatus 1

Dipodomys ordii 1

Lepus californicus 102

Lepus townsendii 4

Neotoma cinerea 1

Oberholseria chlorura 2

Peromyscus maniculatus 1

Sylvilagus spp, 52

Sylvilagus auduboni 14

Sylvilagus idahoensis 3

Sylvilagus nuttalii 12

Thomom:xs bot ta e 2 78

Table 5, Haemaphysalis leporispalustris adults taken from hosts other than leporids in the western United States,

Host

Homo sapiens 1

Lynx rufus 1

Marmota flaviventris 1

Neotoma cinerea 1

Peromyscus maniculatus 1

Speotyto cunicularia 1 79

Table 6. Numbers of Haemaphysalis leporispalustris in all developmental stages taken from Lepus spp. and Sylvilagus spp. in Utah.

Lepus spp.

Larvae Nymphs Adults

Total 944 407 416

Average/stage 4.0 4.0

Average/rabbit

Sylvilagus spp.

Total 1105 436 387

Average/stage 13.8

Average/rabbit 24.1 80 in Utah the leporids are the preferred hosts.

In the western United States Ha leporispalustris adults collected from hosts other than leporids are very few (See Table 5). The adults have been collected just once from each host indicated. This would suggest that these hosts are seldom parasitized by the rabbit tick. On the other hand, Table 7 indicates the numbers of larvae and nymphs that have been collected from non-leporid hosts in the western United States, The list is longer and includes a small variety of rodents and ground-nesting birds, sug- gesting that the adult is more host-specific than the immature stages,

Camin (1963) has suggested that the reason the adult is more host-specific than the immatures is because it is a nidicole, dropping off the host and remaining in the nest when the host is not present, The small numbers of adults found on hosts other than leporids may perhaps be answered by this supposition,

Of the leporids collected in Utah, 81 were from the several species of Sylvilagus in the state, and 106 from the species of Lepus. The average numbers of~, leporispalustris in all stages collected from these two genera are given in 81

Table 7" Immature Haemaphysalis leporispalustris taken from hosts other then leporids in the western United States,

Host Larvae Nymphs

Citellus Sp" 1

Citellus variegatus 1

Dipodomys ordii 2

Eutamias minimus 11

Mustella satuvata 1

Melospiza lincolni 1

Neotoma cinerea 2 6

Oberholseria chlorura 11

Passerella iliaca 7 1

Pipilo fuscus 1

Pipilo erythrophthalmus 120 5

Pica pica 2 1

Peromyscus maniculatus 2

Thomomys bottae 16

Thomomys umbrinus 1

Thryomanes bewicki 31 82

Table 60 These data show that the greatest difference in average number existed in the larval stage and nymphal stage.

The adult averages only varied by 0.800 If the adult is a nidicole in the western United States then one would expect

to find more Sylvilagus parasitized 9 since it nests, than the non-burrowing Lepus. From these data, Ho leporispalustris does not seem to prefer one genus over the other,

Haemaphysalis chordeilis

Geographic Distribution

No specimens of H. chordeilis were collected by

Brigham Young University survey parties during this study.

Those data supplied by various agencies and interested per- sons show that part of the distribution of this tick is in

Montana 9 Washington and Wyoming (See Figure 21), From the literature (Bequaert, 1946; Cooley, 1946; and Carpenter,

1956) this species has been reported from California and

Oregon, Extensive collection data are lacking, and without this a proper picture of its distribution in the western states cannot be achieved. A more concentrated effort to collect known hosts of H. chordeilis in all the western 83

Fig. 21. Geographic distribution of Ji. chordeilis in the Western United Sta.tea. 84 states may appreciably extend its known distribution.

Seasonal Occurrence

Data are lacking to properly evaluate the activity on

a state basis. Figure 22 represents the western United States 9

and shows that adults have been collected in January 9 April,

Juney August 1 and September; nymphs in June and September; and larvae in September and October. The January collection is from Northern California. Reference to Table 2 will show the exact dates and localities of each collection.

Host-Parasite Association

Various species of grouse seem to be a prevalent host of H. chordeilis. But the data at hand are not numerous, and an accurate picture of host-parasite relationships is not clear. Nine records are represented in this study with H. chordeilis being collected from the grouse from four records.

This tick has also been taken from a gopher, Thomomys sp. 9 and from a cow, Bovis sp. (See Table 8).

Anatomical and Morphological Studies

The genus Haemaphysalis is distinguished from other 85

80 .Adults - Nymphs_ Larvae ::::::::::?

0 J F M 0 N D

Fig. 22. Seasonal occurrence of H. chordeilis by month in the Western United States. 86

Table 8. Hosts of Haemaphysalis chordeilis in the western United States.

Blue grouse--Dendragapus obscurus

Domestic cow--Bos indicus

Grouse

Pintail prairie chicken--Tympanuchus cupido

Sage grouse--Centrocercus urophasianus

Sharp-tailed grouse--Pediocioecetes phasianellus

Gopher--Thomomys sp.

White-crowned sparrow--Zonotrichia leucophrys 87 g~nera of hardbodied ticks in the western United States by

the post-anal groove 9 reflexed palpal aritcle II and the lack of eyes. The characteristics referred to in this study may be seen by turning to the appropriate figures of H. chordeilis and H. leporispalustris as directed.

Adult Characters. Haemaphysalis chordeilis male is easily distinguished from the male H. leporispalustris by the extended fourth coxal spur. Other anatomical structures which are also of value are the lack of ventral cornua

(H. chordeilis); and a rather long, pointed, retrograde spur on palpal article III which is found on the ventral surface and extends medially and posteriorly on H. chordeilis. There is only an indication of such a spur on H. leporispalustris.

When obvious, it is short and blunt. The second palpal article on H. leporispalustris is extended laterally at the base and is sharply pointed, whereas on H. chordeilis the lateral extension at the base is not as great, and the edge is bluntly rounded"

The female H. chordeilis lacks ventral cornua as in the male. On the dorsum the porose areas of the basis capituli are large and oval, as opposed to the smaller, elongate 88 porose areas of the female H, leporispalustris, The spur on coxa I in H, chordeilis is long, narrow and sharply pointed,

The female H, leporispalustris coxa I spur is shorter and bluntly rounded, Palpal article II and the ventral spur of palpal article III in the females of the two species are similar to those of the males,

Nymphal Characters, The most diagnostic features are the basis capituli and the ventral retrograde spurs on palpal article III, In the nymph of H, chordeilis the basis is drawn out to a blunt point laterally, whereas in H, leporispalustris the basis is quadrate and lacks such a lateral projection, Likewise, the ventral retrograde spur is pointed and well defined in H, chordeilis and only indicated in H, leporispalustris,

Larval Characters. The important diagnostic structures in this stage are similar to those used to distinguish the nymphal stages: the lateral points of the basis in H, chordeilis and the ventral spur? as opposed to the lack of these in H, leporispalustris, 89 The reader is referred to detailed Figures 23 to 29 for an explanation of the structures employed in the following key. Key to the Species of Haemaphysalis in the Western United States

Ma lEs

1. Coxa IV with long spur; ventral cornua absent; ventral retrograde spur on article III long and well-d~fined (See Figure 26) " . " " . . . Ho chordeilis

l'. Coxa IV without long spur; ventral corrua present; ventral spur on article Ill not well defined (See Figure 29) . . . . . l-L lepori.spalustris 90

Females

L Ventral cornua absent; well defined ventral spur on article III; coxa I spur long and pointed; porose areas large and oval (See Figure 25) . . . . H. chordeilis _j CJ l'. Ventral cornua present; ventral sp~r on article 111 absent or not as above; spur on coxa 1 as broad or broader than long; porose areas not as above (See Figure 28) . H. leporispalustris 91 Nymphs

L Basis capituli drawn to a point laterally; well de~ fined ventral retrograde spur on article III (See Figure 24) , " , Ho chordeilis

I' l

l', Basis capituli not drawn to a point laterally; spur on article III absent or not well defLned (See Figure 27) , Ho .l c:poris pa lus tris 92

Larvae

1. Basis capituli drawn to a point laterally; dcfinit.e ventral retrograde spur on article III (See Figure 23) H, chordeilis

l'. Basis capituli not drawn to a point: laterally; spur on article III absent or not well defined (See Figure 23) H . l c per is pa 1 us tr i s 93 ,-..,,..._:::1-r------Article III Retrograde Spur Basis Capituli (lateral point) Coxal Spur t ctS , Setae (I;J t /J " .._., @)

' ~ ' Anus

t ..._ , ,

¾------Article III -HFL-+-~.,,.c.----+------Article II .v--~~_,~-----Scapula

Fig. 23. Dorsal and ventral views, larval H. chordeilis (upper) and larval H. leporispalustris (lower). '\.:i:ill~'---i'-:'RIMr-'""il--'~---'-'-:'-~t-r----ArticleIII Retrograde Spur

~~s-;;;:---Lateral Point of Basis Capituli

Fig. 24. Dorsal and ventral views, nymphal H. chordeilis. 95

Article III Spur

Porose Area

Coxal Spur

female H. chordeilis. Fig. 25. Dorsal and ventral views 96

~~'+--,J~~~'------Article III Retrograde Spur

.· 0 .

- 0

.. Q' . 0 . .0 0

.• •· :"·

~,-.,-C--'---,--f~..,.c.....~=-----c-·~•~·~~...:._:,_~--;•_·~....,..~~-Fourth Coxal Spur •• 0

Fig. 26. Dorsal and ventral views male H. chordeilis. 97

Article III

Basis Capituli

t.·.·.. I .

. . o ..

,· ,.-

9 ·: . o· -~'?·::. :_..

Q. .~ ·. ·. 0 . .· o.' .. 0 ....·. 0 . 0 . : 9 . .· : ·.o. . 0. .. ·o . 00 .. ,' o· O· . o· ... . . 0 ·. o.. ·. 0 .o . o .. ·.· .... 0 . ·. o· ·b

Fig. 27. Dorsal and ventral views nymphal H. leporispalustris. 98

Article III

Ventral Cornu

'0' -~·

.o. . ·o. 0

o. 0 .0 0. 0 ... ·. o·.

, .· O· ·. '. ·o '.d .. .. 0 :o·

0 0. O·

Fig. 28. Dorsal and ventral views female H. leporispalustris. 99

Article III

·c ·• 0 0

" ' fJ ~-• " ,.G ·.~ •• 0 .. • 0 . 0 • 0 Q' ·•

I) Spur • 0 .....- ·' o.• . .. • Jo \J • s • " a C'; ~- <,I ,;, ~ i " ~ " ~ . " (, ., " .. I ~ 0 ,• • Q Ii ...... • "· • • ,. '& ' • 11, ~. ~- ii .. •. " ll (il

e ..

Fig. 29. Dorsal and ventral views male H. leporispalustris. 100

CONCLUSIONS

The two species of Haemaphysalis occurring in the western United States were studied with respect to geograph-

ical distribution, seasonal occurrence, host-parasite assoc-

iation, and external anatomy and morphology.

Haemaphysalis leporispalustris was found to be widely distributed throughout the west, especially in lowlands, valleys, deserts, and foothills. Further study is necessary to determine the altitudinal distribution of this tick, and possible reasons for limitations in altitudinal distribution.

Records for some states are not abundantj and though it appears that H. leporispalustris may be found in locations elsewhere similar to those where it has been collected, fur-

ther data are necessary to corroborate such suppositions.

Weather appears to influence the seasonal activity of

the "rabbit tick." In the southwest and along the Pacific

Coast to Oregon this tick has been observed during the winter months. On the other hand, in Montana and northern Utah where hosts of!:!_. leporispalustris have been collected throughout the year, activity ceases during the months of 101

December, January and parts of Februaryo It is probable

that two life cycles per year can be observed in the warmer

parts of its distribution, such as in southern Utah to south-

ern California 1 and along the Pacific Coast"

Possibly each developmental stage has a peak activity

in the western United States as was observed for Illinois and

Wisconsin by Mohr and Lord (1960) and Javidpour (unpublished

thesis) respectively. Further study is in order to definitely

conclude on this pointo Where larval activity was first ob-

served by the above workers in June in areas where the tick

hibernates, in the northern parts of the western states larval

activity was first observed in March.

In the western United States the preferred hosts are

leporids. Haemaphysalis leporispalustris does not appear to

show any preference for a genus of leporid. Incidental hosts

are rodents and some ground-inhabiting.birds that are only

rarely parasitized by the adult and more frequently by the

immature stages of this tick.

Drawings are made of a 11 developmenta 1 stag es of !::!•

leporispalustris and H. chordeilis for comparative purposeso

Keys are constructed to be useful in separating one species

from another. The external anatomical and morpholocigal 102 structures of taxonomic value are included in the keys ac- companied by labeled drawings,

During this study no specimens of H. chordeilis were collected by the Brigham Young University survey parties,

Data were obtained through correspondence with other workers,

a review of information at the Rocky Mountain Laboratory 1 and from loans of specimens. The majority of collection sites were from Montana, though a small number included Oregon and

Washington, One collection record represented northern

California. The data are not abundant, therefore geographic distribution analysis is limited, In the western United

States H. chordeilis seems to be confined to the northwest,

Haemaphysalis chordeilis has been observed during five months of the year in one developmental state or another in

California, Montana 1 Oregon, Washington 1 and Wyoming, Data are lacking to conclude anything significant for seasonal activity. One record showed that it was present during

.January, This would not be unusual in view of moderate climatic conditions existing in that area,

Though host records compiled for H. chordeilis are few, it is observed that birds are most frequently parasi- tized1 and this is in agreement with the literature. Its 103 common name is the "bird tick" because of its preference for these hosts, A small number of records exist for hosts other than birds, 104

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49~691-692.. 116

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(n. comb.) and of H. papuana kinneari Warburton (n.

comb.) Ixodoidea 9 Ixodidae) of southern Asia and

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and rabbit populations. Ill. Nat. Hist. Surv. Div.

Bull., 25(6):351-382. ABSTRACT

Two species of Haemaphysalis occurring in the western

United States were studied with respect to geographical distribution, seasonal occurrence, host-parasite association and external anatomy and morphology,

~- leporispalustris is widely distributed in the west.

Abundant records for some states are lacking and should be provided to extend the knowledge of this tick's distribution.

Weather apparently influences the seasonal activity of H. leporispalustris. In the southwest and along the

Pacific Coast to Oregon, this tick has been observed during the winter months. In Montana and northern Utah it has not

been observed during DecE:mber 9 January, and February, although its hosts have been collected. It is probable that two life cycles per year may be observed in the warmer parts of its distribution where it is active the year round.

In the study area th~ preferred hosts are leporids with no apparent preference for one genus. Incidental hosts are rodents and some ground-inhabiting birds which are seldom 2 parasitized by the adult tick but commonly by the immature stag es.

Drawings were made of all developmental stages of both H. leporispalustris and H. chordeilis for comparative purposes. Keys were constructed to distinguish one species from the other.

No specimens of H. chordeilis were collected by

Brigham Young University survey parties during this study.

Data were obtained through correspondence with other workers and agencies and from loans of materiaL The majority of

collection sites are in Montana 1 though a small number in-

clude California, Washington 9 and Wyoming. Geographical distribution analysis is limited because data are not abundant.

In the states where H. chordeilis has been recorded 1 it has been observed during five different months of the year. Data are lacking to conclude anything significant on seasonal activity. One record showed that it was present during January. This would not be unusual in view of moderate-climate conditions existing in that area.

Although host records compiled for H. chordeilis are

few 1 it was observed that birds are most frequently parasitized. 3

A small number of records exist for hosts other than birds.

Approved;

1qGJ Date